Front Cover
 Table of Contents
 List of Tables
 List of Figures
 Systematic paleontology
 Age and correlation
 Biographical sketch
 Back Cover

Title: Blancan Mammals from Haile XVA, Alachua County, Florida
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00097740/00001
 Material Information
Title: Blancan Mammals from Haile XVA, Alachua County, Florida
Physical Description: xii, 156 leaves. : illus. ; 28 cm.
Language: English
Creator: Robertson, Jesse Steadman, 1934-
Publication Date: 1970
Copyright Date: 1970
Subject: Mammals, Fossil   ( lcsh )
Paleontology -- Florida -- Alachua County   ( lcsh )
Zoology thesis Ph. D
Dissertations, Academic -- Zoology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
Thesis: Thesis -- University of Florida.
Bibliography: Bibliography: leaves 146-154.
Additional Physical Form: Also available on World Wide Web
General Note: Manuscript copy.
General Note: Vita.
 Record Information
Bibliographic ID: UF00097740
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000570662
oclc - 13717410
notis - ACZ7641


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Table of Contents
    Front Cover
        Page i
        Page i-a
        Page ii
        Page iii
    Table of Contents
        Page iv
        Page v
    List of Tables
        Page vi
        Page vii
    List of Figures
        Page viii
        Page ix
        Page x
        Page xi
        Page xii
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
    Systematic paleontology
        Page 10
        Page 11
        Page 12
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        Page 125
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    Age and correlation
        Page 127
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
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    Biographical sketch
        Page 155
        Page 156
        Page 157
    Back Cover
        Page 158
        Page 159
Full Text






To Shirley


Fossil. material was examined from the American L.uc-seumr

-of Natural Hi.story, the University of California Museum of

Paleontology, the Florida State Museum, and the University

of Kansas. I wish to thank the persons in charge of these

collections for the opportunity to study the ma-te -ials in

their care, and to acknowledge the excellent wor-X of

Mr. Russell Parks in preparing the plates.

I also wish to thank Dr. S. David Wveb'b, Dr. '. .,

Patton: and Dr. II. K. Brooks for tle.iir encouragement and

help during the course of this study.

. i. L


Pagt q








Order Isectivora
ryptoisis P-rva .
Scalocus aquaticus.

Order Edentata
Glosso Lherium cha padmalensi s ...
KracL0 ievichi a paranensis . .
Review of the chlamythl eri ie.
Evolution of the chlamythea.es.

Das.uls bellus. .

Order L:,go'-.orpha
Sylvj.laqus sp. .

Order Rodentia
Peauria sp. . .
Castor canrad- nss .

.Si__g;io_,don inedius .

Or(ce C.ari :or
Canid. 2 . . .

Smini lcdon cgrac :i. s




. . . . . . . . . 1

. . . .. .
. . . ... ]0


. . . . 79
. . . . 87
. . . 88

. . 90
. . 90
. . . . 95



Order Proboscidca

Gomphotheriideo . . .

Order Pcrissodacty.1o
NannippU17 ph.l:on . . .
P]Lesyippus simplicidens. .
Equus (Asinus) sp. ..
Tapirus sp. . . .

Order ArtioCact.yla
l.y1ohyus flor.idanus . .
HIImiauchenia cf macrocephala
Odocoileus virginianus. .





. . . 97

. .. 99
. . . . 99
. . . . 109
. . . . 111

. . . . 111
. . . . 117
. . . . 122


1 135

S. 140

. . 142




Tab l Pageo

1 Mammalian Faunal List and Minimum Number
of Individuals . . . . . . . 11

2 Measurements (in mm) of the Lower DenrLition
and Mandible of UF 17466 and Other Fo:sil
and Recent Cryntotis parva . . . ... 12

3 Measurements (in mmr) of the Upper and Lower
Dentitions of Three Species of Glosso-
therium. . . . . . . . . . 24

4 Limb Measurements (in mm) of Glossotherium
chapadmalensis and Glossotherium hirlani 26

5 Measurements (in mm) of the Bones of the
Manus and Pes of Glossother:ium chapadmaicnss
From Haile XVA and Glossotherium harlani 28

6 Cranial and Dental Measu:ements (in mm) of
Kraglievichia and Chlamiyterim. .. . 39

7 Measurements (in mm) of Limb Eoements of
Kragli.cvichia paranensis, UF 30902,
Haile XVA . . .. . . . . . 44

8 Measurements (in mm) of Limb Elements of
-ra q1ievichi a paranonsis From Various
Florida Localities . . . . . . 53

9 Dental Measurements (in mm) of Dasypias
bellus . . . . . . . . . 73

10 2^u.-:rem-ents. (in mm) of the Femora of
Fossil -and iclcent Casto;- canadensis. ..... 89

11 Dental.. MeIaslrenmcnts (in mm) of Siqmodon
mec:ius nc Sifmodon minor .. . .... 9i


Table Page

12 Measurements (in mm) of the IlHuecus of
Fossil and Recent Pteronura and Recent
Lutra . . . . . . . . . 96

13 Measurements (in mm) of the Astragali of
Smilodon gracilis From Ilaile XVA and
Port Kennedy . . . . . . . 98

14 Measurements (in mm) of Nannippus phlegon
Cranial and Postcranial Material From
Haile XVA. . . . . . . . . 102

15 Measurements (in mm) of the Upper Dentition
of Plesippus simplicidens From Haile XVA . 106

16 Measurements (in mm) of Plesi ppus simp.li-
cidens Postcranial Material From IHaile XVA 107

17 Measurements (in mmu) of Equus (Asinus) sp.
Cranial and Postcranial Material From
Haile XVA . . . . . . ... . 0

18 Measurements (in mm) of the Dentition of
Mylohyus floridanus. . . . . . . 136

19 Measurements (in mm) of the Upper Dentition
of Hermiauchenia cf macrocephala From Iaile
XVA . . . . . . . . . . . 21

20 Measurements (in mm) of Postcranial Elements
of Odocoileus virginianus From Several
Florida Pleistocene Localities . . .. 1.25

23 Faunal Comparisons of ilaile XVA and OLher
Blancan Localities . . . . . . 129

v :i i



















Geologic Section at Haile XVA. .. ..

Glossotherium chapadmalensis Upper
Dentition . . . . . . .

Glossotherium chapadmalensis Right
Mandible, Lateral View . . . .

Glossotherium chapadmalensis Right
Mandible, Occlusal View. . . . .

Kraglievichia paranensis Cranial Mater

Kraglievichia paranensis Front Limb
Elements . . . . . . . .

Kraalievichia paranensis Hind Limb
Elements . . . . . . . .

Kraglievichia and Chlamytherium Calcan

Kraglievichia and Chlamytherium Femora

Kraglievichia and Chlarmytherium Dermal
Plates . . . . . . . .

Dasypus bellus Mandible. . . . .

Dasypus bellus Astragali . . . .

Petauria sp. Right Mandible. . . .

Petauria sp. Right M3. . . . .

Ptcronura Humerus. . . . . .

Nannijp_!s p hSeon Upper Molar .. .



. 16

S. 19

. 21

ial. 34

. . 42

S. 48

ea 51


. 64

. 71

. 75

. 82

. 84

. 94

. 101


Fig cre Page

17 Pl.iS~us simpl.icidns Upper Dentition . 105

18 MyJ]ohyus floridanus Mand:ibular Ramus . .13

19 Mylohyuus floridanus Mand.ibular Syrmprysjs . 115

20 Ilemiauchenia cf racrocephala Upper
Dentition . .. . . . . . . 119


Abbreviations of institutions and collections used

throughout this work are as follows:

AMNH: American Museum of Natural History

U.C.M.P.: University of California Museum of Paleon-

UF: University of Florida Collections

UK: University of Kansas

H.C.T.: University of Houston Collection

Abstract of DisserLatLion ]?resented to tl-h Graduate Council
of the University of .l.or.i da in ParLial ]Fulfillment of the
Requirements for the Degree of Doctor of Phi]osophy



Jesse Steadman Robertson, Jr.

December, 1970

Chairman: Dr. S David Webb
Major Department: Zoology

The Iaile XVA fauna is the first Gulf Coas-L.tl Plain Blancan

fauna to be described. Among the mammals, which are repre-

sented by 8 orders, 17 families, and 18 genera, are two

edentates which are new to the North American fauna. Glos-

sotherium chapadmalensis (Kraglievich), a small South

American ground sloth species, is reported from North

America for the first time. This species was previously

know from thle ChapadmaJalan (early Pleistocene) fauna of

Argentina. It is proposed that this form may be ancestral

to both C. harlani (Owen) and G. robustus (Owen), Kray~li:-

v.ichi paranonsis (Kraglievjch) a giant, extinct almainl.o

previously known~ only from Miocene and Pliocene deposits of

South America-., is reported for the first time in Nort.h

America. T)c: laile XVA specimen is described as a noe\w s!b)-

species. D I pu be.1 us (Si-mjson), a smaller extinnct arinU.-

dillo, is rc.po.rtLod for the first time in Ilhc Blancan, marxk-

in'j it-s earliest occurrcnc', anywhere.


A brief review of the Subfamily Chlamytheriinae is pre-

sented, and it is suggested that Plaina (Castellanos) is a

synonym of Kraglievichia (Castellanos) and that Hoffstetteria

(Castellanos) is a synonym of Chlamytherium (Lund). Chla-

mytheres are shown to have been present in North America

continuously since the re-establishment of the late Cenozoic

land bridge between North and South America, and it is

proposed that the Pleistocene evolution of these forms

occurred simultaneously on both continents.

The Old World Flying squirrel Petauria is reported for

the first time in the New World. It was previously known

only from early Pleistocene deposits of Bavaria.

Several of the Haile XVA taxa are closely allied with

South American Plio--Pli stocene forms, which further

strengthens the previously established correlation between

the Chapadmalalan stage of South America and the Blancan

stage of North America.

The abundance of aquatic non-mammalian vertebraLes

indicates that the environment of deposition was probably

an open stream. The mammalian members of the terrestrial

community include the tropical or subtropical indicators

Kraqclievichia and Dasypus. Forest indicators include

fj'y1chiyus, CasLor, Petaurjo, and Glossotherium. The zoo-

geography of the more exoLic forms is discussed.


Florida has long been famous for its Pleistocene fossil

vertebrate deposits. Most of these sites have yielded

faunas represenLing the later stages of the Pleistocene.

Classical examples of these are Vero, Indian River County

(Sellards, 1917; Hay, 1917; 1928; Wiegel, 1962), Melbourne,

Brevard County (see Ray, 1957, for numerous references),

and Seminole Field, Pinellas County (Simpson, 1930) which

are all of late Rancholabrean age. Among the slightly

earlier Rancholabrean faunas are Haile VIIIA, Alachua

County, and Bradenton, Manatee County (Robertson, in press).

An Irvingtonian site, Coleman IIA, Sumter County, has

recently been described by Martin (in press). To date, this

is the earliest Florida Pleistocene fauna to be fully de-

scribed. Another excellent Irvingtonian fauna, Inglis IA,

Citrus County, is presently being described by Mr. Jean Klein,

a graduate student in geology at the University of Florida.

The SontLc Fc I locality, Gilchrist County, has long

been conjsidcered to be ,.,f Dl:n-.can age, but the fauna is ;s

yet undcr;rciLn:d except for thie description of T'ianit., a

giant, flightless bird (Brodkorb, 1963). A faunal list for

this locality is provided by Webb (in pressa. Other Florida

deposits which are considered to be of Blancan age, but

which have not been studied, are Santa Fe IV and Santa Fe

VIII, Gilchrist County.

Correlation of Florida Pleistocene deposits is diffi-

cult at best, because of their mode of deposition. The two

most common types of Florida Pleistocene fossil vertebrate

deposits are filled sinkholes in limestone and river bottom

deposits. The various fissure fillings have trapped sedi-

ments representing many stages of the Pleistocene, and

adjacent sinkholes may contain faunas which vary greatly in

age. River bottom deposits are usually heterochronous

accumulations of bones which have eroded out of fossilifer-

ous strata and settled at low energy areas of the stream.

This lack of orderly stratigraphic sequences for most of the

Florida fossil vertebrate deposits necessitates the use of

faunal correlation.

Usually, a reasowrabl y accurate age assignment can be

made based upon the presence or absence of certain genera

andl species. The presence of Bison, for example, indicates

a Rancholabreon age (Savage, 1951). Further refinement of

the age could be made by deicrmining the species of Bison

present. The. presence o f Bi. latilrons would represent early

Ranncholabrean, ). ant iquus late Rancliol abrecan, and B. bio3,

sub-ReconL or Recent (Robertson, in press)

In addition to fauna] correlation, the effects of sce

level changes due to Pleistocene glaciation may be employed

as an aid to dating Florida Pleistocene deposits. Glacial

ecstatic fluctuations of sea l.eve.] have caused "terraces"

or "abandoned shore lines" which are dominant geomor]phir-

features of the Florida landscape (Cooke, 1945; Mac-Ceil,

1949; WVhite, 1958; Alt and Broohs, 1965). The location of

a deposit relative to these structures can be quite helpful

in age determination. If, for example, a coastal terrestrial

deposit is presently lying at or below sea level, then it

must represent a time when sea level was as low or lower

than it is now. This would be an indication that -he rde-

posit was laid down during glacial times, as these weve the

times of low sea level. Once this is determined, fauna].

elements will indicate which glacial stage is represented.

Inland deposits of higher elevations present a problem

in thai. they could have been deposiLed dnrjijn ejiller a

glacial or intcrglcacial stage. If, however, .it can be sbho'..n

thidt a partic-ular inland deposit was coastal at its time

of deposit-.io.n, then it must represent an interglacial si~.ge,

when sea 3 ,-,. 1. was high. As a rule it is difficult to as.c

certain whether or r.o a tc rreoL.rinal fauna rcpy u:; 'nts i,.:and

or near-shore conditions. Auffenberg (1953), however, has

been able to do this for certain deposits. Working with

fossil Terrapene carolina, he has shown that particular

intergradient populations of this species reflect near-

shore conditions. If members of these intergradient popu-

lations are present in an inland deposit of higher ele-

vation, the deposit probably represents an interglacial

stage. Evidence for near-shore conditions is not always

this conclusive, however, and sometimes one is forced to

rely upon more tenuous means.

Of particular interest among the Florida fossil verte-

brate deposits are those near the settlement of Haile,

Alachua County, conveniently located only 15 miles from the

University of Florida campus. In a radius of only a mile .re

clustered numerous vertebrate localities, traditionally

designated by roman numerals in the University of Florida

vertebrate paleontology collections. Most of the Haile

localities have produced fossils of Rancholabrean age.

Notable exceptions to this are localities V and VI .which are

considered to be Pliocene (Auffenberg, 1955) and locality

XVA" which is regarded as Blancan. The fossil vertebrate

assemblage recovered froii the Haile XVA locality constitutes

the subjecL for this dissertation.

The Ha:l.e 'VFA locality is situated on the property of

Parker BroLhic.rs' Limestone Products, Inc., near Haile, R17E,

T95, S25, Alachua County, Florida. It was discovered in

1964 by Mr. Phillip Kinscy of Jacksonville Beach, Florida,

who has long been a friend of, and a contributor to, the

Florida SLate Museum. Mr. Kinsey did extensive collecting

at Haile XVA, and, upon recognizing it as an unusual local-

ity, brought it to the attention of the museum staff.

Further excavations were carried out in the same year by

Dr. S Ddavd Webb, Mr. Robert Allen, and the present author,

with the support of NSF Grant GB 3862. The resulting

collections, in addition to the large quantity of bones so

generously donated by Mr. Kinsey, are now housed in the

University of Florida Collections.

In addition to the liammialian fauna described in this

report, the locality has also contributed extensive fish and

reptile faunas. A fair amount of amphibian and avian re-

mains is also present. Among the reptiles are the type an.d

abundant mzuter.ial of Chrysemys platymarginata (Weaver and

Robertson, 1967).


The Haile XVA deposit, lying at an elevation of about

90 feet above sea level, is a filled sinkhole in the Ocala

(Eocene) Limestone. The deposit extends approximately 30

feet along its east-west axis; its north-south dimension is

narrower and irregular in width.

Underlying the fossil-bearing matrix is a layer of dark

brown, unfossiliferous hardpan which begins at a depth of

approximately 20 feet and extends to a depth of at least

six feet from its upper margin (Figure 1).

The fossil-bearing matrix extends from about five to

20 feet below present ground level, and consists of two

units: an upper layer of heavy, greenish clay; and an

underlying layer of coarse, poorly sorted, gravelly sand-

stone, containing lenses of clay. Where these two units

merge, an alternation of clay and sand lenses occurs. The

lowermost massive sand is the most productive part of the

fossiliferous sequence. It contains much calcareous cement

and calcareous cemented concretions. Many of the larger

bones v'cere en"trusted with cemented sand and gravel. The


Figure 1

Geologic Section at Haile XVA

A. Sandstone alternating with gray clay

B. Heavy greenish clay

C. Coarse, poorly sorted, gravelly sand--

D. Dark browvnr hardpan

E. Ocala Limestone (Eocene)

-- - o


l r : . . .

. . . .

sandstone and clay were shown to be esscnLially contemporca-

neous by the occurrence of fossils which extended from the

sandstone up .into the clay.

Overlying the fossiliferous sand and clay sequence is

a 3ayer of thin, brown sandstone alternating with gray clay.

This layer is unfossiliferous.

Since the original collections were made, mining ac-

tivities have scattered the remaining matrix over a wide

area. On a recent trip, for example, a tooth of Glosso-

therium belonging to the associated skeleton taken from

the original deposit in 1964 was found on one of the ad-

jacent piles of matrix. The site is presently covered ]by

several feet of sand washed in from the overhanging lime--

stone ledge. There is little, if any, of the matrix -e-

maining in the original site.


There are 20 taxa representing a minimum number of 34

individuals in the Haile XVA fauna (Table 1) These are

considered on the following pages.

Order Insectivora

Family Soricidae

Cryptotis parva Say

Material: UF 17466, right mandible with M -M,

This specimen is morphologically indistinguishable fircm

specimens of C parva from the Irvingtonian Coleman IIA and

from recent specimens. The Haile XVA specimen is slightly

larger than the previously mentioned specimens (Table 2),

but not significantly so. The only other Blancan record of

Cryptotis is the Rexroad fauna (Hibbard, 1937, 1941), Meade

County, Texas.

Family Talpidae

Scalopus acuaticus Linnaeus 1758
Material: UF 17466, 2 left M ; UF 17465. right humerus.

A comparison of the three iHaile XVA specimens with

corresponding modern S. aquaticus from the southeas-tern

Table 1.

Mammalian Faunal List and Minimum Numbner of Individuals

CryptoLis parva 1

Scalopus aquaticus 2

Glossotherium chapadmalensis 1

Graq] ievichia paranensis 3

Dasypus be]lus 1

Sylvilaqus sp. 3

Petauria sp. 1

Castor canadensis 1

Siqmodon medius 2

Canidac 2

Pteronura sp. 1

Smilodon gracilis 1

Gomphlotheriidae 1

Nannippus phloeon 2

Plos:i ppos simplicidens 2

Equus (Aslnus) sp. 2

Tapirus sp. 1

y14~ohyus floridanus 1

eminiauchon-c J. cf macroccphala 1

Odocoileus vjir gnianus 5

Table 2

Measurements (in rmm) of the Lower Dentition and Mandible of UF 17466
and Recent Crvptotis parva

X = mean; 0. R. = observed range; number of specimens in

and Other .Fossil


US 17466

Condyloid to mental foramen

Ccndyloid to Mi

Condyloid to M2


I1 -M

z J






C. parva* (20)

X O. R.

6.1 5.9-6.7

6.9 6.7-7.4

5.7 5.5-6.0

2.6 2.4-2.7

3.4 3.3-3.6

2.2 2.1-2.4

C. parva* (fossil)

X 0. R.

6.6 6.1-7.0 (8)

7.1 6.8-7.6 (8)

5.8 5.5-6.1 (8)

2.7 2.5-2.9 (8)

3.6 3.5-3.6 (2)

2.3 2.2-2.3 (2)

"measurements after Martin (in press)

United States revecll no significant rcifierenc-ies in size

or morphology.

Scalopus is also present in the Blancan Rexroad fauna

of southwest Kansas (Hibbard, 1941). The only other mole

from the Rexroad fauna is Hesperoscalops, which is based

upon a partial lower dentition. A direct comparison could

not be made with this genus as only upper teeth are known

from ilaile XVA. As Hibbard (1941) points out, however,

mole humeri of various genera can be distinguished from one

another, so it would be illogical to assume that a Scalopus-

like humerus would belong to another genus. Because of this,

and since Scalopus is known from other Blancan and Irving-

tonian deposits, it seems reasonable to ascribe this ma-

terial to the living species.

Order Edentata

Family Mylodontidae

Glossotherium chapadmalensis

Kraglievich 1925

Mater-ial: UP 10922, partial skeleton

There i.s no doubt that the Haile XVA specimen repre-

sents one individual as there is no duplication of elements,

and the left and right elements agree very closely in all

mcas urcnent,. Parts of this specimen were collected by

several pc(',..! c~,vr a number of years, b)i L the bulk of the


specimen was taken from the sand layer of the fossiliferous

sequence and was semi-articulated.

In the following discussion, terminology regarding sloth

dentitions will be based upon that used by Hirschfeld and

Webb (1968): the first tooth (upper and lower) will be

referred to as the caniniform; all the rest of the teeth

will be referred to as molariform. No homologies are im-


Upper dentition: The upper dentition of the Haile XVA

specimen (Figure 2) is complete except for the left upper

caniniform. The caniniform is triangular in cross-section

and well developed. As in the Argentine specimen of G.

chapadnmalensis, this stoutness of the caniniform is re-

flected by the expansion of the anterior portion of the

maxilla. The anterior portion of the maxilla is very much

reduced in G. harlani, which tends to have a reduced or

lost caniniform.

The first upper molariform tooth is large and 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. Due to the very

well-developedlower caniniform which in part occludes with

this tooth, the anterior oblique wear surface is much more

accentuated '-ha. in G. harlani. In several specimens of

Figure 2

Glossotherium chapadmalensis Upper Dentition

X 1.13 (linear)

6' *r L

1 ;1


*: . -.
!. r


* r

V ;
.r- ~
'B.',. :
r _L

R '
..J ";-.-. r
: *


--w. - --, -


C. robusLus observed, this tooth was worn off smoothly ju-L

above the alveolus. A very short diastcma separates the

caniniform from the first molariform tooth in the H!aile '1\.'A


The second upper molariform tooth of UF 10922 is tri-

angular in cross-section, with the base of the triangle

placed lingually. The anteroposterior distance is less L.an

the transverse diameter in this species, while the opposite

is true in G. harlani. Of the two specimens of G. robustus.

for which measurements are provided, one agrees with G.

chapadmalensis while the other is nearly equal in both

diameters. The third upper molariform tooth of UF 10922

is similar in shape to the second and agrees morphological,..ly

with the corresponding teeth in both G. harlani and G.

robustus. The fourL.h molariform tooth is relatively small

compared to the Argentine specimen of G. chanadmalensis al-

though the shape is the same.

The palate is very much co,,itricted in the area of the

last upper teeth and very much expanded at the anterior elnd

of the maxilla. Both of these characters are more extreme

in UP 10922 than in the Chapadmalalan specimen, or in speci-

mens of G. harlani and G. robustus.

Lower doe-nLition: The lower caniniform of UF 10922

(Figures 2, 4) is.. vcry wvell dcveIlopcd and di ectod labially.

Figure 3

Glossotherium chapadmalensis Right Mandible, Lateral View

X 0.82 (linear)



. A YC I

I k

~ '" v
op A A ,.C, 5

Figure 4

Glossotherium chapadmalensis Right Mandible, Occlusal View

X 0.82 (linear)




-'..! ..
d "'

ii .


This tooth is relatively slightly larger than the correspond-

ing tooth in the South American specimen of G. chlaji~na eC-

sis. The tooth is pointed when viewed laterally due to the

dual occlusion with the upper caniniform and the first

molariform tooth.

The first lower molariform tooth is also relatively

robust. It is somewhat triangular in cross-section and pro-

trudes fairly high above the alveolus. This tooth is also

turned slightly to the outside and closely resembles the

lower caniniform.

The second molariform tooth is rectangular and is turned

lingually, so that its long axis does not parallel the tooth

row. Stock (1925), in discussing this tooth, used the long

axis as the anteroposterior length while Kraglievich (..925)

used the shorter axis of the tooth for this measurement.

For comparison, Kraglievich's method of measuring is used

here for the Haile XVA specimen and Stock' s method for

measuring G. robustus. It was necessary to reverse the

measurements of G. robustus provided by Kraglievich (1928)

so they would be consistent with the other material con-

sidered. Thus, the seemingly great difference in the meas-

urements of the second molariform teeth is actually an ex-

pression of the fact that the orientation of the teeth is

different in ;the species and does not indicate gross

morphological 0jtifoerences i- Lhe teeth.


The third lower molariform tooLl is of a slightly dif-

ferent shape in G. chapadinalensis. The two main colulnns are

separated by a thin bridge, while in G. harlani and G.

robustus this bridge is thicker.

Mandible: Unfortunately, the mandibular symphysis is

not preserved in the Haile XVA specimen. The portion of the

mandible present agrees in morphology with the other two

species. Measurements of the upper and lower dcntitions are

presented in Table 3.

Cranial material: The remainder of the Haile XVA

cranial material consists of part of the occipital bone

and portions of the left and right periotic bones. The

occipital fragment contains the left condyle and a portion

of the otic region. The hypoglossal canal, jugular foraion,

and auditory meatus show no observable differences from

those of G. harlani and G. robustus. A portion of the left

squamosal is attached to the occipital. Part of the right

squamosal is present including the zygomatic process.

Vertebrae: The atlas of UF 10922 is well preserved.

According to Stock (1925) this element differs in G. harlani

and G. robuslus. The posterior portion of the laLeral

process does not extend as far forward in G. harlani. The

location of the dorsal foramina also varies: In G. harlani;

they are q it;.l far apart while in G. robustus they are

Table 3

Measurements (in mm) of the Upper and Lower Dentitions of Three Species of

G. chaDad- G. chapad- G. harlani G. robustus
malensis malensis
.1 .2
Haile XVA Argentina California Argentina'
Upper Dentiticn
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 2.40 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

Lower Dentition
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 17.8 18.0 23.8 20.2
1 measurements after Kraglievich (1925)
2 measurements after Stock (1925)
3 measurements after Owen (1842)


close together. UF 10922 definitely agrees with G. harl]ni

in the expression of these two characters.

The remainder of the vertebrae preserved are, for the

most part, in poor condition, and as a result their posi-

tions in the vertebral column could not be determined


Front limbs: Neither scapula of UF 10922 is present.

The right humerus is present. Neither radius is preserved,

although both ulnae are present. The left ulna is so poorly

preserved that it could not be removed from the plaster

jacket in which it was collected.

Several bones of the manus are present in the Haile XV-,

material. The cuneiform appears to be somewhat different

in G. chapadmalensis than in G. harlani. This element in

UF 10922,when viewed from the palmer aspect, is square in

appearance as opposed to being rectangular in the figure of

G. harlani provided by Stock (1925, Figure 72d). Thcse

differences are reflected by the measurements of this el--

ment in Table 4. In UF 10922, the proximal-distal distance

is greater than the distance across the ulnar articular

surface while the opposite is true for 39 specimens measured

by Stock. The cuneiform appears to be a deeper element in

G. chlapclmal onsis than in G. har l.ini The remainder of the

elemenl.ts of 1-h.-! manu. prc;sorved :in the llaiJ e XVA specimens

Table 4

Limb Measurements (in mm) of Glossotherium chapadmalensis and
Glossotherium harlani

G. chapadma!ensis G. harlani*
UF 10922,Haile XVA 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
Scondyles 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 Stoch (1925). The measurements represent the mean of
large samprn PTh nuimer of specimens varied from element
to element u nd in scmc czsc-s was not given.n

are the left scaphoid, right pis:iform, left melacarpal 2ll,

and left phalanx II, digit III.

Hind Limbs: The pelvis is missing in UF 10022. The

left femur is complete while the right is badly fragmented.

Both tibiae, and the left fibula are well preserved.

Several clements of the pes are present. These are

the right calcancum, right astragalus, left and right meta-

tarsal III, left and right phalanx II, digit III, and an

ungual phalanx, digit III.

A comparison of the limb elements of the Haile XVA

specimen with those of G. harlani from various Rancholabrean

deposits of Florida and with the figures of G. harlani pro-

vided by Stock (1925) and figures of G. robustus (Owen.

1842) shows, for the most part, no great morphological

differences. All three of these species appear to be very

similar postcranially. Comparative measurements of UF

10922 and Stock's (1925) Rancholabrean material show the

Haile XVA material to be significantly smaller (Tables 4, 5).

Glossotherium robustus is larger, about the size of G.


Discussion: There has been much confusion concerning

the taxonomy of the genera 1M.yodon and Glossotherium. Krag-

1ievich (392C) has presented a thorough review of the

literature. 'if concludes that MYylodon darwiijii is the geno-

Table 5

Measurements (in mm) of the Bones of the Manus and Pes of
Glossotherium chapadmalensis From IIaile XVA and Glossotherium

Greatest distance across
articular surface
Greatest distance across
dorsal surface from
inner side to outer
Greatest proximal-distal
Distance across articular
surface for ulna
Greatest length
Greatest depth
Greatest width
Metacarpal III
Greatest length
Width, proximal end
Width, distal end
Depth, proximal end
Phalanx II, digit III
Depth, inner condyle
Width, proximal end
Greatest width, anterior
Greatest depth
Anteroposterior diameter
Metatarsal IV
Width, proximal end
Depth, proximal end
Wi.dt'h, distal end
Depth, distal end

G. chapadmalensis
left right

46.2 ----



31.2 ----








G. harlani*

75.3 (39)**

68.5 (42)











140.2 (41)



Table 5 (continued)

G. chapadmal ensis
left right

Phalanx II, digit III
Depth, inner condyle
Width, proximal end
Ungual phalanx, digit III
Proximal-distal distance of
ungual base
Width, proximal end





G. har ani*

35.4 (32)



*measurements after Stock (1925)
numberbr in parentheses refers to the sample size from which
the mean (the numbers in the right-hand column) was

type of Mylodon, and that what Owen (1840) described as

Mylodon robustus more correctly belongs to a distinct genus,

GlossoLherium. However, Kraglievich retains Paramylodon

(Brown, 1903) as a valid genus even though Stock (1925)

showed that the type of Paramylodon merely represented a vari-

ation of what was then called Mylodon harlani. Simpson

(1945) argues that if the North and South American forms

(late Pleistocene) are not generically distinct, then they

all belong to the genus Glossotherium. This view is

followed here. Hoffstetter (1952) retains Paramylodon as a

subgenus of Glossotherium, recognized by its tendency to

lose the upper caniniform tooth, and because of their narro'-

muzzles and long skulls.

The partial, associated skeleton from Haile XVA is

clearly a member of the Subfamily Mylodontinae as indicated

by the single astragalar facet for articulation with the '

calcaneum. Following the above system of classification,

it is a member of the genus Glossotherium. This specimen

so closely resembles Eumylodon (Glossotherium) chapadmalen-

sis that: it is here ascribed to that species. Glossotherium

chapadmalensis is an early Pleistocene ground sloth from the

Chapadmalal fauna of Argentina and is here reported from

North America for the first time. It is a small species

which appcar;; to be very closely related to both G. harlani


and G. robusLtis, differing from these two forms in the nature

of the anterior dentition and in overall size. It agrees

with these two species in the morphology of most of the

postcranial elements with the possible exception of some of

the bones of the manus and pes.

As this species is known from the early Pleistocene of

Argentina and Florida it is thus temporally, geographically,

and morphologically, a plausible ancestor of both G. harlani

and G. robustus.

Family Dasypodidae

Kraglievichia paranensis new subspecies

Holotype: UF 10902, partial skeleton collected by

P. E. Kinsey, S. D. Webb, R. R. Allen, and J. S. Roberts;on

in 1964.

Type Locality and Horizon: Haile XVA, R17E, T95, S25,

Alachua County, Florida, Blancan.

Subh '::cific Diagnosis: A member of the species KrEa-

lievichia paranensis on the basis of all major morphological

characteristics. Differs from the only other subspecies,

represented by the South American members of the species,

in that the fourth tooth is reniform rather than peg-like

and is not turned lingually, but is nearly parallel with the

tooth row.

Referred Iatctcr.al: U) 10902, partial :'h,'1l and jaws,

and associated skeleton, Haile XVA holotypee); UF 17474,

right ulna, also from Haile XVA; UF 10432, right humerus,

Sante Fe I; UF 9354, right humerus, Santa Fe I; UF 10449,

left ulna, 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 XII!; UF

17568, and UF 17569, right naviculars, Haile XVA.

Description: The discovery of the Haile XVA skeleton,

as well as isolated postcranial material from several other

sites in Florida, affords an excellent opportunity to expand

the description of this species. Heretofore, postcranial

material of this species has been scarce.

The following description is based for the most part on

the Haile XVA specimen. In some cases material from the

other localities is used if corresponding elements are

lacking in the Haile XVA material.

Comparisons are made with North American specimens of

Chlamytherium, as well as with descriptions and figures

(particularly Winge, 1915) of South American chlamytheres.

Cranium: The nasals (Figure 5b) of UF 10902 compare

favorably in size and morphology with the South American

specimc-. of K. paranensis described by Castellanos (1927).

They _fcr, hov.ever, in the presence of tiny prou-ubcrances

Figure 5

KSraolivichia paranensis Cranial Material

A. Right premaxillary

B. Nasal

C. Palletal portion of maxillary

D. Zygoma, ventral view

E. Zygoma, material view

F. Right mandible, occlusal view

G. Right mandible, lateral view

H. First lower tooth

I. Fourth lower tooth

Abbrevi nations: iof, infraorbit-~I for.amen;
ju, jugal; nmx, maxil. ary; ;.x-px, maxillary-
premaxilla-y suture; pn-:: premaxilx :ary
n, ]nanals; numbers refer to tooth n j:rq.nc.c.

mx-pmx /
I ,



4 4.




/t ~ l

,ii ?m

L ~7~ r~iiD

- -7




-j 4'

25 mm

on the anterior endsc of the nasals near the medial surface.

These structures are not present on any of the other Chla-

mytherium or Kraciievi.chia specimens or figures studied.

Only the anterior halves of the nasals are present in UP

10902, as the skull had been eroded away, and only parts of

it were recovered.

The prinaxillary (Figure 5a) contains only one alveolus.

The premaxillary-maxillary suture forms the posterior, border

of the first alveolus, as is the case in Chlamytherium. In

Holmesina (=Chlamytherium), one of the distinguishing

features was supposedly the presence of one tooth in the

premaxillary. Two teeth were supposedly present in the

premaxillary of Chlamytherium. The neotype of "Holmesina"

(AYNII 26856, Simpson, 1930) as well as UF 889 (C. septn-

trionalis from IIornsby Springs, Bader, 1957) illustrates

this condition. James (1957) states that in his specimen of

C. septenrtrionalis (H.C.T. 4) there appears to be a suture

both before and after thu second tooth. A Chlamytherium

skull collected in South America by Dr. Gordon Edmund, Royal

Ont;ario Museum, has only one tooth in the premaxilary

(Edmund, personal communication). Thus, even if this charac-

ter should be found in some skull not now known, it would

not serve t sicparate the Noith and South American forms.

The major features of the maxillary compare favorably

with those in Chlamyth erium. The infraorbital foramen

(Figure 5e) is located directly above the sixth tooth in

both genera. The anterior palatal foramina (Figure 5c)

are located between the posterior edges of the fourth teeth

in the IHaile XVA specimen, while their position is somewhat

variable in Chlamytherium. The maxillary process of the

zygomatic arch is directly above the seventh tooth in both


Only a portion of the zygomatic arch is present in Uv

10902 (Figure 5d, e). The zygomatic process of the maxil-

lary turns posteriorly and downward as it leaves the sk11.l.

The anterior portion of the jugal, which borders the zygo--

matic process laterally, then turns upward and expands

posteriorly to accept the aquamosal process. A well-

developed suture present at the posterior end of the jugal

indicates that in Kraglievichia, as in Chlamytherium, the

zygomatic arch is complete. The base of the zygomatic

process of the maxillary is expanded by sinuses, as in


Upper dentition- There are nine upper and nine lower

teeth in each jaw of Kraglievichi-a. One upper tooth is

located in the premaxi1la.ry. Because the incisors of most

matinals occur in the premaxillary, it is tempting to refer


to the first tooth in KraJg.1. evich.i a as an incisor. However,

the terminology "tooth one" through "tooth nine" is con-

tinued in this report.

A conspicuous difference between the HaiLe XVA specimen

and previously described specimens of Kraglievichia and

Chlamytherium is seen in the nature of the anterior upper

dentition. In the 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

Chlamytherium, the first three teeth show this condition, buL

the fourth is usually bilobate and parallel with the rest

of the tooth row. This character is considered to be im-

portant in the evolution of the chlamytheres (Castellanos,

1937; Simpson, 1930). It is obvious, however, that this

trait is variable. In James' (1957) specimen the first four

teeth are oval, but the fourth tooth is parallel with the

rest of the tooth row, rather than being canted lingually.

The peg-like nature of the fourth tooth may represent a

primitive condition, assuming .Kraglievichia is the ancestor

of Chlamytherium. Vassallia, another even more primitive

chlamythcrc, with its first five teet.h being peg-like and

canted lingually, also shows tli, cveolutionary trend.

Costel.lanos (.1937) sees the evolution of chlamythere teeth

progress, .,g from th le back of IICe tooth row.' to Lhe front,


with the teeth tending to become bilobate and the tooth rows

becoming parallel. The Haile XVA specimen appears to be

intermediate between Kraglievichia and Chlamytherium in

this character. The fourth alveolus shows that this tooth

was moderately bilobate and nearly parallel with the

posterior teeth. That this could represent individual

variation should not be overlooked, especially when one

considers the limited number of Kraglievichia specimens

known; but when all the chlamythere specimens known are con--

sidered, the trend toward bilobate and parallel teeth seems

evident. Cranial and upper dentition measurements are pro--

vided in Table 6.

Mandible and lower dentition: Partial left and right

mandibles are preserved in UF 10902. In the right mandible

(Figures 5f, g) the seventh tooth is complete while the

eighth is broken off below the level of the alveolus. Al-

veoli for the sixth and ninth teeth are present. The

fragmentary nature of the two jaws makes them difficult to

compare in detail. The major differences between the

mandibles of Kraglievichia and Cilamytherium, according to

previous workers, are in the shape and orientation of the

anterior teeth. Unfortunately, this portion is lacking in

both mandibles of UF 10902.. There appear to be no differ-

ences in ; he pos:'erior port.i-on of the r:andibles except for

Table 6

Cranial and Dental Measurements (in mm) of YKralievichia and Chlamytherium

Kraglievichia Kraglievichia Chlamytherium Chlamytherium
UF 10902 Ai'TNH 2686 UF 889
Haile XVA Argentinal Florida Florida
Widthi- cf nasals at anterior
border of premaxillary 25.9 27.6 ---- 39.0
'i:dth 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
Wiidth, tooth 1 6.8 6.5 7.6
Marnd ib le
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.
Width, tcoth 3 5.2 --- .9
Lei.gth, 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
Width, tooth 9 10.3 --- 13. ----
Width, tooth 9 5.5 __- 7.8---
measurements after Castellanos (1927)
est imated

size. MeasuremenLs o: the lower dentition are provided in

Table 6.

Vertebrae: The vertebral material from the Haile XVA

skeleton consists of seven thoracic, five lumbar, and four

caudal vertebrae. Five of the thoracic vertebrae are

cemented together in the proper sequence by a coarse, large

grained sandstone. Comparisons of these specimens with the

vertebrae of an excellent Chlamytherium skeleton from

Branford IA, Suwannee County (Rancholabrean), show no sig-

nificant morphological differences except for size.

Front limbs: In discussion of the major limb elements,

a comparison is also made with Dasypus. Kraqlievichia is

approximately the same size as Rancholabrean specimens of

Dasypus bellus, a large extinct Pleistocene armadillo.

Conceivably, limb bones of the two could be confused.

The right humerus of UF 10902 is well preserved except

for the idstal end (Figure 6a, b). The humerus of Kraq-

lievichia is greatly expanded laterally (as seen in the

specimens from Santa Fe I), The supracondylar foramen is

relatively larger than in Dasypus. The articular surface

for the radius is concave in Kraglievichia, whereas it has

a slight convexity in Dasypus. The supinator ridge is rela-

tively narrower in Kraglievichia, and the deltoid ridge

is wider-. Thie pr:oximal end of the humerus is similar in the

Figure 6

Kraglievichia paranensis 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 view%

X 0.60 (linear)



two gencra. Comparison between Kraqlievichia and Chlamy-

therium shows no distinct differences in this element ex-

cept for size.

There are three ulnae from the Haile XVA site. Two of

these specimens appear to be from the same individual as

they are similarly preserved, are of opposite sides, and

agree very closely in measurements (Table 7). The ulna of

Kraqlievichia (Figure 6c, d) is laterally flattened and

possesses a long olecranon process. The articular facets

for the radius and medial condyle of the humej.us are a

single structure in this form, while in Dasypus they are

partially divided. Another striking difference is a lateral

groove which runs the entire length of the ulna in Dasvpu:;,

but which terminates at the upper border of the semilunar

notch in Kraglievichia. Size appears to be the only dif-

ference between the ulnae of Kraglievichia and Chlamytheriurn.

As in Dasyous, the distal end of the radius is massive

compared to the proximal end although it is less flattened

in Kraqlievichia (Figure 6e, f). The proximal portion of

the shaft is relatively thicker and less curved in Kragli--

evichia. Comparison with Chlaamytherium shows only a size


The Jeft motcarpnl II is preserved v-ith the Haile XVA

skcle-to,. 'In o.-.Jic aj] shape and proportion it agrees with

Table 7

Measurements (in mm) of Li.im Elements; of Kraglievichla
paranensis, UF 10902, Haile XVA

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 ---- 13.2
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

Metacarpal IT
Total length 32.8 ..
Width, proximal end 10.9 ..
Depth, proximal end 14.5 ----
Width, distal end 10.5
Depth, distal end 11.2

Metacarpal III
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

Ungual Phalanx, Digit V
Total length ---- 29.1
Width, proximal end 11.4
Depthl, ploximal end 10--- .

Table 7 (continued)

Total length
Lateral width, proximal end
Anteroposterior thickness of greater
Lateral width of shaft at third trochanter
Anteroposterior thickness of shaft at same
Greatest lateral width of articular facets
Greatest anteroposterior width, distal end

Tibia and Fibula
Total length
Lateral width, distal end
Anteroposterior width, distal end

Total length
Width of articular facets for astragalus
Depth of facet for cuboid
Width of facet for cuboid

Greatest lateral width
Greatest anteroposterior distance
Greatest depth

Metatarsal II
Total length
Width, proximal end
Depth, proximal end
Width, distal end articularr surface)
Depth, distal end particularr surface)

Metatarsal III
Total length
Width, proximal end
Dep-h, proximal end
Width, distal end (articalar surface)
Depth, distal end (arLicular surface)

Left Right


---- 41.7
---- 41.3

---- 48.4

--- 121.0
---- 25.1





14. 3

11. 5

Unnual Phalanx, Digit V
Total 1 enY 1------ 24.6
Width, p:.o":Lci1; e nd ---- 17.6
LDc]tj-h, t. P_ -___ ---J-0' _,. 7

the same element of Chlamytherium. The facet for articula-

tion with the trapezoid, however, is quite different. When

viewed laterally, this facet is a smooth curve in Chlamy-

therium, while in Kraglievichia it forms a sharp, V-shaped

indention. The facet for articulation with the magnum is ob-

long in Chlamytherium, while it is round in Kraglievichia.

The facet for articulation with the trapezoid is wider dor-

sally in Chlamytherium. This extra width gives the facet for

metacarpal III a different shape. In Chlamytherium it bulges

out proximally to form a quarter diameter for the facet which

articulates with the trapezoid. It appears that less of the

trapezoid articulates with metacarpal III in Kraglievichia~

Left and right metacarpals III are preserved in UF 10902.

In this element the facet for articulation with metacarpal

II and the trapezoid shows a relatively greater association

with the trapezoid than with the adjacent metacarpal. The

facet for the magnum in both Chlamytherium and Kragllievichia

is convex dorsally and concave ventrally. In Kraqlievichia the

greater portion of the facet is convex, while the opposite is

true in Chlamytherium. The ventral portion of this facet,

when viewed from the ventral aspect, is parallel in a line at

right angles to the long axis of the bone in Chlamytherium

while in Krajlievichia it is oblique.

in. limbs: The femur (Figure 7a, b) and the tibia and

Figure 7

Kraglicviclia paranensis Hind Limb Elements

A. Right femur, anterior view

B. Riyht femur, lateral view

C. Right tibia and fibula, anterior view

D. Right tibia and fibula, lateral view

X 0.60 (linear)



fibula (Figure 7c, d) show no morphologica] differences from

those of Chlamytherium except for size.

The calcaneum of Kraqgievichia is relatively loss ex--

panded at the distal end, and is a relatively less robust

element than that of Chiamytherium (Figure 8). The facets

for articulation with the astragalus are quite different.

In Kraglievichia the facets are connected, forming a double

facet, while in Chlamytherium they are separated by a central

valley. The relative proportions of the two facets also

differ. In Chlamytherium, the lateral facet is much larger

than the medial facet, while in Kraglievichia they are more

nearly the same size.

The navicular of Kraglievichia shows no appreciable

differences from that of Chlamytherium. A comparison of the

three right naviculars from Haile XVA with one another shro,..

only a slight variation in the relative shapes and sizes of

the facets for articulation with the cuneiforms.

Regarding the metatarsal II, the facet for articulation

with the mesocuneiform is relatively narrower ventrally in

Kragqlievichia than in Chlamytherium. In Chlamythorium the

medial border of metatar;a] II forms an unbroken line, while

in K]ag.ievichia there : is an indonLatJon at the proximal

end to accept the pi.oxiimal cend of meLalarsal I. There must

have icci, a crurresponding bulge in the proximal end of

Figure 8

Kraglievichia and Chlam-rLheorium Calcanea

T. Kraglievichia paranensis right
calcaneum, Hai.le XV

B. ?Kraglievichia sp. right calcaneum,
Inglis :CA

C. Chlamnytherium s epten.lrionalis right
calcaneum, Branford IA

All approx. natural size


metatarsal I of Kraqlievichia; no such feature occurs in

metatarsal I of Chlamytherium.

When viewed laterally, the proximal articular surface

of metatarsal III appears rounded in Kraglievichia while in

Cllamyth erium it forms a straight line, perpendicular to the

long axis of the element. Due to the rounding of the

proximal end of this element in Kraglievichia, the articular

surface can be seen when the element is viewed from above;

this is not the case for Chlamvtherium. This would appear

to permit more dorsoventral movement of the toes of Kra_-

lievichi a.

Regarding metatarsal IV, the facets for articulation

with metatarsal III and the cuboid are a single structure in

Kraqlievichia, while they are two separate structures in


Two ungual phalanges of UF 10902, both from digit V,

have been recovered from Haile XVA. One is rather narrow

and pointed, and the other is blunt and broad. Apparently

the ungual phalanges of the front feet were pointed, while

those of the hind feet were blunt. The same is true in

Chlamvtherium. Measurements of limb elements of other

examples of Kraqlievichia from various Florida localities

are presented in Table 8.

Table 8

Measurements (in nun) of Limb Elements of Karaqj.ievichia
paranensis From Various Florida Localities

UF 10432 (right) UF 9354 (right)
Santa Fo I Santa Fe I

Greatest anteroposterior
diameter of shaft
Lateral width of shaft at
same point
Lateral width, distal end
Greatest width of distal
particular surface


Total length
Lateral width, proximal end
Lateral width, distal end
Lateral width at semilunar
Anteroposterior width at
semilunar notch




UF 10449
Sante Fe



(left) UF 17474 (right)
I Haile XVA






Total length
Transverse width, proximal end
Anteroposterior width, proximal end
Transverse width, distal end
Anteroposterior width, distal end


Lateral width, proximal end
Anteroposterior width, proximal end
Lateral width of shaft at third trochanter
Anteroposterior width of shaft at same

Calcan um

ToLal length
Width of ar-ticular facets for astragalus
Depth of: rac>: for culo Ld
Width 'l-j cr' t f-" or, cuboid

UF 10830
Sante Fe


UF 17476 (left)
Haile XII B


UF 17473 (left)
Santa Fe II

Table 8 (continued)


Greatest lateral width
Greatest anteroposterior
Greatest depth

UF 17568 (right) UF 17569 (right)
Haile XVA Haile XVA
37.3 33.6



Metatarsal II

Total length
Width, proximal end
Depth, proximal end
Width, distal end articularr surface)
Depth, distal end articularr surface)

Metatarsal III

Total length
Width, proximal end
Depth, proximal end
Width, distal end articularr surface)
Depth, distal end articularr surface)

Metatarsal IV

Total length
Width, proximal end
Depth, proximal end
Width, distal end articularr surface)
Depth, distal end particularr surface)

Haile XV

UF 17472
Santa Fe

UF 17475
Santa Fe


(ri gh t)

righth )

__1_1____~_ I~_____ _I~_ _I

Review of the ch]amythcriinae

Since the genus Kraqglievichia is reported here for the

first time in North America, this seems to be an approp-riate

place for a review of this group on both continents. The

nomenclatural history of the various chlamytherje genera will

first be discussed in the order of their introduction into

the literature. This will be followed by a brief review of

each genus, in geological order, 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).

In his early works Lund referred to this genus as Chlamy--

therium, but later (beginning in about 1840) he began to

call the genus Chlamydotherium without giving any reason'

(Castellanos, 1927). Later authors used the name Chlamy-

dotherium, apparently not realizing that this name, in the

meantime, had been given to a genus of glyptodonts by Bronn

(1838). In 1875 Amcghino proposed the name Pampatherium

for this genus, recognizing that the name Chlamydotherium

was occupied by the glyptodont genus. lie later abandoned

this name \.hen he realized that Lund had previously named

the genus ChlanmyLherium. Paula Couto (1956) has argued

1thliaL Pam at'horj .iu'- n! s a valiCd name, as Lund meant to call it

Chlanvly0'a Lhae.r,' .

The first North American record of ChlamLyherium was

reported by Leidy (1889a) when he designated the species

Glyptodon septentrionale. In the same year Leidy, in

another work (1889b), referred these specimens to the South

American species C. humboldtii. Sellards (1915) believed

that the North and South American forms represented differ-

ent species and revived Leidy's original trivial name s_-


In 1902 Ameghino described Machlydotherium from the

Eocene of Patagonia.

Castellanos (1927) named two new genera of chlamytheres:

Vassallia, based on an edentulous mandible and several dermal

plates; and Kraglievichia, based upon two skulls, a mandible,

and a small amount of postcranial material. The genotypic

species of Vassallia was originally Chlamytherium minutum

(Moreno and Mercerat, 1891). Kraqlievichia was erected to

include C. paranensis, C. intermedia (Ameghino, 3887), and

C. subintermedius (Rovereto, 1914). The trivial name

paranensis was retained.

Simpson (1930) established the genus Holmesina while

studying excellent material of septentrionalis from the

Seminole Field in western peninsular Florida. Subsequent

authors have tended to refer: to the North American forms as

Holmos:ina and t.o the South American forms as Ch.lamvtherium.

CastelJanos later (1937) named P1 aina, based upon the

type of C. intermedius which he had earlier placed in Krag--

lievichia. The reason for the establishment of this new

genus was his interpretation of the lineage of the chlamy-

theres. He believed that there should be a form intermediate

in size between Kraglievichia and Chlamytherium. Since C.

intermedivs is larger than the other material referred to

Kraglievjchia, he saw it as representing this intermediate


The genus Iloffstetteria of Castellanos (1957) was based

upon a skull collected in Ecuador. This skull had previously

been described as C. occidentalis by Hoffstetter (1952).

Each of the chlamythere genera will now be briefly re-

viewed in geologic order beginning with the oldest, Machly-

dotherium. This form is known from the Eocene of Patagonia

and its relationship to the later chlamytheres is not known

(Simpson, 1945). Chlamytherec are not known frcm the Oligo-


Vass .lia precedes Krcgli evichia temiporally as it is

known from the L; Vcnta faiuna of Miocene age in Colombia.

The author has examined a skull from this locality (U.C.M.P.

40401) -which definitely represents Vassallia. This idenLi-

fication is ba-r.ed upon its small s.ize, and the nature of the

ainterjor o ;:iion, in which the first five tooth are poq-


like and rotated lingually. Vassillia has also been reported

from Auracanean deposits (Castellanos, 1946) so it appears

that this genus did range into the Pliocene.

Porta (1962) has reported Kraglievichia from the La

Venta fauna in Colombia, but this record was based only

upon dermal plates which could just as well represent

Vassallia. Other Miocene records of Kraglievichia have been

reported as some previous works considered this to be the

age of the Auracanean deposits; however, more recent in-

terpretations of South American stiatigraphy (Patterson

and Pasqual, 1968) demonstrate the Auracanean to be Plio-

cene rather than Miocene. Therefore, it appears T:hat Jra; -

lievichia was restricted to the Pliocene in South America.

Castellanos (1927) has reported morphological diff.r-

ences between Vassallia and Kraglievichia, In Vassallia

the first five teeth tend to be peg-like and rotated lin-

qually while only the first four teeth of Kraqglievichia show

this condition. Just how variable this characteristic is

cannot be ascertained presently because of the lac: of

comparative material known for these genera. Vassallia is

also significantly smaller than Kraglievichia. It is sus-

pected here (as Castcllanoc has suggested) that Vassallia

is the ancestor of K'-'r i :--i vichia.

Regarding Plajir. v'which is based on three isolated

dermal plates, Castellanos (1927) was probably correct in

his first judgment when he placed this material in the genus

Kri alievichia. lie considered these plates to be inter-

mediate in size and sculpLuring between Kracq1ievichia and

Ch]amytherium. They seem, however, to fall within the size

range expected for the older armadillo genns. Furthermore.

it seems quite difficult to interpret slight variation in

plate rugosity as being generically significant. It is

therefore proposed here that Plaina is a synonym of Krag-


Thle genus Ieffstetteria (Castellanos, 1957) is based

upon supposed differences in the shape and measurements of

the teeth. The deviations, however, are clearly attributable

to individual variation. Hoffstetteria is here considered

to be a synonym of Chlamyt]Lerium.

In Simpson's (1930) description of Holinesina, he listed

a number of characters in which this new genus differed

from Chla myttheriumn. However, James (1957) has shown that

the characters pointed out by Simpson are not sufficient to

separate the North and South American forms generically, and

that Holmesina is a synonym of Chlamytherium. This report

strer'l;thens James' views on this issue. Simpson (1930) has

also argued thar if Ho.lmesina is noc volid, then all the

Soui:h A ru.i.;an iol0-s shloull be placed in t h genus Chlamny--

therium as they are no more different from each other than

Chlarytheriumn is from Holmesina. There may be some justifi-

cation for doing just this. In this report, however, new

characters of Kraglievichia, different from Chlamytherium,

have been pointed out which, together with what previous

workers have considered generic characters, justify retain-

ing Kraqlievichia as a valid genus.

The genera of chlamytheres, then, recognized in this

report are Machlydotherium (Eocene), Vassallia (Miocene and

Pliocene, South America), Kraglievichia (Pliocene, South

America and early Pleistocene, North America), and Chlamv-

therium (Pleistocene, North and South America).

Evolution of the chlamytheres

The principle morphological trend in Chlamythere evolution

since the Miocene was an increase in size. Vassallia was

the smallest form in the known chlamythere lineage, followed

by Kraglievichia which was slightly larger. An examination

of the two femora in Figure 9 shows the great size difference

between Kraglievichia and Chlamytherium. The intermediate

steps in this size increase are illustrated in Figure 10,

which shows dermal plates from deposits representing various

stages of the Pleistocene. The Blancan Haile XVA plates

are the smallest, followed by those from the Irvingtonian

Inglis TA site. Slightly larger still are the plates from

Figure 9

Kraqlievichia and Chlanytherium Femora

A. Chlamytherium septentrionalis right femur
Branford IA

B. Kraglievichia parancnsj.s right femur,
Haile XVA

X 0.60 (linear)



Figure 10

Kraglievichia and Chlamytherium Dermal

A. Kraglievichia paranesis, Haile

B. ?Kraglievichia sp., Inglis !A,

C. Chlamytherium septentrionamlis,
late Irvingtonian

D. Chlamytherium soptentrj onalis,

X 0.40 (linear)


XVA, Blancan

Early Irving-

Coleman IIA,

Branford IA,


-. -

* *1-

. *t

* .

i.he later Irvingtonian Colcmpn IIA site, and the largest

plates of all are those from the Rancholabrean Branford IA


A trend from peg-like to bilobate teeth, and from

oblique to parallel tooth row axes was previously noted.

3oth of these trends seem to have extended from the rear

of the dentition toward the front, as the number of bilobate

and parallel teeth increases from Vassallia through Chlamy-


Concerning the geography of chlamythere evolution, it

is apparent that these forms originated in South America

as all pre-Pleistocene records of this group are restricted

to that continent.

The earliest known Chlamythere is the Eocene Machlvdo-

therium, and no chlamytheres are known from the Oligoccne,'.

Vassallia is present in Miocene deposits of South America,

and both Vassallia and Kraglievichia were present in the

South American Pliocene. Kralievichia is known from North

American early Pleistocene doposiLs, but its presence in

South America during the Pleistocene is questionable. Chla-

mytherium is known f.:romi the Pleistocenc: of North and South


It has gener.r.lly boc; LeIio\vcd f-hat- the evolution of

the chlamythere. 'ts b'.c .lt-ricLly a South American phcnom-


enon, '.*ith only the end product, Chlamy1thoerium, migrating to

North America during Rancholabrean time. The evidence sup-

porting this hypothesis is, briefly, that all pre-Ranchola-

brean records of chlamytheres have been from South America.

and all genera of chlamytheres except Chlaniytherium seemed

to have been restricted to South America. The study of the

Haile XVA fauna, and other pre-Rancholabrean sites of

Florida, has yielded conclusive evidence warranting modifi-

cation of this Theory.

Apparently Kra glievichia migrated to North America

shortly after the establishment of the late Cenozoic land

bridge between the two continents. This is substantiated

by its presence in the Haile XVA and other Blancan faunas

of Florida.

Three hypotheses can be offered regarding the zooge-

ography of the evolutionary transition from Kraglievichia to

Chlanmytherium. The first hypothesis, already discussed, is

that the evolution of Chlamytherium took place in South

America and this genus then moved into North America in

Rancholabrean times. This interpretation has already been

shown to be in erro .

The second hypothesis suggests that Kragilievichia moved

into North America during the ;latest Pliocene and became

ex;--inct in gouth America. Tijs would make tlh e.\iolultion of

Chlamyllorium str.icLly a North American cveInt. According

to this hypothesis, Chlamy Lherium, a North Amc;.ican form,

then reinvadcd South America in the later Plcistocene. This

would account for the apparent lack of chlamytheres in the

early Pleistocene of South America.

The apparent lack of chlamytheres from t-.e Argentinian

Chapadmalalan fauna should be briefly discussed here. A

dermal plate has been questionably attributed to t)his faurna

(Kraglievich, 1.934); however, the specimen was from a local-

ity near the major deposit and its horizon is actually un-

known. Because of their unusually high number of bony

plates available for preservation, if the chlamytheres we-c-

present at all i.n the fauna, they should be well represented.

It is possible that their absence from the fauna is ecelo.-

ically determined. The early Pleistocene Chapadmalal fauna

represents a temperate part of the continent, whereas the

chlamytheres may have been restricted to the more tropical

parts of the continent. The lack of early Pleistocene de-

posits in tropical South America hampers a definite estab-

lishmennt of the presence or absence of these forms on that

continent at that time.

The third hypothesis to be considered is that the di:e--

tribut2.on of ch lamytheres h1i.as been continuous o:n both con-

tincnts t-hrc,-oi0 the Pleistoenre, and the evolution of


Chlamytherium from Kra gievichia occurred simultaneously in

North and South America. That is the hypothesis favored here.

Dasypus bellus Simpson 1929

Material: UF 16698, left nasal, right maxilla and man-

dible, right astragalus, 51 isolated dermal plates.

The preceding material probably represents one individu-

al as the maxillary and mandible articulate nicely and all

the material (with the exception of some of the plates) was

found closely associated. Comparisons are made here with

other specimens of D. bellus and with its closest living

relative, D. novemcinctus. The only other known D. bellus

material containing teeth is from Crankshaft Pit, Missouri

(UK 15544, Oesch, 1967). Postcranial comparisons are made

with material from a number of sites of various ages in


The nasal bone is represented by only the anterior

portion and is larger than, but morphologically similar to,

that of D. novemcinctus.

In discussing the dentition, teeth will be designated

as follows: T will refer to the first upper tooth, T2

the second upper tooth, and so forth. The same procedure

will be used in referring to the lower teeth.

The maxillary is fragmented and contains onl T3 T6

Except for size, the lateral portion of -hi s specimen coin-


pares favorably with the corresponding portion of the Crank-

shaft Pit specimen. The first three teeth of UF 16698 were

probably transversely flattened as this is the nature of T3

The remaining upper teeth were probably round and peg-like

4 6
as this is the condition in T T. The Crankshaft Pit

skull is from a juvenile individual, and TI has not completely

erupted. In this specimen the first tooth is transversely

flattened, but all the rest are round and peg-like. In D.

novemcinctus the first three teeth tend to be somewhat

laterally flattened, but not as much as in the Haile XVA

specimen of D. be].lus. That tooth number and structure is

variable in Dasypus has been pointed out by Talmage and

Buchanan (1954), as well as by others. In four D. novem-

cinctus skulls examined, the total number of teeth varied

from seven to nine.

The mandible of UF 16698 (Figure 11) is lacking thic

proximal end. The number, shape, and location of the mental

foramina are highJy variable in this genus. In both known

D. bellus mandibles there are two foramina, one large and

one small. In UF 16698, the larger of the two foramina

occurs between T2 and T3. The smaller of the two foramina

is locaLed 38 mm alLterior to TI. In UK 15544, the larger

foramen is anLnerior to the mailerer and is located directly

belnea;.h '. The smaller is .located beneath T,. In D.

Figure 11

Dasypus bells Mandible

UP 1.6698, Hailc XVA

X 2.26 (linear)



K..... ',
d'. .-


C' '


novemcincLus, the number of mental- foramina varies from one

to four. The symphysis of the Haile XVA specimen is weak,

which is characteristic of the genus.

The mandible of UF 16698 lacks T T5. Ti T3 are

laterally flattened ]but; T6 and -7 are round. This differs

from the lower dentition of UK 15544 in which all the teeth

are round.

Except for smaller size and a tendency toward flattening

of the anterior teeth, the mandibles and dentition of the

Haile XVA specimen differ little from other Dasypus material.

Dental measurements are provided in Table 9.

Differences in the astragalus of the Haile XVA D. beljus

and other specimens of D. bellus from various Florida

Pleistocene sites and Recent specimens of D. novemcinctuvs

are insignificant except for size. A comparison of the

astragali of Dasypus from various stages of the Pleistocene

reveals a size trend similar to that of the chlamytheres,

with one exception (Figure 12). The size of Dasypus in-

creased from Blancan through Rancholabrean time, then de-

creased in Recent times when it is represented by D. novem-


The dermal plates of UP 16698 do not differ from any

ether specimens of Dasypus except for size. Martin (in

press) has rv.m7-:sured samples of plates from several Florida

Table 9

Dental Measuremonts (in mmn) of Dasypus bellus

UF 16698
Haile XVA

Total distance, teeth 3-6
Anteroposterior distance, tooth
Transverse distance, tooth 3
Anteroposterior distance, tooth
Transverse distance, tooth 4
Anteroposterior distance, tooth
Transverse distance, tooth 5
Anteroposterior distance, tooth
Transverse distance, tooth 6

Depth of ramus at tooth 1
Width of ramus at tooth 1
Depth of ramus at tooth 6
Width of ramus at tooth 6
Depth of ramus at tooth 8
Width of ramus at tooth 8
Length of tooth row
Anteroposterior distance, tooth
Transverse distance, tooth 1
Anteroposterior distance, tooth
Transverse distance, tooth 2
AnteroposLerior distance, tooth
Transverse distance, tooth 3
Anteroposterior distance, tooth
Transverse distance, tooth 4
Anteroposterior distance, tooth
Transverse distance, tooth 5
Anteroposterior distance, tooth
Transverse distance, tooth 6
Anteroposterior distance, tooth
Transverse distance, tooth 7
Antcroposterior distance, tooth
Transverse distance, tooLh 8

3 3.5
4 4.4
5 4.0
6 4.2

1 3.0
2 3.3
3 3.3
4 4.81
5 4.01
6 4.5
7 4.6
8 4.11

UK 15544
Crankshaft Pit
left right




4. 7




2-<1 i VcO a i-, c'st.i ii t_ -cL
"';CO! 1 co iio.l ele .v, orklpLcd

______ ___

Figure 12

Dasypus bellus Astragali

A. UF 16698, Haile XVA, Blancan

B. UF 13187, Coleman IIA, Irvingtoniar

C. UF 2478, Mefford Cave, Ranchola'brex

X 2.23 (linear)


.'-. . ,, .-

.; X , ,

" :

... Y%^

sites and has shown little overlap in plate size between

D. bellus and D. novemcinctus. His measurements indicate

that the size of the plates do not follow the trends through-

out the Pleistocene which are seen for other parts of the

skeleton. The Coleman IIA plates are small, and overlap

slightly with those of D. novemcinctus. However, this was

probably due to the small sample of plates used. There are

approximately 2,500 plates in a single armadillo carapace

and little or no satisfactory method for determining the

area of the shell from which most of them come. Therefore,

size trends based on plates should involve large samples to

insure accuracy.

Inasmuch as the morphological differences between the

Haile XVA material and other D. bellus material are so

slight, and those which were observed probably represent

individual variation, this material is referred to Dasvpus


Dasypus bellus has typically been considered a southern

North American species of Rancholabrean age. Martin's (in

press) description of the Coleman IIA fauna, however, ex-

tended the temporal range of the species back to Irvingtcnian

time. This report extends this species still further back,

into the Blancan, and is thus the earliest known record.

There is a question as to the relationship between D.


hoellus and D. novemcinctus. These animals are clearly more

related to each other than to any of the other armadillos

(Auffenberg, 1957). The question arises as to whether D.

novemcinctus was derived from D. bellus or existed allo-

patrically with D. bellus during the Pleistocene, replacing

it during the last few thousand years.

Considering the former possibility first, it would have

taken quite rapid selection for smaller size to develop

D. novemcinctus from D. bellus. In Miller's Cave (Patton,

1963), a date of ca 8,000 BP is given for the Travertine

stratum bearing D. bellus remains, and a date of ca 3,000

BP for the Brown Clay deposits containing D. novemcinctus.

According to Patton (personal communicationn, even if the

D. novemcinctus is a Recent intrusive (which he doesn't

believe to be the case), the difference between 8,000 EB

and 3,000 BP, or 8,000 BP and 1,000 BP is not that great--

it is still rapid evolution, but possible, particularly if

the only change is size.

Concerning the second hypothesis, if D. novemcinctus

existed somewhere else during the Pleistocene and replaced

D. be llu-s very late, it should be known from some Central

or South American Pleistocene deposit. To date, however,

D. rnoovemcinctus has no fossil record except for Miller's

Cave, ibut te fo...il. i.ccord in tropi cal America is extroe.cly


That D. bellus is not known from pre-Rancholabrean de-

posits outside Florida is puzzling. Slaughter (.1961) lists

two conditions which he feels were necessary for the occur-

rence of this species: winters could have been no more

severe than those of North Central Texas today; and rainfall

probably had to be more than 20 inches per year. That

temperature and rainfall could have restricted D. bellus to

Florida during the entire Pleistocene is possible, but it

seems more likely that the known distribution of this species

is a reflection of a lack, outside of Florida, of Gulf

Coastal Plains sites of the proper age.

Order Lagomorpha

Family Leporidae

Sylvilaqus sp. Gray 1967
Material: UF 17561, 2 M UF 17562, M3; UF 17563,

lumbar vertebra; UF 17564, 3 innominates; UF 17565, 2

femora; UF 17566, 1 tibia; UF 17567, 2 metatarsals.

Hypolagus and Svlvilagus are two smaller lagomorphs

common in North American Blancan deposits. Unfortunately,

the generic diagnosis provided for ILypolagus by Dawson (1958)

does not include discussion of either M" of i4M, which are

the only teeth preserved in the Haile XVA fauna. A compari-

son of the innominaLes from Haile XVA with the descriptions

and figures of these elements of Iy__o.la-u- in Dawson (1958)

shows th.e IJaile specimens possess a less elevated iJlial

crest, a condition considered to be more advanced. A31

the rest of the Haile XVA material also agrees in size and

morphology with Sylvilaqus.

Unfortunately, a lack of diagnostic elements prevents

a species determination. Two living species are present

throughout Florida: S. palustris and S. floridanus. The

present range of the genus is throughout North America, with

two species being present in South America: S. floridanus

and S. brasielcesis (Hall and Kelson, 1959).

Previous Blancan records of Sylvilaqus include the

Curtis Ranch fauna (Gazin, 1942) and the Broadwater fauna

(Barbour and Schultz, 1937). Numerous Irvingtonian and

Rancholabrean occurrences of this genus are recorded through-

out North America.

Order Rodentia

Family Sciuridae

Petauria sp. Dehm 1962

Material: UF 12353, partial right mandible with M3.

By Miocene Lime flying squirrels had diversified into

two groups (James, 1963). One of these groups has compli-

cated cheek tooth loph patterns. T'ijs is due to the presence

of "protoLoph"u1.cs and metalophules, and especially meta-

lophulcs thai. c:xt.-cnd from the met loph posteriorly to -th

posterior cingulum, dividing the posterior valley into two,

and sometimes three or more smaller valleys." (James, 1963)

This complicated toothed group contains several living genera

including Petaurista, Belomys, Trogopterus, and Sciuropterus,

as well as thefossil genera Pliopetaurista (Kretzoi, 1959),

Petauria (Dehm, 1962), and Pliosciuropterus (Sulimski, 1964).

The second group has simpler loph patterns in its cheek

teeth. This group is represented by the living genera

Hylopetes, Aeromys, Eoglaucomys, Glaucomys, and others, and

by the extinct genus Pliopetes (Sulimski, 1964) and several

extinct species referred to Sciuropterus.

Formerly, fossil species referred to Sciuropteriis in-

cluded both simple and complex toothed types. Hugueney and

Mein (1966), however, have placed the complex toothed fossil

species of Sciuropterus in the genus Pliopetaurista based

on Deperet's (1897) S. pliocenica. The result is that all

fossil species remaining in the genus Sciuropterus in both

the New and the Old World have simple teeth, while the living

species which are restricted to the Old World have compli-

cated teeth. This curious arrangement leaves some doubt as

to the evolutionary unity of the fossil and recent species

referred to Sciuropterus.

The Haile XVA specimen (Figures 13 and 14) marks the

first occu.-rrnc' in the New World of a member of the group

Figure 13

Petauria sp. RighL Mandible

UP 12353, Haile XVA

X 7 (linear)



Figure 14

Petauria sp. Right M3

UP 12353, Haile XVA

4 mm


with complicated tooth. It is clearly uore closely related

to Petaurista than any of the other living genera.

The Haile XVA specimen (UF 12353) differs from P.io-

petaurista in that its 143 is oval in shape as opposed to

being pointed at the posterior end. The metastylid in UF

12353 is only moderately developed as opposed to being well

developed in Pliopetaurista, and the postflexid is not as

well developed in the Haile XVA specimen.

UF 12353 differs from the M3 of Pliosciuropterus in its

shape, which is narrow and tapers toward the posterior end,

as opposed to being oval in the Haile XVA specimen. The

mesoconid is; only moderately developed inthe Haile XVA

specimen while in Pliosciruopterus it is so well developed

that it forms a labial spur. The postflexid is also well

developed in Pliosciuropterus.

UP 12353 is clearly more allied with Petauria than any

of the fossil genera. On the other hand, one striking dif-

ference separates these forms: the mesoconid is completely

absent in Petauria, but it is moderately well developed in

the Haile XVA form (Figure 13 and 14)

The 1;j.ie XVA specimen shares one important characteris-

tic wilth PetaurisLta that does not occur in Pr taur:ia, namely

the pr.c ccnc,- of a n'asocon.i' On th-c' other hand it shares

the follo.'.. characteris i cs w'th P]tox'ij a which do not


occur in Petaurista: the grinding surface of the talonid is

less complex; and the metastyiid is moderately developed

as opposed to the metaconid-metastylid chaos (McKenna, .962).

Finally, it shares the following characteristics with both

PeLaurista and Petauria: the grinding surface of the trigo-

nid is complex, with extra lophids and fossettids; the post-

flexid is poorly developed; the M3 is oval in shape; and

the entoconid is connected to the hypoconid through the


Since the Haile XVA specimen shares characters with

both Petaurista and Petauria, assignment to a genus is dif-

ficult. Whether the lack of a mesoconid is more important

than the presence of extra fossettids and lophids on the

talonid would probably be a disputed point among students of

rodent evolution. However, Petauria is known from only one

dentition, and the variations which may have existed are not

known. Petaurista, being extant, has been studied in

greater detail and its variations have been taken into

account. Thus characters known to be peculiar to Petau-

rists are more reliable than those in the diagnosis of

Petauria. The simple grinding surface of the talonid (not

observed in a very large sample of Petaurista) becomes a

more important character with regard to the Haile XVA

specimlin Tvhan the presence of the mesoconid, \.which may


have occurreC in Petauria. Following this line of reasoning,

.it secms more appropriate to refer the Uaile XVA specimen

to Petauria.

The only specimen of Petauria previously known is from

Bavaria, in the Region of Eichstatt, The deposit was a

fissure filling of red loamy sand in the Solcnhofn Lime-

stone. The filling has been ascribed to the "older Pleis-

tocene" by Dehm (1962).

It is almost certain that the populations of flying

squirrels of Florida and Bavaria during the early Pleisto-

cene were reproductively isolated; thus the Haile XVA

specimen isnot assigned to the type species, P. helleri.

It also seems pointless at this time to describe a new

species based upon this single specimen, as even the

generic characters are difficult to ascertain with this

limited amount of material. Until more material is

available, it seems more prudent to refer to the Haile

XVA specimen as Petauria sp,

Family Castoridae

Castor canadensis Linnaeus 1758

lMateriai: UF 17489, left fmur.nr.

This element, lackin the proxiimal end, represents a

young individaJ It shows no morphological differences

from the :iv.inc specie.r. A comparison of measurements

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