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FIELDIANA

Geology

NEW SERIES, NO. 30

Giant Short-Faced Bear (A rctodus simus yukonensis)
Remains from Fulton County, Northern Indiana

* Ronald L. Richards
William D. Turnbull

With an Appendix by E. J. Neiburger

JUN 2 9 1995

April 28, 1995
Publication 1465

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.

Grubb, P. J., J. R. Lloyd, and T. D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador.
I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601.

Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. Jn Browman, D. L.,and R. A.
Schwarz, eds., Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands.

Murra, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American
Indians. Vol. 2, The Andean Civilizations. Bulletin 143, Bureau of American Ethnology, Smithsonian
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KELLDIA:

Geology

NEW SERIES, NO. 30

Giant Short-Faced Bear (Arctodus simus yukonensis)
Remains from Fulton County, Northern Indiana

Ronald L. Richards

Indiana State Museum
Department of Natural Resources
202 North Alabama Street
Indianapolis, Indiana 46204

William D. Turnbull

Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605

With an Appendix by E. J. Neiburger

Accepted August 22, 1994
Published April 28, 1995
Publication 1465

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY |

© 1995 Field Museum of Natural History
ISSN 0096-2651
PRINTED IN THE UNITED STATES OF AMERICA

GEOLOGY LIBRARY

Table of Contents

PAISTRAGCT 620 to estan ait ee Pee 1
INTRODUCTION 529 Sori st ne Cee aes 1
OCCURRENCE 625s eae Se Aas eee 2
TRSCRIPTION eRe Se Serer Na ee es y
Skull, Dentaries, and Teeth ............. 15
VOTCIT AR i ret ihe owuterE etre atnetats 18
WRG or fosg oe kere ie a ee eC aes 18
Forehimb and Gitte en neces 19
Bind mn and <sirdie. 3 oe ee 19
PRENGIORY nd Be Cire ae vl aoe VER els 23
DISCUSSION AND CONCLUSIONS ............. 25
ACKNOWLEDGMENTS 56 655 300 Se ceae ne. oiane 31
LITERATURE CITED 2400 os) ose es teas 32
PPPENTAR ou hires sot een eee ENS 34
List of Illustrations
b. Skull— Three Views 2 3 oi stone 3
2. Skull— Ventral view and enlarged denti-
TIGR A Sr eto ce eee wile Cate ure ere 4
ER OWOE SAW helt in re bo ears eee oes 5
4. Cervical vertebrae— Atlas, axis, and C-4 6
5. C-6 and three pathologic thoracic verte-
DERG 02 ee ee ee ea ee esa ek rat 7
6. Thoracic and lumbar vertebrae ......... 8
7. Scumbar vertebrae. 5. eee kone eee 9
8. Three pathologic thoracic vertebrae—
articulated, and articular surface of two 10
oe 0c. Seen aan art Aer ee et POL IL Ey Oe Pera 11
CO SCADUIBE! cote aie ee ror coo Pees 12
1A) AS 5 1 Crs For 9 Wiens era VaR delim eA tin oo ar 13
I? Radi and inks. oak eres 14
13..-Pelvis :and:sacrum: (24.5 4:2372-625-5n80).: 15
14. “Hind limb elements: 2.25.55 cs ewe oe 16
PS. WIRE DAINOIOMIES: Fe ote eae ee 17
16. Diagram of rib measurements ......... 18
17. Bivariate graph—Skull length by zygo-
matic width of Indiana specimen com-
Paved: Wi OUIETS eo A acces ee 28
18. Bivariate graph— Length and anterior
WIG OE NO oa arte ot er See 29
19. Bivariate graph— Length and posterior
WIE OLE occ wet siete eee ena 30
20. Bivariate graph— Femur, greatest length
and greatest distal- width... 2525. 752.5- 30

ili

List of Tables
1. Cranial and dental measurements ....... 19
2. Dental measurcments:; .i0...55 1s. y ven es 20
3. Vertebral. measurements. ......6.54.46.%. 22
4; Rid measurements: 2.5962 cs 5 one eek 24
5. Forelimb and girdle measurements ...... 25
6. Hind limb and pelvis measurements 26
7. Podial Measurements 2565-50558 Fetes 26
8. Statistical comparison of the Indiana Arc-
todus specimen with populations from
Potter Creek Cave, Rancho La Brea, and
Fairbanks, “AlgskA. 36 ions okies anes 27
9. Other Arctodus dimensions, ratios, and
COMPALISONS een cee cae bin 31

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a .

Giant Short-Faced Bear (Arctodus simus yukonensis)
Remains from Fulton County, Northern Indiana

Ronald L. Richards William D. Turnbull

Abstract

Major portions of the skeleton of the extinct giant short-faced bear (Arctodus simus yukonensis)
were recovered from Fulton County, northern Indiana, in 1967. The bones provided a radio-
carbon age of 11,500 + 520 years B.P. (apatite fraction). The skeleton was deposited in a shallow
lake, perhaps in or near an open or patchy boreal forest dominated by spruce, after the recession
of Wisconsinan ice. The bear was large, comparable in size to Rancholabrean specimens from
Alaska and the Yukon. It is the only tremarctine recorded from Indiana, is one of two specimens
from the Great Lakes region, and is one of the most nearly complete skeletons known. Most
of the other known specimens are highly incomplete, so the Indiana specimen provides a firm
basis for assessing variation in the subspecies. The skull is relatively short compared to the
limb bones, is markedly wide, and has a very narrow interorbital width. Several of the vertebrae
and limb bones exhibit pathological conditions. The pathologies were described previously and
are further described here and discussed by Neiburger in the Appendix. A full set of measure-
ments is presented. Comparisons are made with other A. simus specimens, yielding important

data that have not previously been reported.

Introduction

In 1967 Mr. William Lane of the Northern In-
diana Public Service Company (NIPSCO) un-
earthed and collected some large bones while op-
erating a backhoe. He contacted the Department
of Biology, North Central Campus, Purdue Uni-
versity, Westville, Indiana, transferring the re-
mains to Dr. Greta Woodard. After identifying the
remains as those of Arctodus simus (giant short-
faced bear) with the aid of one of us (W.D.T.),
plans were laid for a joint report on the bones.
With the untimely death of Dr. Woodard, that
report was not realized, and the bones, returned
to the Field Museum in 1970 by Dr. Woodard’s
spouse, were deposited in the fossil mammal col-
lection (catalogue PM #24880). The specimen was
used by Jay H. Matternes in his reconstruction of
Arctodus simus for an article by Guthrie (1972)
and for a life-sized restoration of the bear in a
mural at the National Museum of Natural History,
Smithsonian Institution. In 1983 we undertook a
study of the specimen, reported herein. A maga-

zine article subsequently illustrated a lateral view
of the articulated skull and jaws (Richards, 1983).

Arctodus simus is a large tremarctine with rel-
atively robust dentition and skeleton and propor-
tionately long legs (Kurtén, 1967). It is known from
more than 100 localities in North America, rang-
ing in age from middle Irvingtonian through Ran-
cholabrean (Kurtén, 1967; Richards et al., in press).
Kurtén (1967) recognized two subspecies, the
smaller Arctodus simus simus and the larger A. s.
yukonensis. There is some debate over whether
the bear was an omnivore/herbivore or a predator
(Emslie & Czaplewski, 1985; Voorhies & Corner,
1986).

The skeleton reported here is the first record of
Arctodus simus in Indiana and one of two speci-
mens from the Great Lakes region. The second
specimen was recently recovered from a cave in
Wyandot County, Ohio (Hansen, 1992; H. G.
McDonald, pers. comm. to Richards, 6 Jan. 1992).
Richards et al. (in press) present an expanded cov-
erage of North American Arctodus simus localities
and materials.

FIELDIANA: GEOLOGY, N.S., NO. 30, APRIL 28, 1995, PP. 1-34 1

Occurrence

The remains were found south of Rochester,
Fulton County, Indiana. The exact site is uncer-
tain. Because the bones were unearthed by a back-
hoe during excavation of a pipeline trench, and
some were recovered from the spoil heaps, the
exact stratigraphic placement is unknown. The land
surface was modified shortly thereafter by earth-
moving equipment.

A complex record of glacial sediments is pre-
served in what is now Fulton County, Indiana,
representing the effects of three lobes of Wiscon-
sinan ice (Gray, 1989). Most of the county is blan-
keted by northeastern-source till of the Trafalgar
Formation, deposited by the East White Sublobe
(Huron-Erie Lobe). A recessional edge moraine of
Trafalgar ice (“‘“Packerton Moraine’’) transects the
southeastern corner of the county (Bleuer & Mel-
horn, 1989). Northern-source tills and moraines
of the Michigan Lobe terminate in northwestern
Indiana to the west of Fulton County, but a com-
plex series of associated alluvial fans and fan heads
(‘““Maxinkuckee Moraine’’) enter the northwestern
part of Fulton County. Both the ““Maxinkuckee”
and ‘“‘Packerton” features were previously be-
lieved to have been terminal moraines of the
northeastern-source Saginaw interlobe (Wayne &
Zumberge, 1965; Schneider & Johnson, 1967),
which is now believed to have terminated much
farther to the north in Indiana (Bleuer & Melhorn,
1989). The primary advance of the Trafalgar ice
margin occurred at approximately 20,000 B.P., and
a recent date suggests that ice was present at the
present site of Plainfield, Hendricks County (about
150 km south of Fulton County), at 17,650 + 190
B.P. (N. Bleuer, pers. comm. to Richards). A later
advance of Lagro ice (Miami Sublobe, Huron-Erie
Lobe) to the east of Fulton County deposited a
classical series of curved recessional moraines in
northeastern Indiana (Gray, 1989; Mickelson et
al., 1983). The earliest Lagro advance occurred at
approximately 15,500 B.p. (after the “Erie Inter-
stade’’), with ice receding to near the eastern bor-
der of Indiana (Fort Wayne Moraine) by approx-
imately 14,800 B.p. (Mickelson et al., 1983).

The initiation of the Lagro sequence ca. 15,500
B.P. indicates the period when Fulton County was
deglaciated of Wisconsinan ice. Deglaciation left
numerous kettle depressions across the landscape,
many of which held water. Deposition of calcar-
eous clay (marl) and peat are typical in “kettle
lakes” of northern Indiana (Wayne, 1971; Schnei-
der & Moore, 1978). Sediments and the occasional

remains of mastodonts or other vertebrates de-
posited in these kettles typically date from the past
13,000 years (Wayne, 1963; Wayne & Zumberge,
1965).

Calcareous clays containing aquatic snails (in-
cluding Gyraulus, Valvata, and Heliosoma /Pla-
norbella) adhering to some of the bones and filling
their cavities indicate that the Fulton County Arc-
todus remains were deposited in a shallow kettle
lake. The period of deposition is indicated by a
radiocarbon date of 11,500 + 520 B.p. (Geochron
Laboratories, GX-12483; C-13 corrected) on Arc-
todus bone apatite (a portion of the left #7 rib).

Description

Genus Arctodus Leidy, 1854

Arctodus simus (Cope, 1879)

Arctodus simus yukonensis (Lambe, 1911);
Kurten, 1967

The skeleton is clearly assignable to Arctodus
simus using the criteria of Merriam and Stock
(1925) and Kurtén (1967). The remains are illus-
trated in Figures 1-16, and measurements are pre-
sented in Tables 1-7. These tables include mea-
surements of some elements (i.e., atlas, axis,
sacrum, and scapula) that have previously been
measured only in the smaller-sized Potter Creek,
California, Arctodus simus population (Merriam
& Stock, 1925) or that were entirely lacking in the
literature (e.g., pelvis, caudal #1, ribs). Kurtén
(1967) presented scant size information on these
elements for the larger A. s. yukonensis.

Most North American Arctodus simus occur as
isolated individuals, but Rancholabrean-aged
population samples are available for Potter Creek
Cave and Rancho La Brea, California (Merriam
& Stock, 1925), and from the Fairbanks, Alaska,
area (Kurtén, 1967). Table 8 presents comparisons
of the Indiana specimen with those population
samples using data from Merriam and Stock (1925)
and Kurtén (1967) and additional information from
Richards, Churcher, and Turnbull (in press). The
Potter Creek population is composed predomi-
nantly of females (Kurtén, 1967), the Rancho La
Brea population may be composed largely of fe-
males and adolescent males (Agenbroad & Mead,
1986), and the Fairbanks material appears to in-
clude both sexes. Although there are some robust
skeletal elements from Rancho La Brea, the Fair-

FIELDIANA: GEOLOGY

ERE
Fic. 1. A. Skull, dorsal view. B. Skull with dentaries articulated, rostral view. C. Skull, left lateral view. Scale in cm.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 34

Fic. 2. A. Upper incisors, occlusal view. B. Upper right premolars, occlusal view (scale to right). C. Upper left
tooth row, occlusal view. D. Skull, palatal view. Scale in cm.

banks material assigned by Kurtén (1967) to the
subspecies A. s. yukonensis appears to be of larger
size than either of the California samples. The
large size of the Indiana remains compares closely
to that of the Fairbanks, Alaska, material. Arcto-
dus simus teeth, however, display a great range in
size and do not segregate readily into the two sub-
species that are indicated by limb bone length.

Teeth of the Indiana specimen, with the exception
of the large canines, fall within the middle range
of size. Figures 17-20 show size relationships of
selected elements of PM #24880 within a broader
sample of North American Arctodus simus, in-
cluding large specimens of Irvingtonian age. (An
expanded variety of elements and measurements
is presented in Richards et al. [in press].) The large

FIELDIANA: GEOLOGY

D

Fic. 3. A. Right dentary, lingual view. B. Right dentary, occlusal view of tooth row. C. Right dentary, labial view.

D. Left dentary, occlusal view of tooth row. Scale in cm.

size of the Indiana remains compares closely to
that of Irvingtonian-aged specimens from the cen-
tral and northern Great Plains and to that of Ran-
cholabrean-aged specimens from Alaska and the
Yukon, while tooth dimensions appear not to be
diagnostic. Differentiation between the two sub-
species rests on the length of the skull, jaws, and

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA

limb bones. Based on the long limb bones, the
Indiana skeleton is referred to the subspecies A. s.
yukonensis.

The excellently preserved, medium brown, hard
bones of the specimen include the nearly complete
but broken skull with jaws and most of the major
skeletal elements. Some elements show patholog-

Fic. 4. A. Atlas, dorsal view. B. Atlas, posterior view. C. Atlas, ventral view. D. Axis, dorsal view. E. Axis, left
lateral view. F. Axis, anterior view. G. Fourth cervical vertebra, anterior view. H. Fourth cervical vertebra, posterior
view. Scale in cm.

6 FIELDIANA: GEOLOGY

Fic. 5. A. Sixth cervical vertebra, anterior view. B. Sixth cervical vertebra, left lateral view (scale below). C. Sixth
cervical vertebra, posterior view. D. Third thoracic vertebra, anterior view. E. Third thoracic vertebra, posterior
view. F. Fourth thoracic vertebra, anterior view. G. Fourth thoracic vertebra, posterior view. H. Fifth thoracic
vertebra, anterior view. I. Fifth thoracic vertebra, posterior view. Scale in cm.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA i

Fic. 6. A. Tenth thoracic vertebra, anterior view. B. Tenth thoracic vertebra, left lateral view (scale to left). C.
Tenth thoracic vertebra, posterior view. D. Eleventh thoracic vertebra, anterior view. E. Eleventh thoracic vertebra,
posterior view. F. Thirteenth thoracic vertebra, anterior view. G. Thirteenth thoracic vertebra, left lateral view (scale
to left). H. Thirteenth thoracic vertebra, posterior view. I. Second lumbar vertebra, anterior view. J. Second lumbar
vertebra, left lateral view (scale to left). K. Second lumbar vertebra, posterior view. Scale in cm.

FIELDIANA: GEOLOGY

H |

Fic. 7. A. Fourth lumbar vertebra, anterior view. B. Fourth lumbar vertebra, left lateral view (scale to left). C.
Fourth lumbar vertebra, posterior view. D. Fifth lumbar vertebra, anterior view. E. Fifth lumbar vertebra, posterior
view. F. Sixth lumbar vertebra, anterior view. G. Sixth lumbar vertebra, posterior view. H. Seventh lumbar vertebra,
anterior view. I. Seventh lumbar vertebra, posterior view. Scale in cm.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 9

Fic. 8. Pathologic detail of thoracic vertebrae. A. Fourth thoracic vertebra, anterior centrum face pathology. B.
Fifth, fourth, and third thoracic vertebrae articulated, right lateral view. C. Fifth thoracic vertebra, posterior centrum

face pathology. Scale in cm.

ical conditions of varying severity, which we de-
scribe here. Rothschild and Turnbull (1987) re-
ported a positive treponemal reaction in some of
the thoracic vertebrae, and Neiburger and Turn-

10

bull (1990, and unpublished) made a differential
diagnosis of all of these pathologic conditions. The
ulnar pathologies are discussed by Neiburger in
the Appendix. The fragmented skull, scapulae, pel-

FIELDIANA: GEOLOGY

13

12
1

A

VERE aia a RIGHT
Fic. 9. Ribs, numbered in position, posterior view. Scale in cm.
vis, tibia, and ribs, among others, without sedi- of inexperience on the part of the collector, who

ment impacted into the bone interiors, appear to nevertheless did well, inasmuch as this specimen
have been broken either upon their discovery or is one of the more nearly complete and best pre-
during subsequent excavation. Missing elements, served skeletons of Arctodus in existence.

or broken-off parts of bones, were lost as a result The bones lack rodent gnawing and other ob-

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 11

Fic. 10. A. Left scapula, lateral view. B. Left scapula, medial view. C. Right scapula, lateral view. D. Right

scapula, medial view. Scale in cm.

vious indications of predepositional disturbance.
It is apparent that one individual is represented,
that the bones were in relatively close association
(most probably in articulation) when discovered,
and that the skeleton must have undergone a rel-

12

atively rapid burial. Considering the hand-col-
lected method of recovery, without screening or
washing of the sediments or following of anatom-
ical relationships to ensure completeness of wrist,
ankle, and foot bones, the lack of small elements

FIELDIANA: GEOLOGY

A. Left humerus, anterior view. B. Left humerus, posterior view. C. Right humerus, anterior view (note

shaft pathology). D. Right humerus, posterior view (note shaft pathology). E. Left humerus, superior view. F. Left
humerus, inferior view. G. Right humerus, anterior view of pathology. H. Right humerus, medial view of pathology.

RiGiht:

Scale in cm.
RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA

13

Fic. 12. A. Left radius, anterior view. B. Left radius, posterior view. C. Right radius, anterior view. D. Right
radius, posterior view. E. Left ulna, right lateral view. F. Left ulna, left lateral view. G. Right ulna, left lateral view.
H. Right ulna, right lateral view. I. Left ulna, proximal end, anterior view. J. Left metacarpal IV, right lateral view.
Scale in cm.

14 FIELDIANA: GEOLOGY

Fic. 13. A. Pelvis, superior view. B. Pelvis, inferior view. C. Pelvis, right lateral view. Scale in cm.

or associated fauna (except molluscs) is not sur- skull lacks a section of the left parietal wall, right
prising. The bones have been consolidated with sphenoid, and most of that portion from the vo-
glyptal, carried by acetone. mer area forward, including the nasal turbinates,

most of the palatines, the posterior part of the right
maxilla, and the anterior section of the right zy-
gomatic arch. The skull and its dentition represent

The skull, broken by the backhoe and recovered an early middle-aged adult. Externally, sutures are
as front and rear halves, is nearly complete. The closed, most with some “bridging” of the ele-

Skull, Dentaries, and Teeth

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 15

Fic. 14. A. Right femur, posterior view. B. Right femur, anterior view. C. Right femur, superior view. D. Right
femur, inferior view. E. Left tibia, anterior view. F. Right fibula shaft. G. Right tibia, anterior view. H. Right tibia,
right lateral view. I. Right metatarsal V, right lateral view. J. Left ectocuneiform. K. Right calcaneum, left lateral
view. L. Right calcaneum, superior view. Scale in cm.

16 FIELDIANA: GEOLOGY

Fic. 15. A. Right ulna, left lateral view of pathology. B. Left ulna, posterior view of pathology. Scale in cm.

ments, and some sutures are nearly obliterated
(e.g., Squamosoparietal and anterior maxillopre-
maxillary). The sagittal crest rises to a maximum
of 50 mm above the braincase. Features typical of
the species include the short, wide snout (including
nasal), the large narial opening, the high-vaulted
skull, three anterior palatine foramina, and double
infraorbital and large parietal foramina (Merriam
& Stock, 1925). The frontals are somewhat inflat-
ed.

Both dentaries are complete. The premasseteric
fossae, typical of the Tremarctinae, are well de-
veloped, as are multiple mental foramina. The large
mental foramen is below p3 on the right dentary
and below the p3—p4 junction on the left. The right
dentary has four and the left dentary three small

accessory foramina. The angles project posteriorly
beyond the condyles.

The dental formula for Arctodus is that of the
Ursidae: +++. The P4 metacone forms somewhat
of a shearing blade, as does the m1 trigonid (Mer-
riam & Stock, 1925). The P4 protocone has a rel-
atively anterior position (Kurtén, 1967). Several
teeth had been shed while the bear was alive, the
alveoli closing or containing bony trabeculae: LI1
(trabeculae); LP1 (alveolus closed); L,Ril (?tra-
beculae), and Lp4 (alveolus closed) (loss of ante-
rior premolars is a usual condition in bears). Sev-
eral teeth have been lost from their alveoli post
mortem: RP2; RP3; L,Ri2; L,Ri3; L,Rp1; Lp2;
and L,Rp3. Crowns fractured from the roots (pre-
sumably by the backhoe) and lost post mortem

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA ae

A El

‘ B

Fic. 16. Right rib, showing measurements for data compiled in Table 4 (after Kurtén, 1966). Scale in cm.

include RC; RP4; RM1; and RM2. Tooth wear
ranges from slight or moderate on the molars to
moderate on the carnassials and incisors, where
the dentine is exposed.

The upper premolars are relatively crowded.
Premolar spacing in the left maxilla is as follows:
LC alveolus-LP2, 11.49 mm (LP1 shed); and LP2,
LP3, and LP4 are in contact. LP2 is twisted
obliquely inward anteriorly, its long axis being
turned inward 55°. The posterior end of LP3 is
located slightly medial to the anterior end of LP4.
Premolar spacing in the right maxilla is as follows:
RC-RP1 alveoli, adjacent; RC-RP2 alveoli, 11.15
mm; RP2-RP3 alveoli, 3.44 mm; and RP3 alve-
olus adjacent to the anterior RP4 root and alve-
olus. RP1 is the largest of the first three anterior
premolars, and RP2 and RP3 appear to be slightly
larger (by alveoli) and less crowded than those of
the left series.

The lower premolars are not as crowded. Pre-
molar spacing in the left dentary is as follows: Lc-
Lp! alveoli, 2.3 mm; Lp1-Lp2 alveoli, contacting;
Lp2-Lp3 alveoli, 7.63 mm; and Lp3-Lm1 alveoli,
13.68 mm (Lp4 shed, no alveolus). Premolar spac-
ing in the right dentary is as follows: Rc-Rp1 al-
veoli, 3.3 mm; Rp1-Rp2 alveoli, 2.08 mm; Rp2-
Rp3 alveoli, 9.16 mm; and Rp3-Rp4 alveoli, 4.33
mm, with Rp4 contacting Rm1. The anterior end
of Rp2 is twisted slightly inward. Thus, the longest
lower tooth row diastemata are between p2 and
p3, a condition noted in the Frankstown Cave
(Peterson, 1926) and Hay Springs (Kurtén, 1967)
materials.

18

Vertebrae

Recovered vertebrae include the atlas, axis, cer-
vical vertebrae C-4 and C-6, thoracic vertebrae
T-3, T-4, T-5, T-10, T-11, and T-13, lumbar ver-
tebrae L-2, L-4, L-5, L-6, and L-7, and five fused
sacral vertebrae with a sacralized caudal #1. The
preserved vertebrae posterior to the atlas and axis
were determined as to position by comparison with
one modern skeleton each of Tremarctos ornatus
and Ursus americanus (14 thoracic vertebrae; Kur-
tén, 1966) and based on a thoracic count of 13 in
Arctodus (Merriam & Stock, 1925; Kurtén, 1967).
The recovered vertebrae are nearly complete. T-3,
L-2, L-4, L-6, and L-7 each lack the tip of the
neural spines. Transverse processes are fragment-
ed or missing on T-5 (R), L-2 (L,R), L-5 (L,R),
L-6 (L), and L-7 (R). T-3, T-4, and T-5 display
extensive osteophytosis (discussed later). Mea-
surements of all preserved vertebrae are presented
in Table 3. Few vertebral measurements are avail-
able from the literature (Kurtén, 1967).

Ribs

A relatively complete set of ribs is present. Based
on 13 pairs of ribs, allotment is the following: R#1,
L#2, L,R#3, L#4, L,R,#6, L,R#7, L#9, L,R#10,
L,R#11, R#12, and L,R#13, and a distal rib sec-
tion. Lacking distal sections are the following: L#2,
L,R#3, L,R#6, L#7, L#9, L#10, and L#13. Heads

FIELDIANA: GEOLOGY

of several anterior left ribs have minor patholog-
ical exostoses.

Kurtén (1966) compared several of the ribs of
Tremarctos ornatus, T. floridanus, and Ursus
americanus, but those of Arctodus simus have not
been described. The ribs are not morphologically
different from those of other ursids but are distinct
in size and proportions. Comparison of measure-
ments in Table 4 to those of Kurtén (1966) for
ribs #1, #4, #9, #11, and #12 show those of A.
simus to be considerably longer than those of the
above species, with all of the A. simus ribs from
at least #4 through #13 exceeding all ribs of the
above species in length. Though rib #9 is incom-
plete, #10 appears to be the longest rib in A. simus,
as may be the case with the above species. The
ribs of A. simus also appear to be distinct from
those of the above species in their relatively small
amount of lateral “bow” (radius; Kurtén, 1966),
falling equal to or less than those of 7. floridanus
and one specimen of U. americanus in absolute
values. Arctodus simus is also unusual in that the
radii of the posterior rib series (#12 and #13) are
less than that of #4; in the above bears, the radii
of the posterior rib series (including #9 through
#12) are greater than that of #4. Rib proportions
thus suggest a deep, relatively narrow chest in Arc-
todus simus.

Forelimb and Girdle

Forelimb elements include both scapulae, hu-
meri, ulnae, radii, and L metacarpal IV. The right
scapula is nearly complete, lacking the anterior
midsection of the blade. The calcified suprascap-
ular cartilage of the R scapula is unfused to the
blade on its inner surface. The left scapula lacks
much of the anterior and posterior blade margins
and the spine above the coracoid process. The
limb bones, especially the ulnae and radii, are rel-
atively long and slender. The ulnae, radii, and hu-
meri are complete, the latter bearing the entepi-
condylar foramina characteristic of the
Tremarctinae (Merriam & Stock, 1925). The R
humerus, both ulnae, and the L radius display
pathology. The L metacarpal IV lacks the distal
end.

Hind Limb and Girdle

Hind limb elements include the pelvis, R femur,
L, R tibiae, R fibula shaft, R calcaneum, L ecto-

TABLE 1. Arctodus simus (PM #24880)—Cranial and
dentary measurements (mm).

Skull
Basal length of skull 396
Condylobasal length of skull 422
Extreme length of skull 453
Zygomatic width 319
Rostral width at canines 126.4
Interorbital width 135.3
Width over postorbital processes 170.7
Width over postorbital constriction (esti-

mate) 94
Postorbital height (top of frontal to posterior

inferior rami of palatines) 174
Width of nasal opening (inner rim of pre-

maxillae) 86.4
Front of premaxillae to hinder ends of M2 174.5

Bottom of glenoid fossa to junction of frontal
and occipital sutures (height of cranium) 207

Braincase width (estimate) 141
Width across occipital condyles 88.8
Occipital height (base of condyles to top of

occipital crest) 181.6
Mastoid breadth (estimate) 218
Width across base of incisors 68.6
Height of foramen magnum 30:5
Width of foramen magnum 35
Dentaries Left Right
Length, anterior edge of canine to rear

of condyle 299 297.3
Length, tip of angular process to ante-

rior end of symphysis 324 325
Depth of coronoid process (from bot-

tom of angle) 153 155
Depth at diastema 64.1 63.7
Ramus depth between m1 and m2

(lateral face) 64.7 68.2
Ramus depth behind m3 86.4 89.9
Length, p4—m3 _ 94.9

cuneiform, and R metatarsal V. Much of the pelvis
is present, the right innominate (in two parts) lack-
ing the pubic symphysis area; the left is repre-
sented by the ischium only (in two parts) con-
tacting a small scrap of the ilium. The slightly
fragmented sacrum is co-ossified with both in-
nominates (suture obliterated), unifying the pelvis,
which was fragmented into more than five parts
by the backhoe.

The pelvis was compared with descriptions of
materials of Tremarctos (Kurtén, 1966) and of
Arctodus simus (Merriam & Stock, 1925) and with
FMNH specimens 123369 (7remarctos ornatus),
63803 (Ursus arctos gyas), and 27268 (Ursus arc-
tos). The ilium of Arctodus simus is relatively short

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 19

TABLE 2. Arctodus simus (PM #24880)— Dental measurements (mm).

Upper teeth Left Right
C

Width at base of enamel 23.03 _

Length at base of enamel 32:75 —

Width at alveolar border 25.42 _

Length at alveolar border 39.04 —
Pl

Length — 10.24

Width — 7.34
2

Length 7.61 —

Width 4.73 _
P3

Length 8.92 -

Width 5.44 _
P4

Length 23.47 --

Width 16.8 —

Paracone height 9.8 _
Ml

Length 26.06

Anterior width 24.04 —_

Posterior width 25.64 —

Paracone height 10.02 _

Metacone height 8.37 —
M2

Length 36.52 -

Anterior width 24.50 _

Central width 21.68 _

Paracone height 9.28 a
Il, transverse diameter — 8.88
12, transverse diameter 9.56 9.79
13, transverse diameter 14.13 14.8
Length C-M2 150.9 os
Length, posterior edge of C alveolus to an-

terior edge of M1 48.2 _
Length, M1-M2 61.4
Length, posterior edge of C to anterior edge

of P4 27.6 —
Lower teeth* Left Right
(s

Width at enamel base 24.22 24.14

Length at enamel base 32.9 34.1 (e)

Width at level of alveolus 24.39 24.82

Length at level of alveolus 41.67 37.82
pl

Alveolar length a 6.5

Alveolar width 5.6 6.8
p2

Length 8.1 (alv.) 9.04

Width 6.2 (alv.) 5.94

6.0 (alv.)

p3

Alveolar length 8.1 5.6

20

FIELDIANA: GEOLOGY

TABLE 2. Continued.

Lower teeth (cont.) Left Right
Alveolar width 4.5 4.6
p4
Length — 12.59
Width _ 7.0
ml
Length 33.1 33.92
Anterior width 16.1 17.0
Posterior width 16.92 17.04
Trigonid length 23.03 23.69
Protoconid height
Labial 15312 15.18
Lingual 14.33 13.95
Hypoconid height 11.8 ca. 10.14 (f)
m2
Length 30.35 31.8
Anterior width 21.14 21.48
Posterior width 19.75 19.63
Protoconid height 12:57 ca. 10.58 (f)
Hypoconid height 9.21 10.16
Metaconid height 9.62 11.16
m3
Length 22°31 22.19
Width 16.55 16.50
il, transverse alveolar diameter 7.9 est. 8.5 est.
i2, transverse alveolar diameter — 7.4 est.
i3, transverse alveolar diameter 10.1 9.5 est.
Length, c-m3 170.8 L717
Length, posterior edge of c alveolus to an-
terior edge of m1 44.4 48.8
Length, ml—m3 83.4 82.8

*e, enamel limit difficult to determine; f, cone fractured or worn; alv., alveolar measurement; est., estimate.

(Merriam & Stock, 1925), and the more complete
Rochester specimen displays ilia that flare antero-
laterally more than those of the three FMNH spec-
imens, distinctively producing a relatively large
width across the outer iliac blades for the size of
the pelvis. The neck of the ilium immediately in
front of the acetabulum is massive, and relatively
more so than in the FMNH specimens. The acetab-
ulum is very large, but not relatively larger than
in Ursus. The Rochester specimen displays a more
pronounced development of the ventral shelf run-
ning from the lower border of the acetabulum to
the obturator foramen than do the FMNH speci-
mens, a condition noted to be absent in 7remarc-
tos (Kurtén, 1966). The ischial bar above the ob-
turator foramen is similar to that of Ursus and not
compressed from the sides as in Tremarctos (Kur-
tén, 1966). Unlike a pubis of A. simus which was
relatively narrower than that of Ursus or Tremarc-
tos (Merriam & Stock, 1925), the Rochester pubis

is proportioned similar to that of Ursus.

The interpretation of five sacral vertebrae with
a sacralized caudal #1 might alternatively be in-
terpreted as four true sacrals with one totally fused
caudal #1 and an additional sacralized caudal #2.
By comparison, two FMNH specimens displayed
four true sacrals and a sacralized caudal #1 (Trem-
arctos ornatus, 123369; Ursus arctos gyas, 63803),
and another specimen was composed of five true
sacrals (Ursus arctos, 27268).

The right femur and right tibia are complete.
The left tibia is present as proximal and distal
ends, lacking the middle of the shaft, and the fibula
lacks both proximal and distal ends. The calca-
neum, ectocuneiform, and R metatarsal V are
complete. The hind limb bones, especially the fe-
mur, are relatively long and slender.

Epiphyses on all postcranial elements are closed,
with the lines of union obliterated, indicating full
adult stature.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 21

TABLE 3. Arctodus simus (PM #24880)— Vertebral measurements (mm).

Atlas
Greatest width across lateral process 243
Greatest width across anterior articulating facets 97
Greatest width across posterior articulating facets 91
Least anteroposterior diameter of dorsal arch 44
Anteroposterior diameter of ventral arch at midline 38
Greatest anteroposterior length (including lateral processes) 12
Greatest depth 64
Least width between atlantic foramina 64
Length from anterior end of articulation for condyles to posterior end of articulation for axis 89
Axis
Greatest ventral length, including odontoid process 104
Ventral length of odontoid process from transverse ridge 39
Greatest width across anterior articulating facets 86
Greatest width across postzygapophyses 78
Greatest transverse width across posterior epiphysis of centrum 58
Depth of centrum at midline of posterior surface 39
Least anteroposterior diameter of pedicle of neural arch 53
Anterior width of odontoid process (at base) 24.5
Extreme posterior depth of dorsal edge of neural spine 121
Anteroposterior length of neural spine 121
C-4

Cervical vertebrae (or 5) C-6
Ventral length of centrum from midpoint of anterior surface 53 53
Length of centrum measured normal to posterior face and along median line 48 46
Greatest width of posterior end of centrum 61 61
Depth of centrum at midline of posterior surface 43 43
Depth of centrum measured normal to floor of neural canal and across posterior epiphysis 42 42
Length from end of anterior zygapophysis to end of posterior zygapophysis 83 88
Width across anterior zygapophyses (not the supporting bosses) 94 98
Width across posterior zygapophyses 95 96
Greatest width of neural canal at anterior end ca. 33 ca. 42
Greatest width across outer ends of transverse processes 182 --
Height from middle of ventral border of posterior epiphysis of centrum to top of neural

spine 117 120
Thoracic vertebrae T-3 T-4 T-5 T-10 T-11 T-13
Ventral length of centrum

from midpoint of anteri-

or surface 43 ca. 40* 39 47 49 56
Greatest width of posterior

end of centrum 87* 86* 82 91 85 87
Depth of centrum at mid-

line of posterior surface 48 48 49 59 56 58
Depth of centrum mea-

sured normal to floor of

neural canal and along

median line of posterior

epiphysis ca. 48 ca. 48 49 533 56 By)
Length of centrum mea-

sured normal to posterior

face and along median

line 43* ca. 41* 38* 47 49 Si
Length from end of anterior

zygapophysis to end of

posterior zygapophysis iS cavi2 76 84 84 99

pes

FIELDIANA: GEOLOGY

TABLE 3. Continued.

Thoracic vertebrae (cont.) T-3 T-4 T-5 T-10 T-11 T-13
Greatest width across ante- (facets)

rior zygapophyses 59 Di) 53 56 54 80
Width across posterior zyg-

apophyses 59 56 61 55 78 62
Greatest width, across outer

ends of transverse pro- (metapophyses)

cesses 150 143 _ 141 135 LS
Greatest length from end of metapophyses to end of anapophyses

Left 88

Right 85
Greatest anteroposterior diameter of outer end of transverse process (vertical diagonal)

Left - 40 _ 45 51 86

Right 43 43 40 44 56 88
Height of neural canal at

anterior end 28 29 30 33 3) 32
Height from middle of ven-

tral border of posterior

epiphysis of centrum to

top of neural spine (cen- — 207 207 203 189 195

trum vertically) 229* 21'5*
Depth of centrum mea-

sured normal to floor of

neural canal and along

median line of posterior

epiphysis (including pa-

thology) 54* 66* 61* — ~ —
Lumbar vertebrae L-2 L-4 L-5 L-6 L-7
Ventral length of centrum from midpoint of anterior

surface 58 59 60 61 59
Length of centrum measured normal to face of pos-

terior epiphysis and along median line 60 63 62 62 59
Greatest width of posterior end of centrum 98* 95 100 107% 101*
Depth of centrum at midline of posterior surface 61 63 62 59 55
Depth of centrum measured normal to floor of neu-

ral canal and across posterior epiphysis 61 63 64 61 56
Greatest length from anterior end of metapophyses

to end of posterior zygapophyses 104 100 98 100 96
Greatest width across metapophyses 120 119 122 131 139
Greatest width across posterior zygapophyses 67 81 86 89 106
Width across transverse processes _ 240 — — —
Height from middle ventral border of anterior epiph-

ysis of centrum to top of neural spine —_ — 218 aaa _

* Measurements include or influenced by pathology.

Pathology

Pathological lesions occur in the lower jaws, the
vertebral column and associated ribs, and the bones
of both forelegs. Minor abscesses are present be-
tween m1 and m2 of both dentaries. The left den-
tary has some bone resorption and small drainage
channels for the abscess. Abscessing was less de-
veloped in the right dentary, with only minor re-
sorption of the alveolar edge.

An irregularly shaped area approximately 65 mm
long and 46 mm high on the labial surface of the
right dentary (below p4, m1, and the anterior '2
of m2) is penetrated by approximately 40 to 50
minute pits ranging from 0.5 to 1.0 mm in di-
ameter and is likely the result of disease. One ac-
cessory mental foramen (below the anterior root
of p4) appears to have been invaded with this
diseased condition, resulting in somewhat of an
hour-glass-shaped foramen.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 23

TABLE 4. Arctodus simus (PM #24880)—Rib measurements (mm).

Ribst
Rib No. A B Cc D E
R#1 171 170 7A 64 50
L#2 - _ _ 66 —
L#3 — —- = 68 -
R#3 _ _ frag. 67+
L#4 440 455 167 13" frag. 24+
L#6 — —_ _ 76 ~_
R#6 - — _ Td —
L#7 — - — — —
R#7 493 503 182 80 31
L#9 — _ _ 79 _
L#10 505 (+?) 512 (+7) 181 (+?) 78 _
R#10 501 508 185 78 _
L#11 500 503 178 74 31
R411 495 497 186 75 33
R#12 467 471 166 59 30
L#13 a _ = 48 —
R#13 399 399 135 46 41
? Left distal end 33

* Measurements include or influenced by pathology.

+ Rib measurements (see also Fig. 16 for illustration): A, chord length (to head); B, chord length (to tubercle); C,
radius; D, length from external edge of tubercle to end of head; E, greatest width, distal end (parallel to outer surface).

There is extensive erosion on the centra faces
of thoracic vertebrae T-3, T-4, and T-5 (especially
the posterior face of centrum T-3). These same
vertebrae exhibit spondylosis deformans (osteo-
phytosis; Morse 1978), but eburnation is not ap-
parent on the involved centra. Bony proliferation
had begun to bridge the three centra on ventral
and ventrolateral surfaces, and compression of the
vertebrae initiated reactive bone growth between
adjacent centra faces. The inferior edges of T-3
and T-4 were closely appressed, the posteroinfer-
ior margin of the centrum of T-3 fitting into an
eroded pit in the centrum of T-4, resulting in a
slight upward hunch between the vertebrae, sim-
ilar to Pott’s disease in humans (tuberculosis spon-
dylitis). Centra of T-4 and T-5 were also closely
appressed, with the prezygapophyses of T-5 drop-
ping ca. 3 mm downward, slightly constricting the
neural canal from above. The greatest reactive bone
buildup was on the ventral surfaces of the centra:
T-3, 9.8 mm; T-4, 17 mm; and T-5, 13.4 mm. On
the ventral and ventrolateral surfaces of the centra,
reactive bone encircled the rib heads, with most
of the rib facets (except for R#3) slightly eroded.
Loss of the supportive function of the centra ap-
parently allowed sufficient slippage to have pre-
cipitated such bone reaction. Rothschild and
Turnbull (1987) suggested that these and other
lesions of the skeleton resulted from infection of
the animal with syphilis or yaws, and Neiburger

24

and Turnbull (1990) have completed a differential
diagnosis that suggests tuberculosis, osteomyelitis,
arthritis, or some mycotic (fungal) infection, either
singly or in combination, as additional possible
causes.

The heads of ribs L#2, L#3, and L#4 have mod-
erate exostoses, and there is slight development
on the heads of L,R#6.

The skeleton shows little osteoarthritis, though
minor “‘lipping” is present on the centra of the
atlas, axis, C-4, C-6, T-10, T-11, T-13, L-2, L-4,
L-7, and the sacrum and on articular facet margins
of the radii and ulnae. All “‘lipping”’ appears to be
the result of ontogenetic stress on the joints of this
early middle-aged adult bear.

The R humerus and both ulnae display lesions
and possibly traumatic injury. A bony flap 190
mm long (46 mm wide at the middle) projects
medially from the anteromedial edge of the shaft
ca. 260 mm from the distal end of the R humerus.
It terminates proximally in a bony spur and dis-
tally in a muscle scar. Arguably this could be a
healed fracture as originally thought. However, ra-
diographic evaluation suggests an unusual form of
periosteal reaction rather than fracture. The shaft
is predominantly swollen in the anteroposterior
plane at the distal end of the pathology. Interest-
ingly, shaft alignment and total length are identical
to that of the L humerus.

Both ulnar shafts have abscesses. The abscess

FIELDIANA: GEOLOGY

of the L ulna (15.9 mm x 6.5 mm, and 12.7 mm
deep) orients with the long axis of the shaft and
expands in size below the smooth-edged orifice. It
centers 253 mm from the distal end of the shaft.
The abscess area of the shaft is swollen over 10
mm greater than that of the R ulna. Some reactive
bone formation is present in the abscess area. The
abscess on the posteromedial margin of the R ulna
is 11.7 mm long aligned with the shaft axis, 5.6
mm wide, and 10 mm deep. The orifice edges are
rounded, and there are osteophytes and small bony
spurs around the abscess area. The abscess centers
116 mm from the distal end of the shaft. The shaft
in the abscess area is swollen in the anteroposterior
(somewhat diagonal) plane 6 mm more than that
of the L ulna. Both abscesses appear to have a
common origin. Rothschild and Turnbull (1987)
suggested that these and other skeletal lesions orig-
inated as a pathogenic (treponemal) infection
(Benditt, 1989). Neiburger (1984, 1988, and the
Appendix) does not accept that diagnosis and fa-
vors origin by an infected wound, and Neiburger
and Turnbull (1990) give a full differential diag-
nosis that implicates osteoarthritis and tubercu-
losis, in addition to the contracted syphilis with
long-term infection. The convex side of the L ra-
dius shaft is rough and thickened just below the
level matching the abscess on the L ulna. Perhaps
it represents an adjustment to the weakening of
the L ulna.

Discussion and Conclusions

The Indiana specimen is the only A. s. yuko-
nensis fossil known east of the Mississippi River.
The Frankstown Cave, Pennsylvania (Peterson,
1926), and at least one of the two Kentucky finds
(Proctor Cave; Wilson, 1981) represent the smaller
A. s. simus. The Sheriden Pit, Ohio, specimen has
the small size of simus females from Rancho La
Brea (McDonald, pers. comm. to Richards, 6 Jan.
1992). Both subspecies may have co-inhabited (and
likely interbred in) the lower Great Lakes region
for a short time in the very late Pleistocene.

Kurtén (1955) related that the canines showed
the strongest sexual dimorphism in the dentition
(and skeleton) of Ursus spelaeus (cave bear). As-
suming like conditions for Arctodus, Kurtén (1967)
suggested that Arctodus simus canines wider than
19.5 mm [tended to] represent presumed male in-
dividuals. The Indiana canine teeth range from
23.0 (upper) to 24.2 (lower), indicating a male bear
by this criterion.

TABLE 5. Arctodus simus (PM #24880)—Forelimb

and girdle measurements (mm).

Measurement Left Right
Scapulae
Greatest length, from coracoid pro-

cess to top of scapula, measured

along spine axis 462+ 478
Distance from inner border of glenoid

cavity to top of spine 409 409
Length, from inferior rim of glenoid

articulation to dorsal border,

along axis of spine 431 436
Greatest anteroposterior width of fac-

et of glenoid cavity 96 97
Greatest transverse diameter of gle-

noid cavity 70 13
Least anteroposterior width of neck at

articulating end 125 126
Least thickness of neck at articulating

end 46 46
Transverse width of glenoid articula-

tion to outer side of acromion

process 141 152
Greatest width taken normal to the

anterior edge of the spine to the

greatest extension of the posterior

blade part of the scapula —_ 225
Humeri
Total length 594 594
Greatest proximal width 126 128
Greatest proximal anteroposterior di-

ameter 145 146
Least width of shaft (in plane of axis

of distal end) 55 55
True least width of shaft 51 50
Minimum midshaft circumference 196 —
Greatest width across epicondyles 160 161
Greatest transverse width of distal ar-

ticulation surfaces 125 WAT
Greatest transverse width of distal ar-

ticulation (facets only) 116 119
Ulnae
Greatest length 542 545
Proximal diameter from tip of coro-

noid process to Margo dorsalis 108 is
Smallest diameter from bottom of

semilunar notch to Margo dorsalis 61 64
Inner diameter of semilunar notch 49 48
Least transverse diameter of shaft

above capitulum 36 34
Radii
Greatest length 481 479
Proximal end, long diameter 69 68
Proximal end, short diameter 56 56
Middle of shaft, long diameter 44 43
Middle of shaft, short diameter 34* 30
Distal end, long diameter 92 95
Distal end, short diameter 60 61

* Measurement includes or influenced by pathology.

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA

pas

TABLE 6. Arctodus simus (PM #24880)—Hind limb

and pelvis measurements.

TABLE 6. Continued.

Right fibula
Left Right
innom-_ innom- Least diameter in distal half of
Pelvis inate inate shaft 10.5
Greatest length _ 516
Length of ilium from anterior
margin of acetabulum _ 260 ,
Greatest length of ischium, from Comparison of the measurements of the largely
posterior border of acetabulum 170 169 disassociated elements plotted in Figures 17 and
sguceenes end of ilial blade, long on 20 suggests that the Indiana specimen has a rel-
iameter - : : :
Least depth oF ilial neck *s 83 atively small skull in relation to the length of the
Racat thickness of iial neck ea, 67 long bones. The few skulls with associated long
Least depth of ischium bar above bones include presumed male individuals from
obturator foramen 49 ow! Hay Springs, Nebraska (Kurtén, 1967), and Hot
Least width of same 37 37 Springs, South Dakota (Agenbroad & Mead, 1986),
perenne acctabulues tantero- 90 and a presumed female from Cueva Quebrada,
y : Texas (Lundelius, 1984) (Table 9). Comparison
tie, greatest length 466 with those specimens indicates that the Indiana
elvis, width from outer edges of : ; ; : :
ischia 234 specimen has a relatively short, wide skull that is
Pelvis, width from tubercles, ante-
rior edge of acetabula ca. 274
Pelvis, width across outer iliac TABLE7. Arctodus simus (PM #24880)—Podial mea-
blades (left ilium absent; surements (mm).
R-midline = 267 mm) ca. 534
Sacrum Right calcaneum
Central length 225 Greatest length 135
Central depth, anterior 50 Greatest width, obliquely across sustentacu-
Central width, posterior 33 lum — 81.5
Greatest width of third sacral ver- Least width of tuber calci (“neck”) 30.1
tebra across transverse processes ca. 156 Depth of tuber calci (mid-“neck”’) 57.8
Centrum length, caudal #1 ca. 34 Width of cuboid facet 47.6
Length, anterior end of sacrum to Depth of cuboid facet (difficult to align) 31.4
posterior end of caudal #1 259 Long diameter of sustentacular facet 49.5
Short diameter of same 37.8
Right femur Width of inner astragalar facet 29.6
Width of tuber calci (terminal) 53.8
Greatest length 651 Depth of tuber calci (terminal; difficult to
Greatest proximal width 174 align) 62.5
Caput diameter (anteroposterior) 719
Least transverse width of shaft 57 Left ectocuneiform
Minimum midshaft circumference 171
Greatest distal width over epicon- Greatest dorsoventral depth 56:2
dyles 145 Greatest width 33.4
Least depth of collum 67 Greatest proximodistal length (dorsal) 2123
Metatarsal facet, long diameter 40.2
Tibiae Left Right
Left Right
Greatest length _ 478 meta- meta-
Greatest proximal width — 140 carpal tarsal
Least width of shaft — 49 Metapodials IV V
Greatest distal width, measured in
plane of articulation 114 113 Greatest length a 135
Anteroposterior diameter of distal Proximal width (with distal end
end 65 64 “‘squared up’’) 29.8 45.1
Proximal anteroposterior diame-
Right fibula ter 41.8 51.8
Least transverse width of shaft 21.0 Wiz
Least diameter in proximal half of Greatest distal width over epi-
shaft 13.6 condyles _ 31.4

26

FIELDIANA: GEOLOGY

27

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BASAL LENGTH SKULL

Fic. 17. Bivariate graph of Arctodus simus—Skull length by zygomatic width of Indiana specimen compared with
others. For explanations of abbreviations, symbols, and type localities, see below.

Abbreviations and symbols: RAN, Rancholabrean; IRV,
Irvingtonian; UNK, unknown. Primary reference(s) for
measurements follows age. Boxes encase the range of
measurements from Rancho La Brea (Locality 9) and
Potter Creek Cave (Locality 12). Symbols: solid circle,
A. simus simus (RAN); hollow circle, A. simus yukonen-
sis (RAN); hollow square, A. simus yukonensis (IRV);
“winged” circle, subspecies uncertain (RAN); “‘winged”’
triangle, subspecies and age uncertain; star, Fulton Coun-
ty, Indiana (PM #24880). Symbols along graph margins
show isolated, singular measurements.

Key to type localities for Figures 17—20:

1. Gold Run Creek, Yukon Territory (RAN; Kurtén,

1967; Harington, 1977).

New Fern Cave, Jackson County, Alabama (RAN;

Ray, 1969).

Upper Cleary River Beds, Alaska (RAN; Kurtén,

1967; Richards et al., in press).

Ester Creek, Alaska (RAN; Kurtén, 1967; Richards

et al., in press).

Engineer Creek, Alaska (RAN; Richards et al., in

press).

6. Cleary, Alaska (RAN; Richards et al., in press).

. Goldstream (“G. S. Dredge’’), Alaska (RAN; Rich-
ards et al., in press).

. Keams Canyon, Navajo County, Arizona (RAN;
Ray, 1969).

. Rancho La Brea, Los Angeles County, California
(RAN; Merriam & Stock, 1925; Kurtén, 1967;
Agenbroad & Mead, 1986).

. McKittrick, Kern County, California (RAN; Schultz,
1938; Kurtén, 1967).

2:
3:
4.

3;

28

1M ls

12:

13.

14.

26.
Oa ke

Irvington, Alameda County, California (IRV; Kur-
tén, 1967).

Potter Creek Cave, Shasta County, California (RAN;
Merriam & Stock, 1925; Kurtén, 1967; Ray, 1969).
Jinglebob, Lone Tree Arroyo, Meade County, Kan-
sas (RAN; Kurtén, 1967).

Arkalon, Seward County, Kansas (IRV; Richards et
al., in press).

. Bat Cave, Pulaski County, Missouri (RAN; Hawk-

sley, 1965; Kurtén, 1967).

. Perkins Cave, Camden County, Missouri (RAN;

Hawksley, 1965; Kurtén, 1967).

. Hay Springs, Sheridan County, Nebraska (IRV;

Kurtén, 1967; Richards et al., in press).

. Labor-of-Love Cave, White Pine County, Nevada

(RAN; Emslie & Czeplewski, 1985).

. Frankstown Cave, Blair County, Pennsylvania (RAN;

Peterson, 1926; Kurtén, 1967).

. Hot Springs Mammoth Site, Fall River County,

South Dakota (RAN; Agenbroad & Mead, 1986).

. Cueva Quebrada, Val Verde County, Texas (RAN;

Lundelius, 1984).

. Rock Creek (2 mi north of Equus quarry), Briscoe

County, Texas (IRV; Kurtén, 1967).

. Silver Creek, Salt Lake County, Utah (RAN; Nelson

& Madsen, 1983).

. Natural Trap Cave, Big Horn County, Wyoming

(RAN; Agenbroad & Mead, 1986).

. Little Box Elder Cave, Converse County, Wyoming

(RAN; E. Anderson, 1968) (as Ursus arctos).
Zacoalco, Jalisco, Mexico (UNK; Aviiia, 1969).
Tequixquiac, exact locality unknown, Mexico (RAN;
Freudenberg, 1910; Richards et al., in press).

FIELDIANA: GEOLOGY

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LENGTH M2

Fic. 18. Bivariate graph of Arctodus simus—Length and anterior width of M2. For explanations of abbreviations,

symbols, and type localities, see p. 28.

slightly shorter in relation to the femur than in the
other individuals. This is apparently not a sexual
characteristic, as the Cueva Quebrada specimen
(considered to be a female) has the longest skull
in proportion to the femur length. A peculiarity
of the Indiana specimen is the extreme narrowness
of the postorbital constriction.

In limb proportions, the radius/humerus (100
x radius/humerus) and the tibia/femur (100 x
tibia/femur) indices (values of 80.8 and 73.4, re-
spectively) are within the range reported for other
A. simus specimens (Kurtén, 1967; Emslie & Cza-
plewski, 1985).

In life the Indiana Arctodus specimen would have
stood more than 158 cm (5’2”) at the shoulder.
Using the combined minimum midshaft circum-
ference of the femur (171 mm) and humerus (196
mm) as predictors of body mass (J. F. Anderson
et al., 1985; Roth, 1990; Damuth & MacFadden,
1990), the weight of the Indiana specimen was
calculated at 766 kg. This is about the weight for
the largest of the Alaskan brown bears (Ursus arc-
tos, up to 780 kg; Walker, 1964), and heavier than
the values for A. s. yukonensis, based on the area
of the cross-section of the femoral shaft presented
in Kurtén (1967; 590-630 kg) and Nelson and
Madsen (1983; 620-660 kg).

The radiocarbon date of 11,500 + 520 B.P. is
among the few direct dates available on Arctodus
bone and is among the youngest dates for the spe-
cies. Other young dates on materials associated

with Arctodus simus are 11,220 + 110 to 11,420
+ 110 B.p., Huntington Dam, Sanpete County,
Utah (Gillette & Madsen, 1992); 12,650 + 350
B.P., Lubbock Lake, Lubbock County, Texas, where
A. simus foot bones have aboriginal cut marks
(Kurtén & Anderson, 1980; Johnson, 1987); 12,650
+ 70 B.p., Silver Creek, Salt Lake County, Utah
(Nelson & Madsen, 1983); and ca. 11,500 B.P.,
Burnet Cave, New Mexico, where A. simus was
associated with a Clovis point (Hester, 1967). Em-
slie and Czaplewski (1985) reported a date of 5,320
+ 120B.p. for the Labor-of-Love Cave, White Pine
County, Nevada, specimen but did not accept it
as giving the true age of the specimen. Churcher
et al. (1993) summarized other radiocarbon dates
available for Arctodus simus.

Pollen spectra from several Indiana localities
suggest the dominant regional vegetation at the
time the bear at the Rochester site was deposited.
Marls from a kettle lake at the Wells Mastodont
Site, Fulton County (about 11 km northwest of
the Arctodus locality), were dated at 12,000 + 450
B.P. in the mastodont level (Gooding & Ogden,
1965). Pollen spectra were dominated by spruce
(Picea), with ash (Fraxinus), fir (Abies), and birch
(Betula) represented, until 10,500 B.p. when the
coniferous forest gave way to oak (Quercus).

At another site, the Kolarik mastodont locality,
Starke County (about 35 km northwest of the
Rochester Arctodus locality), mastodont bones were
associated in a sandy muck with pollen, indicating

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 29

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ig a cS eal en ie es
5 1,27
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ah St ee ee
29 30 31 32 33 34 35
LENGTH m1

Fic. 19. Bivariate graph of Arctodus simus—Length and posterior width of m1. For explanations of abbreviations,

symbols, and type localities, see p. 28.

an open or patchy boreal forest dominated by
spruce, with a high percentage of fir, ash, and herbs
and a low percentage of pine, oak, and other hard-
woods (Jackson et al., 1986). This sequence, dated
at >12,000 B.P. to at least 11,000 B.P., was suc-
ceeded by a dominance of pine and other hard-
woods. Clayey marls preceded the bone-bearing
sandy muck level.

The Christensen mastodont locality, 145 km to
the south, preserved an open, white spruce-dom-.
inated boreal forest at 14,000-—13,000 B.P. asso-
ciated with the bone levels. This forest was fol-
lowed thereafter by the immigration of hardwoods
(Whitehead et al., 1982). Similar pollen spectra
are known throughout the central Great Lakes re-
gion (Bailey, 1972; Bailey & Ahearn, 1981; King,

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A | poy | 123
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1 bet | | ms |
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480 500 520 540 560 580 600 620 640 660 680 700 720 740

FEMUR: GREATEST LENGTH

Fic. 20. Bivariate graph of Arctodus simus—Femur, greatest length and distal width. For explanations of abbre-

viations, symbols, and type localities, see p. 28.

30

FIELDIANA: GEOLOGY

TABLE 9. Absolute dimensions and ratio (%) of skull length to the other skull dimensions and to other select
elements among associated Arctodus simus specimens. (Measurements in millimeters.)

Hay Springs, Hot Springs, Cueva Quebrada,

Measurements Rochester, Indiana Maine South Dakota Texas
Skull, basal length 396 a413 402 343
Skull, zygomatic breadth 319 (124%) _ 244.5 (164%) 207 (166%)
Skull, width over postorbital

processes 170.7 (232%) al84 (224%) 167 (241%) -_
Dentary, canine to condyle 298 (av.) (133%) 295 (140%) _ _
Femur, greatest length 651 (61%) 658 (63%) _ 508 (68%)
Tibia, greatest length 478 (83%) 492 (84%) —_ a

1981) and the eastern United States (Delcourt &
Delcourt, 1981). Thus, the Quaternary palynology
of the region suggests that the Rochester Arctodus
likely existed in an open or patchy boreal forest
dominated by spruce.

Kurtén (1967) visualized Arctodus simus as a
long-legged, fast-running, powerful predator, ca-
pable of preying on bison, deer, horse, or ground
sloth. Emslie and Czaplewski (1985), however,
suggested that A. simus was an omnivore or her-
bivore, highly capable of scavenging, and that the
proportions of the long limbs were not necessarily
adapted for running, but perhaps for pulling down
vegetation or for increased visibility within tall
ground cover in open habitat. Voorhies and Cor-
ner (1986) felt that the skull morphology and jaw
mechanics of A. simus confirmed its predaceous
nature. Shaw and Cox (1994) believe that A. simus
was more carnivorous than living bears, except
the polar bear. With most modern ursids being
omnivorous, it seems to us unlikely that a ten-
dency for the omnivorous way of life would have
been abandoned by Arctodus, even though its teeth
and limbs strongly suggest that it also had spe-
cialized for a predatory life-style. Thus, we suggest
that Arctodus simus was a highly predaceous om-
nivore, with a preference for larger-sized prey.

Harington (1980), examining the geographic
distribution of A. simus, suggested that the species
occupied high, well-drained grasslands. Both Kur-
tén (1967) and Nelson and Madsen (1983) have
suggested that A. simus inhabited open prairie or
savanna environments. The context of the isolated
Indiana specimen suggests only that short-faced
bears occupied the open boreal forests of northern
Indiana ca. 11,000—13,000 years ago. Apparently
Arctodus occasionally also frequented lowland lake
and swamp environments such as the depositional
situation described in this paper suggests.

Potential local prey may have included the large
species: Jefferson’s ground sloth (Megalonyx jef-

fersonii), giant beaver (Castoroides ohioensis),
American mastodont(Mammut americanum), Jef-
ferson’s mammoth (Mammuthus columbi jeffer-
sonii), tapir (Tapirus sp.), horse (Equus sp.), flat-
headed peccary (Platygonus compressus), long-
nosed peccary (Mylohyus nasutus), stag-moose
(Cervalces scotti), elk (Cervus elaphus), white-tailed
deer (Odocoileus virginianus), caribou (Rangifer
tarandus), ancient bison (Bison bison antiquus),
Harlan’s musk ox (Bootherium bombifrons), and
tundra musk ox (Ovibos moschatus) (Richards,
1984). Potential plant foods have not been inves-
tigated.

Cave finds of A. simus are common (Emsilie,
1985; Emslie & Czaplewski, 1985; Hansen, 1992;
Hawksley, 1986; Kurtén & Anderson, 1980; Nel-
son & Madsen, 1983; Puckette, 1976), and some
“‘bear beds” present in Missouri caves appear to
indicate denning (Hawksley, 1965).

Acknowledgments

The late Dr. Greta Woodard, Department of
Biology, North Central campus, Purdue Univer-
sity, Westville, Indiana, in responding to the re-
covery of an unknown skeleton brought about the
preservation of this important Arctodus specimen.
Both Clayton E. Ray, National Museum of Nat-
ural History, and Richard H. Tedford, American
Museum of Natural History, kindly reviewed ear-
lier drafts of this manuscript and made helpful
recommendations for our study. We are particu-
larly indebted to Richard Tedford for encouraging
one of us (R.L.R.) to examine and measure ma-
terials in the AMNH collections (including Arkalon,
Kansas, and Alaska) and to present data not pre-
viously published. We gratefully acknowledge the
counsel of Ellis J. Neiburger and Bruce Rothschild
in the interpretation of skeletal pathology. Ned
Bleuer, Indiana Geological Survey, Bloomington,
helped greatly with the interpretation of northern

RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA 31

Indiana glacial geology. James McKean, Indiana
State Museum, produced Figures 1-15, with some
supplements by Fred Lewis. Dave Rieger pro-
duced Figures 17-20. Linda Badger, Indiana State
Museum, typed the final drafts of the manuscript.

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RICHARDS & TURNBULL: ARCTODUS SIMUS FROM INDIANA eS

Appendix

Osteomyelitis in the Ulnae of the
Bear, Arctodus simus

E. J. Neiburger!

Lesions on the Ulnae

Each ulna exhibits an area of osteolytic-osteo-
plastic activity.

1. The right ulna shows an area of periostitis
with one defect located 11 cm from the distal end
of the bone. The defect is an oval opening 1.5 cm
x 0.8 cm in size and dissecting through the cortex
to a depth of 2.0 cm. The opening is surrounded
by a 0.5-cm elevation of bone extending in an area
3 cm in diameter around the opening. This bone
is quite dense.

There are four exostoses (0.2-0.8 cm) located
on the edge of the defect, probably caused by hy-
perplastic bone formation responding to an infec-
tion.

2. The left ulna presents three related lesions
encircling the midshaft area, 24 cm from the distal
end. The largest lesion is 1.0 cm in diameter with
a slightly raised lip and a few small (0.1-0.3 cm)
exostoses protruding from the edge. The lesion
dissects 2.0 cm into the cortex. The second lesion
is located 2 cm posterior to the first. It is 0.5 cm
in diameter and 0.5 cm deep. The third lesion is
located 1.0 cm posterior from the second. It is 1.0
cm wide and 0.5 cm deep.

These lesions do not appear interconnected by
blunt probing but occupy a slightly elevated area
of bone that “‘rings”’ the shaft. The bone surround-
ing the lesions in both ulnae is dense and has little
porosity and no fracture lines or other defects.

' 1000 North Avenue, Waukegan, Illinois 60085.

34

Interpretation

These lesions are quite typical of the abscess-
fistula formation seen with osteomyelitis, a chron-
ic infection of the bone.

These lesions are localized and have not spread
to the medullary portions of the bone shaft. They
are well circumscribed, only slightly hyperplastic
and dense. These conditions are typical of a long-
standing (multi-year) infection in which the bear’s
resistance is just able to contain the infection but
not strong enough to heal it. Such infection results
from either lowered host resistance or a very vir-
ulent pathogen. In this situation, the area becomes
a localized sore that constantly drains, repeatedly
flares up, and then partially heals, only to flare up
again.

The area of the lesion is occasionally painful but
not totally debilitating. Such wounds could be ini-
tiated by a penetrating or crushing injury that
breaks the skin surface and becomes infected.

In wild animals, these wounds quickly heal or
progress to a fatal condition. It is rare to see such
wounds, as in this case, remain in a static state for
such a length of time as to produce the hyperplastic
bone formations noted.

FIELDIANA: GEOLOGY

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