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PLATE 1

BULLETIN 171

U. S. NATIONAL MUSEUM

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Topographic map showing location of Cumberland Cave and its relation to the physiography.
The cave is indicated (X) on north end of spur just south of Corriganville, Md. Map is
Scale, 1/62500, or about

taken from U. S. Geological Survey Frostburg Quadrangle.
Contour interval, 20 feet.

1 mile to the inch.

SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM

BuLuetin 171

THE PLEISTOCENE VERTEBRATE
FAUNA FROM CUMBERLAND CAVE
MARYLAND

BY
JAMES W. GIDLEY anp C. LEWIS GAZIN

UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON : 1938

LSS La ee
For sale by the Superiatendent of Documents, Washington, D.C. - - - - <= - - Price 25 cents (paper)

The scientific publications of the National Museum include two
series, known, respectively, as Proceedings and Bulletin.

The Proceedings series, begun in 1878, is intended primarily as a
medium for the publication of original papers, based on the collec-
tions of the National Museum, that set forth newly acquired facts
in biology, anthropology, and geology, with descriptions of new
forms and revisions of limited groups. Copies of each paper, in
pamphlet form, are distributed as published to libraries and scientific
organizations and to specialists and others interested in the different
subjects.

The dates at which these separate papers are published are recorded
in the table of contents of each of the volumes.

The Bulletin series, the first of which was issued in 1875, consists
of separate publications comprising monographs of large zoological
groups and other general systematic treatises (occasionally in several
volumes), faunal works, reports of expeditions, catalogs of type speci-
mens and special collections, and other material of similar nature.
The majority of the volumes are octavo in size, but a quarto size has
been adopted in a few instances in which large plates were regarded
as indispensable. In the Bulletin series appear volumes under the
heading Contributions from the United States National Herbarium, in
octavo form, published by the National Museum since 1902, which
contain papers relating to the botanical collections of the Museum.

The present volume forms No. 171 of the Bulletin series.

ALEXANDER WETMORE,
Assistant Secretary, Smithsonian Institution.

WasHINGTON, D. C., January 26, 1988.
Ir

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Z\ASONIA,
ZAM 7 Ay

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( FEB

CONTENTS

Matroductione. oes oo a> oe LEE SESS Sea ee Sea en oe
History of investigation....--.------+---~------+-+<22----2+-----=---

Location of Cumberland Cave

Accumulation and preservation of migterial] os Oe ee

Relationship of cave to physio

graphy... -----------------------------

Gumberland Cave fauna.--_-----=++-------~-----=-45-+-=+2---+-=---
Environment of the fauna --------------------------------------

Age and relationships of t

Heal Ase ee i eS aa aa

Systematic description of vertebrate fauna__._-2_--------+-----=------
Order Loricata. 4.2222 2221-22222 242-2 2-2-2 2 2 - eee nnennae =
Grocadwaid tees oi ae 22s as See nee eee

Order Serpentes.=+-----------2425-+--42-22--25- 25-5 eee noone
Ophidian remains------.-------------------=----------------
@rder-Galliformes= 2+ ---.--22 22242. =. 2222-2 2 = eee enna
Family Tetraonidae-_---------------------------------------

Bonasa umbellus

(Linmseus) 224 222222. Sa === =

@rderinsectivoralt= 22 2.==52 == 222225--2-2--=------ == nae
amily Soricidae=++2-++=-2--+2=+----==+-----2---=---=""-=-
ores Spuse=sh22-5-¢5-chhe2 set essene Sse e-- 6 Sana anee

Blarina brevicauda (Say) --------------------------------

@ider @hiraptera- 2-22 S222 22-2 2S Sele ae == Siete a
Family Vespertilionidae------------------------------------
Eptesicus cf. grandis (Brown) 22 22-282) RU seo-- ===
Corynorhinus alleganiensis Gidley and! Garin? Seee so te ee

Grder Garmivotal 2-22: 35-2429 =2225-552522242=2=--~--- ===
Family Canidaessss22: $2222 2222222-222+22255~---=-- 22S oe

Canis armbruster

feGidlay sa fusees -Satlsks 4S SoSe Ses

Canis ef. priscolatrans Cope-.----------------------------
BamilysUreidaess) 22522 F242 23 22c Baas sash == 4 —-- 2 oo e

Euarctos vitabilis
Arctodus haplodo
Family Mustelidae - -
Martes parapenn
Mustela cf. vison
Gulo gidleyt Hall

(Gidley) i242 222 shs22se 235 ~ = 22 = == =
fh (Cope) Be Saas eee ea a= oe
anti Gidley and Gazin- sy Ne \ sR

Sehirebers= se ee 2 ee Pe eae ase

Lutra parvicuspis Gidley and Gazin_---------------------

Brachyprotoma p

ristina Brown: 2222222222252 22=-2-=-=---=-

Spilogale marylandensis Gidley anid) Cazine =) Sewers ee Se
Taxidea marylandica Gidley and Gazin-------------------
anilysWelidwe>\] sss DUNE rears ore en ee aa
Felis cf. inexpectata (Cope) ------------------------------
Foliemear atrov Leidy 22 2! = 2222 2522252 s24==-3422--2-2-==

IV BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Systematic description of vertebrate fauna—Continued. Page
Order: Rodentisia=2. 232 = ee ae ee ee ee ee eee 53
Hamily Sciuridaes." 2. oa eae ee ee ee ee 53
Marmotavmnonacc) (ui seus) arse: aye er eee eae 53

Citellus ef. tridecemlineatus (Mitchill)......._....._________ 54

Tamias cl. striatus: (hinnaens) eee see ee ee 54

Sciurus (Tamiasciurus) tenwidens Hay__._____.__._______- 54
Glaucomys: spe ae ee Se ae Sl a See ee ee ee 56

Family -Geoomyidae.s 3 = ese a ee ee 57
Plesivithomomys potomacensis Gidley and Gazin__-__________ 57

Family" Castoridae. =: sso 2ees 2222 5. ee eee eee 59
Castonm canadensis Kohl a a= ee ee 59

Family ‘Cricetidae. 22 2 S22 2s. ee 59
Peromyscus cf. lewcopus (Rafinesque)...._._......._--_-_._ 59
Neotomasmagester: Dard eee es oe ee ee 59
Parahodomys spelaeus Gidley and Gazin.-_._.._..__--_.-- 60
Synaplomys cl. coo pen Dard. anes a nae oe ee 62

SUNG DLOMYS MACON YS) Spe ae ee ee 63
Microtus (or Pitymys?) cf. involutus (Cope). ___---._.------ 63
Ondatrarctannectens: (Brown) 29a" soe ae ee 65

Family, Zapodidacs— 222 so. es Se 67
LODUS BD ea ee ee ae ree eee, Seah a 67
Napacozapusch. tasrgnis. (Miller) S22 7 9. Se ee eee 67

Kamiuly Erethizomtidaes 2 32.2 ea ee 67
Hrethizon cl. dorsatum.(einnaeus)) == 3 ee ae 67

Order Lagomorpha _-______------ Cert ee i: ee Bak anes ria 68
Family Ochotonidae_______- wc oN) eS SD ele ne pe ee 68
OCTIOLON GUST) Baars te te Se rn ee ey or eee 68

Family Leporidae: _... =... et Saxe ipl he Os ee glee eee 69
Lepus cf. americanus Mrxiebens. = 2.2. = enon a eee 69

Order Proboseides: . 6. 23-22. Saale es seen oy ee oe 70
Family Mastodontidae so: )\ 422 ae eee 2 eee ee ee 70
Mamimutsck.. americanwin: (Nernst 70

Order Perissodactyla 224 8 2 ae ee ee ee es 70
WamnilyBiguidae.s. == ot ete ee oes ee get een a 70
FguusiSD. osc ce coe ee 32 a a ee see 70

Family Tapiridaé.. #s2“25.. 2 22-2 eee ee eS 70
TODiTUS Spi. ee oe ee A ee ee ee 70

Order Artiodactyla: 2.282232. ee = So a ee 71
Vamily Tay assuidae 3 soe. 5) 2 a ee 2 71
Platygonus cumberlandéensis Gidley. -22-— ee tee 71
Platygonus vets (2) eid 5) 532 ae ee ee ee 82
Malohyus: exortivus Gidléy=— 22 5a seer See eee 83
Mylohyus cf. pennsylvanicus (Leidy)__........--.-------- 86

Family ‘Cervidae 23 <2 oe scat be eee ee ee 86
Cervus'S Pies 2nd 2 See eS ee ee eee 86
Odocoileus cf. virginianus (Boddaert)__.._.._.__._._-.-------- 86

Family, Bovidsaeteeeu2- 52 S642 2 ee ee ee ee ee 86
Euceratherium(?) americanum (Gidley) _._..-...---.------ 86
Literature cited: 2. 224.45 eh ee eee re ee 91

MARS ae Sr So nye ee Are, nT ec a 95

—_

16.

he
18.
Ls
20.
21.
22.
23.
24.
25.
26.
27.
28.

29.
30.
31.

32.
33.
34.
35.
36.
37.
38.
39.

ILLUSTRATIONS

FIGURES

_ Skull and upper dentition of Corynorhinus alleganiensis Gidley and

WSkullbol Canisianmorustert: Gidley 2s 45 = eas ee eee eee eee
. Skull and lower carnassial of Canis armbrusteri Gidley _-------------
Per cunt tak Oras Mc TiT7a Nee SC Ty Gx IL a pen eee
Parle ot Ga rau sai L0 Tess Cerium Onl CU Cys eee ee ae
. Left ramus of mandible of Canis armbrusteri Gidley -_--------------
. Right ramus of mandible of Canis armbrustert Gidley---------------
. Left ramus of mandible of Canis armbrusteri Gidley __--------------
. Right ramus of mandible of Canis armbrusteri Gidley----~----------
. Right ramus of mandible of Canis armbrustert Gidleysn nse ae we
. Right ramus of mandible of Euarctos vitabilis (Gidley) --------------

Skull and mandible of Fuarctos vitabilis (Gidley) ---~----------------

PASiculOt archos UILQUULUS, (GiGIGy a2 oe epee ee
PaSkullof uanrchos vraurits. (GiGlCY 222 ae ce ee ee ee ee
. Anterior portion of male skull, right rami of female mandibles, and left

ramus of male mandible of Martes parapennanti Gidley and Gazin_-
Anterior portion of skull and right ramus of male mandible of M ustela

GIA i STET SCHROMEIS Best aa ea eee a ate Beer Aree Seen
Anterior portion of skull and mandible of Mustela ef. vison Schreber_-
Right ramus of female mandible of M/ustela cf. vison Schreber--------
Skull ane menditsteof Gauloigzuareys tall 2-2) 2
Slaull"@f Grlovgidlere: tial Sa St ea A a eee
Maxillary portion of skull of Lutra parvicuspis Gidley and Gazin_---
Skull of Brachyprotoma pristina Brown__-_--_-----------------------
Left ramus of male mandible of Brachyprotoma pristina Brown_------
Right raraus of mandible of Brachyprotoma pristina Brown- ---------
Right ramus of mandible of Spilogale marylandensis Gidley and Gazin_ -
Skull and mandible of Taxidea marylandica Gidley and Gazin__------
Skull of Taxidea marylandica Gidley and Gazin_----~--~-------------
Upper carnassial and left ramus of mandible of Felis cf. inexpectata

(SONG) ee ea ee ee pple rgd Noe erg SL Ped
Hoot DOMES OL MELTS MEAT GETOT Ln Cyt ames ee eee ere
Skull and mandible of Sciurus (Tamiasciurus) tenwidens Hay _~------
Left ramus of mandible of Plesioithomomys potomacensis Gidley and
Left ramus of mandible of Neotema magister Baird. __--------------
Right ramus of mandible of Parahedomys spelacus Gidley and Gazin_-
Right ramus of mardible of Parahodomys spelaeus Gidley and Gazin_
Right ramus of mandible of Microtus (or Pitymys?) ef. involutus (Cope) -
Left ramus of mandible of Ondatra cf. annectens (Brown) ------------
Skull and mandible of Erethizon cf. dorsatum (Linnaeus) - - - ---------
Rastilower molarot.laperus spot -) 522 ee ee ee eee ea
Skull of Platygonus cumberlandensis Gidley ------------------------

Page

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vI BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Page
40. Skull of Platygonus cumberlandensis Gidley_.._.--.---.------------ 74
41. Skull of Platygonus cumberlandensis Gidley.....-.--.-------------- 75
42. Skull of Platygonus cumberlandensis Gidley_..-..---.-.------------ 76
43. Skull of Platygonus cumberlandensis Gidley_..-.-.---------------- a
44. Skull of Platygonus cumberlandensis Gidley----..-.-.-------------- 78
45. Left hind foot of Platygonus cumberlandensis Gidley __....---.------ 79
46. Right ramus of mandible and cheek teeth of Mylohyus exortivus Gidley - 83
47. Cheek teeth of Mylohyus exortivus Gidley__..-.....-..---.--..---- 84

48. Superior deciduous premolars and first permanent molar of right side
of Mylohyus exortious Gidley 2...) 32 Se 84
49. Superior cheek teeth of Euceratherium(?) americanum (Gidley) -__-_-- 87

50. Left ramus of mandible and right lower premolars of Euceratherium(?)
americanum (Gidley) = 32626222 a id Be eee 88

PLATES

Facing page

1. Topographic map showing location of Cumberland Cave and its rela-
tion tothe pliysiopraphy.2_- <= ee ee ee ee ee Frontispiece

2. Cut made by Western Maryland Railroad Co., which uncovered the

Camberland Cave deposit 22. 3.22222 os.52 cae ee eee

3. View from opposite side of railroad cut, showing fossil deposit at bot-
tom, near track, and traces of ancient opening at top of cliff_____-- 3
4. Near view of part of excavation made in fossil deposit___.._.__.-__-- 6
5. Later view of opening to excavation made in fossil deposit. _______-__ ti

6. Small portion of cave interior, showing bones sticking in clay and
stalactitic material. 2-23.52 So cee oe ee er 10

7. View from spur in which cave is located, looking north across Wills
Greek Valley. 2c ce ae ay hg ee a oe ee ee Bl

8. Mounted skeleton of bear (Euarctos vitabilis Gidley), from Cumber-
land Cave, exhibited in U. S. National Museum_____-___-------- 26

9. Mounted skeleton of wolverine (Gulo gidleyi Hall) from Cumberland
Cave, exhibited in U. S. National Museum________._____.__----- 27

10. Mounted skeleton of peccary (Platygonus cumberlandensis Gidley)
from Cumberland Cave, exhibited in U.S. National Museum- - - ---- 70

THE PLEISTOCENE VERTEBRATE FAUNA FROM
CUMBERLAND CAVE, MARYLAND

By JAMES W. GIDLEY and C. LEWIS GAZIN

INTRODUCTION

LimMESTONE caverns have played a major role in preserving a record
of the Pleistocene life in the eastern region of North America. Except
for the remarkable deposits in Florida, our knowledge of the com-
position of Pleistocene mammalian faunas from this region has been
limited largely to occasional and fortunate cave finds. This mode
of accumulation rivals that of the tar pits in presenting a broad cross.
section of the land life belonging to the geclogic period immediately
preceding the present.

Caves are abundant in the limestone areas of the Appalachian and
Mississippi Valley regions, but very little systematic exploration has
been done with a view toward obtaining representative collections of
Pleistocene mammals. Strong impetus for exploratory work has come
from the field of archeological research, and many of the important
paleontological finds were made during the investigations for the
remains of man. The search for saltpeter has also led to the exam-
ination of many caves, but in this work the importance of fossil re-
mains found has undoubtedly been overlooked in the majority of
instances.

Cumberland Cave was first found by workmen in excavating for a
railroad cut near Cumberland, Md. Considerable fossil material was
destroyed by steam shovel and dynamite before the significance of
the find was known, but the abundant material subsequently obtained
includes a remarkable variety of mammalian forms, many of which
are represented by unusually well preserved remains.

The junior author is greatly indebted to C. W. Gilmore, curator
of vertebrate paleontology at the National Museum, for advice and
criticism during the preparation of this report, and to Dr. J. B.
Reeside, Jr., and Dr. W. C. Alden, of the United States Geological
Survey, for a better understanding of the geologic and physiographic
relations of the cave. Acknowledgment is also made of the courtesy
extended by G. S. Miller, Jr., Dr. Remington Kellogg, and staff of
the division of mammals in the National Museum and Dr. H. H. T.
Jackson, EK. A. Goldman, and A. H. Howell, of the United States
Biological Survey, in giving generously of their time and advice and

2 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

in permitting unreserved use of the collections of recent mammals,
Dr. Walter Granger, of the American Museum of Natural History,
and Dr. C. L. Camp and R. A. Stirton, of the University of California,
have aided greatly in loaning type material from their cave collections.
The drawings included with the text were made by Sydney Prentice
and Rudolf Weber.

HISTORY OF INVESTIGATION

The attention of the United States National Museum was directed
to the occurrence in 1912 by Raymond Armbruster, of Cumberland,
Md., and by George Roeder, of Swetnan, Va. Investigation of the
occurrence was undertaken by Dr. J. W. Gidley, and quarrying
operations in the cave were conducted by him at intervals during
the years 1912 to 1915. For a part of this time he was assisted in
the excavation work by Mr. Armbruster. The exploration work
carried on at the cave is interestingly described in three popular papers
published in “Explorations and Field-Work of the Smithsonian
Institution” for the years 1913, 1914, and 1915.

Early results of Gidley’s study of the fauna were published in two
short papers that appeared in 1913. These included a description
of an “eland’”’ (1913a) (see also Gazin, 1933), and new species of wolf
and black bear (1913b). In later papers Gidley (1920a, 1920b) gave
a general account of the occurrence and described the peccary remains
taken from the cave. Dr. Alexander Wetmore (1927) reported the
remains of a ruffed grouse included in the collection.

Since the death of Dr. Gidley in 1931 it has become the junior
author’s privilege to continue the investigation of the Cumberland
Cave fauna, completing the manuscript and describing those portions
of the fauna that had not been studied. In 1933 a revised list of the
forms encountered in the investigation was published (Gidley and
Gazin, 1933) together with descriptions of the remaining new forms
in the fauna.

LOCATION OF CUMBERLAND CAVE

Cumberland Cave is located in Allegany County, western Maryland,
about 4 miles northwest of Cumberland (see pl. 1). It is situated in
a cut made by the Western Maryland Railroad through the north
end of a ridge just south of Corriganville, near where Jennings Run
joins Wills Creek. Its location as determined from the topography
shown on the Frostburg quadrangle, U. S. Geological Survey, is at
about latitude 39°41’ N. and longitude 78°47’ W.

The cave is exposed in the cut at the base of an escarpment of
Devonian limestone about 75 feet high. Dr. J. B. Reeside, Jr., of
the United States Geological Survey, informs the writer that the

PEATE, 2

BULLETIN 171

U. S. NATIONAL MUSEUM

‘yisodap JAV) puevylequin’) oy potaAooun Yor M “40{5) PVOIT[IL | puvyAlryy uJ So

M Aq ope

w yng

PLATE 3

BULLETIN 171

U. S. NATIONAL MUSEUM

near track, and

at bottom

deposit

ssil

fo
Photographed in 1913 by Raymond Armbruster.

showing

ilroad cut

Tal

yosite side o
traces of ancient opening at top of cliff.

1e@W fre mM OPT

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 3

limestone at this point belongs to the Keyser member of the Helderberg
formation, and is about 50 feet stratigraphically below the base of
the Oriskany. The limestone is structurally a part of the steeply
dipping west flank of the Wills Mountain anticline. In the vicinity
of the cave the strata are standing nearly vertical, a fact that probably
aided greatly in the penetration of surface waters.

ACCUMULATION AND PRESERVATION OF MATERIAL

The following account of the conditions encountered at the cave is
taken from Gidley’s preliminary report (1913b, pp. 94-95) on the
occurrence:

In making the railway cut, several small chambers at bigher levels than the
one containing the bones were encountered, and before the work of excavation
began there was said to have been an opening to the surface on the crest of the
hill directly above the middle of the present roadbed. This opening so nearly
overhead probably at one time served as a trap through which were introduced
the animals whose remains are now in the deposits of the bone cavern. ‘There
are other openings along the line of outcrop of the ledge, one of them at about
the same level with the bone-bearing deposits, appearing at the north end of
the ridge where it slopes abruptly down into the Wills Creek Valley. These
openings may or may not have communicated at one time with the caverns
intersected by the railroad cut, but probably had nothing to do with the accumu-
lation of material in the latter.

From Brown’s [1908, p. 163] account of the Conard Fissure, it would seem
that the conditions governing the accumulation of material in the Cumberland
Cave were quite similar. The bones for the most part are much broken, yet
show no signs of being water worn. They are found scattered fairly uniformly
throughout the entire mass of unstratified accumulations which consist entirely
of cave clays and breccias, unevenly hardened and more or less cemented together
by stalactitic materials. There is an almost entire absence of admixture of sand
or gravel, or in fact anything that would suggest the possible aid of stream currents
in sorting or placing the material during the process of accumulation.

RELATIONSHIP OF CAVE TO PHYSIOGRAPHY

Five geographic provinces are recognized in Maryland (Fenneman,
1930): Coastal Plain, Piedmont Plateau, Blue Ridge, Valley and
Ridge Province, and Appalachian Plateau. The region about
Cumberland Cave belongs physiographically in the western part of
the middle section of the Valley and Ridge Province.

Physiographic studies of western Maryland and vicinity (Abbe,
1900; Clark and Mathews, 1906; Stose and Swartz, 1912; Stose and
Miser, 1922) show that several surfaces, representing periods of ces-
sation of downward stream-cutting, have been developed. Rem-
nants of the various upland surfaces recognized are commonly
represented by sets of ridge tops having accordant levels. More
recent stages in the process of stream erosion are seen in the different
cravel-strewed terraces.

4 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Perhaps the oldest surface recognized, which has been referred to
the Schooley Plain, includes the higher portions of the Allegheny
Mountain section of the Appalachian Plateaus. A somewhat lower
level may be represented by the crest of Wills Mountain (see pl. 1)
just east of Cumberland Cave and having an elevation varying from
1,400 feet to 1,800 feet. A still lower surface is believed to be indi-
cated by the crest of Shriver Ridge just north of the town of Cumber-
land and by a spur of Knobly Mountain to the south of the town.
W. B. Clark and E. B. Mathews (1906, p. 89) correlated these ridge
crests with the Harrisburg surface, and Cleveland Abbe, Jr. (1900,
p. 52), considered them as representing a surface which he called the
Shenandoah Plain. The ridges have an elevation of about 1,100
feet near the Potomac River, and if the surface represented were
extended up Wills Creek Valley at the present stream gradient it
would have an elevation of about 1,200 feet at Cumberland Cave.
It may be represented, as suggested by Clark and Mathews, by the
ridge west of Wills Mountain, in which Cumberland Cave is located.
This ridge crest varies in elevation from about 1,040 feet near the cave
to about 1,300 feet farther south. The elevation of the cave deposit,
as determined by O. P. Hay (1923) is 837 feet. Assuming practically
no gradient for the surface between Cumberland and Cumberland
Cave, we have a minimum difference of nearly 300 feet between the
elevation of the cave and that of the surface.

During later stream history (Abbe, 1900) two important terraces
were developed in the vicinity of Cumberland at elevations of about
800 feet and 650-700 feet. The higher of these may have had an
elevation of about 900 feet on the spur in which the cave is located.
This is probably very close to the elevation at which the cave mouth
was located. The lower terrace may have been 750-800 feet in
elevation near the cave. This level appears to be slightly lower than
the cave deposit. It seems evident that the fossil material did not
accumulate until cutting by Wills Creek had at least reached the
stage represented by the higher terrace, and probably at a somewhat
later date when the drainage system had dissected this surface.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 5
CUMBERLAND CAVE FAUNA

The classified list that follows includes all the Pleistocene vertebrate
fauna identified from the Cumberland Cave and herein treated:

Loricatva:
Crocodylid.
SERPENTES:
Ophidian remains.
GALLIFORMES:
Tetraonidae:
Bonasa umbellus (Linnaeus).
INSECTIVORA:
Soricidae:
Soreaz sp.
Blarina brevicauda (Say).
CHIROPTERA:
Vespertilionidae:
{Eptesicus cf. grandis (Brown).
tCorynorhinus alleganiensis Gidley
and Gazin.
CARNIVORA:
Canidae:
Canis armbrusteri Gidley.
tCanis cf. priscolatrans Cope.
Ursidae:
}Huarctos vitabilis (Gidley).
tArctodus haplodon (Cope).
Mustelidae:
+Martes parapennantt Gidley and
Gazin.
Mustela cf. vison Schreber.
tGulo gidleyz Hall.
tLutra = parvicuspis
Gazin.
tBrachyprotoma pristina Brown.
+Spilogale marylandensis Gidley
and Gazin.
}Taxidea marylandica Gidley and
r Gazin.
Felidae:
}Felis cf. inexpectata (Cope).
{Felis near atrox Leidy.
RODENTIA:
Sciuridae:
Marmota monaz (Linnaeus).
Citellus ef. tridecemlineatus (Mitch-
ill).
Tamias cf. striatus (Linnaeus).
tSciurus tenuidens Hay.
Glaucomys sp.
Geomyidae:
tPlesiothomomys potomacensis Gid-
ley and Gazin.

Gidley

+ Extinct species.

t Extinct genera.

RopeEntT1A (Cont.):

Castoridae:
Castor canadensis Kuhl.
Cricetidae:
Peromyscus cf. leucopus (Rafin-
esque).

{+ Neotoma magister Baird.
tParahodemys spelaeus Gidley and
Gazin.
Synaptomys cf. cooperi Baird.
Synaptomys (Mictomys) sp.
+Microtus (or Pitymys?) cf. in-
volutus (Cope).
tOndatra cf. annectens (Brown).
Zapodidae:
Zapus sp.
Napaeozapus ef. insignis (Miller).
Erethizontidae:
Erethizon ef. dorsatum (Linnaeus).

LAGOMQRPHA:

Ochotonidae:
Ochotona sp.
Leporidae:
Lepus cf. americanus Erxleben.

PROBOSCIDEA:

Mastodontidae:
{Mammut cf. americanum (Kerr).

and | PerrssoDACTYLA:

Equidae:
{Equus sp.

Tapiridae:
{Tapirus sp.

ARTIODACTYLA:

Tayassuidae:
tPlatygonus cumberlandensis Gid-
ley.
tPlatygonus vetus(?) Leidy.
t{Mylohyus exorlivus Gidley.
tMylohyus ef. pennsylvanicus
(Leidy).
Cervidae:
Cervus sp.
Odocoileus ef. virginianus (Bod-
daert).
Bovidae:
tEuceratherium(?) americanum

(Gidley).

6 BULLETIN 171, UNITED STATES NATIONAL MUSEUM
ENVIRONMENT OF THE FAUNA

Evidence of the environmental conditions- that prevailed ir the
region of the cave during the time in which the fossil material accumu-
lated might reasonably be expected from a consideration of the various
forms present in the fauna. Many of the species recognized are
closely related to living forms, and it seems probable that their habits
were not greatly different from those of living representatives.

The fauna from the cave includes a peculiar assemblage of animals.
Many of the species are comparable to forms now living in the vicinity
of the cave, but others are distinctly northern, or Boreal, in their
affinities and some are related to species peculiar to the southern, or
Lower Austral, region. The fauna now living in this part of Maryland
is composed principally of species representative of the Carolinian or
Upper Austral region and of the Alleghenian division of the Transition
Zone (Merriam, 1900, p. 292).

The wolverine, Gulo, and lemmings of the group Mictomys are
distinctly northern in range, and it seems highly improbable that they
were contemporaneous in the same region with a crocodylid and a
tapir. Such an association might be plausible with sufficient topo-
graphic relief, but from a consideration of the physiographic history
it seems evident that during the time in which the cave deposit
accumulated relief was not greater than at present, as streams may
not have cut so deeply below the old upland surfaces. A more
probable explanation of the apparent association is that the fauna is
not entirely contemporaneous and that the entombment extended
over a period of time sufficient to allow important climatic changes to
take place. The cave may have received material during a portion
of a glacial and of an interglacial stage. Gidley (1920a) contended
that the forms were contemporaneous in the same region, postulating
greater relief, but the physiographic evidence does not seem to
justify his reasoning.

Included among the forms that may well have been associated
with wolverine and lemming mice in the colder fauna are the long-
tailed shrew, fisher, mink, red squirrel, muskrat, porcupine, jumping
mice, pika, hare, and elk; although in present distribution several of
these extend well into the southern region. Suggesting warmer
climatic conditions are bats and peccaries, in addition to crocodylid
and tapir. Among those forms whose most closely related living
representatives are now found much farther west are coyote, badger,
pika, a pumalike cat, and a pocket gopher that resembles Thomomys
more closely than it does Geomys.

The fauna as a whole is strongly indicative of a wooded region with
no lack of moisture. The floristic condition is suggested particularly
well by the spermophiles and jumping mice, and the plentiful supply

P71 Pe AtiE A

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PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE a

of water is evident from the presence of fisher, mink, otter, beaver,
bog-lemmings, and muskrat, as well as of the crocodylid. The greater
part of the fauna is not unsuited to humid conditions and an abun-
dance of vegetation. However, the horse, coyote, badger, and hare
are suggestive of a more open terrane, perhaps bordering a wooded
area. It should be noted, moreover, that coyotes and perhaps
badgers are commoner in arid regions.

AGE AND RELATIONSHIPS OF THE FAUNA

A Pleistocene age seems clearly indicated by the assemblage, but a
more critical recognition of the stage represented in the absence of
stratigraphic evidence is not entirely satisfactory. Our knowledge of
the sequence of mammalian forms during Pleistocene time is very in-
complete, and it is only through direct correlation of fossil occurrences
with known glacial and interglacial deposits that the succession of
faunas can be determined. The appearance of new migrants and the
extinction of old types appear as valuable criteria in recognizing
various stages, but in any case the absence of forms can never be
regarded as more than negative evidence.

The number of extinct forms recognized in the fauna and the diffi-
culties encountered in identifying as modern several of the species so
classified do not favor a very late stage at Cumberland Cave. Many
typical Pleistocene forms, however, as saber-toothed cais, sloth,
camel, elephant, and extinct species of bison and musk-ox, are not
included in the assemblage. Here again the absence of types is not
positive evidence. It is a noteworthy fact that remains of animals
larger than a black bear are exceedingly scarce in the collection,
probably owing to the mode of accumulation and possibly m part to
regional environment.

Gidley (1913b, p. 95) regarded the assemblage as pre-Wisconsin in
age and possibly as old as that from Port Kennedy (Cope, 1899) m
Pennsylvania. A comparison of the Cumberland Cave faunal list
with that of the Port Kennedy fauna, and with the list of mammals
from the Conard Fissure in Arkansas (Brown, 1908), follows.

8 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

CUMBERLAND OAVE

Sorez sp.

Blarina brevicauda
Eptesicus cf. grandis

Corynorhinus alleganiensis
Canis armbrusteri
Canis cf. priscolatrans

Euarctos vitabilis
Arctodus haplodon
Martes parapennantt
Mustela cf. vison

Gulo gidleyi
Intra parvicuspis
Brachyprotoma pristina

Spilogale marylandensis

Taridea marylandica

Felis cf. inerpectata
Felis near atror

Marmota monar

Citellus ef. tridecemlineatus
Tamias cf. striatus

Sciurus tenuidens

Glaucomys sp.
Plesiothomomys potomacensis
Castor canadensis

Peromyscus cf. leucopus
Neotoma magister
Parahodomys spelaeus
Synaptomys cf. cooperi
Synaptomys (Mictomys) sp.

Microtus (or Pitymys?) cf. in-

volutus
Ondatra cf. annectens
Zapus sp.
Napaeozapus cf. insignis
Erethizon cf. dorsatum
Lepus cf. americanus

Ochotona sp.
Mammut cf. americanum

Port KENNEDY

Scalopus sp.

Blarina simplicidens
Myotis(?) sp.!

Canis dirus(?)

Canis priscolatrans
Vulpes latidentatus
Urocyon cinereoargenteus

Euarctos americanus
Arctodus haplodon
Martes diluviana

Gulo gidleyi

Tutra rhoadsit
EBrachyprotoma obtusata
(?) Mephitis nigra

Osmotherium spelaewm
Pelycictis lobulatus
Tazidca tarus
Smilodontopsis gracilis
Smilodontopsis mercerii
Felis inerpectata

Felis eyra

Lynx calcaratus

Sciurus calycinus

Castor canadensis

Peromyscus leucopus(?)!

Anaptogonia hiatidens

Sycium cloacinum

Microtus diluvianus

Microtus speothen

Microtus (or Pitymys?) dideltus
Microtus (or Pitymys?) involutus

Zapus hudsonius(?)
Erethizon dorsatum(?)
Sylvilagus floridanus

Ochotona(?) palatina
Mammut americanum

' According to O. P. Hay (1923, p. 312).

CONARD FIssuRE

Scalopus aquaticus

Sorex personatus

Sorex personatus fossidens
Sorex obscurus

Sorex fumeus

Microsorer minutus

Blarina brevicauda ozarkensis
Eptesicus grandis

Myotis subulatus(?)

Canis occidentalis(?)

Vulpes fulva(?)
Urocyon(?) sp.
Procyon lotor
Euarctos americanus

Martes pennanti

Mustela vison

Mustela cicognanti angustidens
Mustela gracilis

Brachyprotoma pristina
Mephitis mesomelas newtonensis
Spilogale interrupta(?)

Smilodontopsis troglodytes
Smilodontopsis conardt
Felis couguar

Felis longicrus

Lynx compressus

Lynz rufus(?)

Marmota monar

Citellus tridecemlineatus(?)
Tamias nasutus

Sciurus hudsonicus

Geomys parvidens

Castor canadensis
Reithrodontomys simplicidens
Peromyscus sp.

Neotoma czarkensis

Microtus ochrogaster

Ondatra annectens

Erethizon dorsatum
Lepus americanus(?)
Lepus giganteus
Sylvilagus floridanus(?)

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 9

CUMBERLAND CAvE—Con. Port KENNEDY—Con. CONARD FissurE—Con.

Equus sp. Equus complicatus ! Equus scotti(?)
Equus pectinatus |

Tapirus sp. Tapirus haysii
Platygonus cumberlandensis Platygonus(?) tetragonus
Platygonus vetus(?)
Mylohyus exortivus Mylohyus nasutus Mylohyus browni
Mylohyus cf. pennsylvanicus Mylohyus pennsylvanicus Mylohyus sp. indet.

Mylohyus sp. indet.
Teleopternus orientalis

Cervus sp. Cervus canadensis(?)
Odocoileus laevicornis Odocoileus hemionus(?)

Odocoileus cf. virginianus Odocoileus virginianus Odocoiletis virginianus(?)
Bison sp.!

Buceratherium (?) americanum
Symbos australis
Megalonyx lorodon
Megalonyr wheatleyi
Megalonyx tortulus
Megalonyz scalper
Mylodon harlani(?)

The Cumberland Cave fauna exceeds that of other caves in number
of mammalian genera recorded but includes a slightly smaller number
of species than recognized in either the Port Kennedy or Conard
Fissure occurrence. Of the 41 genera listed from Cumberland Cave 8
are extinct, and of 46 species about 28 are believed to represent extinct
forms. Thirteen of the 39 genera and about 37 of the 51 species in the
Port Kennedy fauna as listed are extinct. This suggests a somewhat
greater antiquity for the Pennsylvania assemblage. Thirty-eicht
genera are recorded from the Conard Fissure, and of this number 4
are extinct, and about 19 of the 50 species are extinct. The Conard
Fissure fauna may be appreciably younger than either of the other
two, although O. P. Hay (1923, p. 14) believed that the Arkansas
occurrence antedated Cumberland Cave. Saber-toothed cats were
found in both Port Kennedy and Conard Fissure deposits and not in
Cumberland Cave, but tapir, mastodon, and Arctodus included in the
Cumberland Cave and Port Kennedy occurrences are absent from
Conard Fissure. It seems odd that sloth remains were not found in
either Conard Fissure or Cumberland Cave. Megalonichid material
is frequently encountered in caves, and species of this group of sloths
probably lived until near the close of Pleistocene time.

If the Port Kennedy occurrence is considered as early Pleistocene,
that of Cumberland Cave may well be near middle Pleistocene in
age. However, the particular stage or stages represented remains
undetermined.

1 According to O. P. Hay (1923, p. 312).

10 BULLETIN 171, UNITED STATES NATIONAL MUSEUM
SYSTEMATIC DESCRIPTION OF VERTEBRATE FAUNA

Order LORICATA
Crocodylid

A single reptile tooth in the Cumberland Cave collection was recog-
nized by C. W. Gilmore as belonging to a crocodile or an alligator.
The importance of this tooth is seen in its occurrence farther north
than the present limits of this group, suggesting a more equable
climate than at present.

Order SERPENTES
Ophidian remains

Several small vertebrae in the collection were identified by Gilmore
as snake. There is no certain evidence in the material that more than
one form is represented.

Order GALLIFORMES
Family TETRAONIDAE

BONASA UMBELLUS (Linnaeus)

A bird bone (U.S.N.M. no. 11690) in the collection was identified
by Dr. Wetmore (1927) as the distal portion of a left humerus belong-
ing to a ruffed grouse.

Order INSECTIVORA
Family SORICIDAE

SOREX species

A single maxillary fragment (U.S.N.M. no. 12468) including three
teeth, apparently P*, M', and M’, was kindly identified by Dr. H.H. T.
Jackson as belonging to the long-tailed shrew, Sorex. The specimen
is too fragmentary to permit specific recognition, but no differences
could be cited to distinguish it from the living species Sores: cinereus,
which inhabits the region of the cave today.

BLARINA BREVICAUDA (Say)

Representing the short-tailed shrew are one rostral portion of a
skull with the greater part of the dentition, two maxillary fragments
with teeth, and eight lower jaws. The fossil material indicates a size
somewhat greater than the average in Recent specimens and in several
characters resembles the form described by Brown from the Conard
Fissure as Blarina b. ozarkensis (Brown, 1908, pp. 170-171, pl. 15).
The heel on the last lower molar is reduced as in the Arkansas form
but not more so than in some modern individuals of B. brevicauda.
The presence of this heel clearly separates the Cumberland form

U. S. NATIONAL MUSEUM

BULLETIN

171

PLATE 6

hotographed from near opening, showing bones sticking in clay and stalactitic material.

cave interior p

Small portion of ¢

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BULLETIN 171

U. S. NATIONAL MUSEUM

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PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE Si

from B. simplicidens Cope (1899, pp. 219-220). The longitudinal
profile of the lower border of the mandible varies in the series of Jaws
from a smooth curve, as in the Conard Fissure specimen figured by
Brown, to a margin distinctly angulate below M3. The variation,
however, does not appear to be greater than in living Llarina brevi-
cauda, although modern specimens show a greater tendency toward a
sharper angulation below M3. Furthermore, there is an average
difference in the form of the large lower incisor, which in the fossil
shrew is less turned upward at the point than is usual in living species.
As noticed by Brown, the crowns of the teeth show pigment as in more
recent specimens.

Order CHIROPTERA
Family VESPERTILIONIDAE

EPTESICUS cf. GRANDIS (Brewn)?

The large vespertilionid E’ptesicus is represented in the collection by
13 skull portions and 55 mandibular rami. About half the lower jaw
specimens are exceptionally well preserved, and one pair of rami was
associated with a nearly complete skull.

The two best-preserved Cumberland Cave skulls (U.S.N.M.
nos. 12432 and 12433) are distinctly larger than the living bat Eplesicus
fuscus, in which respect they resemble E. f. grandis from the Conard
Fissure. Brown considered his form a subspecies of #. fuscus, but the
characters outlined by him suggest that the form might well deserve
full specific rank. Furthermore, if the Cumberland Cave form is
conspecific with the Arkansas Pleistocene type, as seems evident,
additional characters are to be seen in the crania of the better-preserved
skulls from Maryland.

As in E. grandis, the Cumberland Cave form shows the following
differences from £. fuscus: Skull larger, ridges and depressions on the
dorsal surface of rostrum usually more pronounced, width of zygoma
greater, teeth somewhat larger than average, depth of masseteric
fossa slightly greater, condyle on mandible relatively larger, antero-
internal basal cusp on P, better developed, and mandible of greater
relative depth. Additional characters not determinable in the Conard
Fissure material but present in the Cumberland specimens include a
wider brain case, greater separation of the occipital condyles, and
slightly larger bullae. The only difference observed between the
specimens from the two localities is that the upper canine in the
Maryland form is slightly smaller. However, the number of speci-
mens in which this tooth is preserved is very limited in both collec-
tions, and the canine in recent individuals shows a noticeable variation
in size.

21 Brown, 1908, pp. 174-175, pl. 15.
16541—38——-2

12 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

TABLE 1.—Measurements (in millimeters) of skulls and mandibles of Eptesicus

grandis and E. fuscus

Eptesicus Eptesicus Eptesicus
ef. grandis grandis fuscus '
A.M.N.H. U.S.N.M.
U.S.N.M. no. 11795 no. 255580
no. 12432 (type) (Div. Mamm.)
Distance from anterior surface of canine to occipital condyle - 18.:4)./eeon 2s eee 18.2
Width of postorbital constriction---.-..-------------------- 4.3 4.3 4.6
Width) of: brain’ casez= =5 = teen ee eee ee eee 94a aes eee 8.5
Distance across occipital condyles....--....---------------- CAN hata re eee eee 5.8
Distance across zygomatic arches. ....----.--....-=-------- TSN lea e ee see oe 12.9
U.S.N.M.
no. 12438
Length of upper dentition, P4 to M3__..............------- 6.1 26.1 5.9
U.S.N.M.
no. 12447
Length of lower jaw from li to condyle--.-.-.-------------- 15,2 15; 2 15.0

Length of lower dentition from C to M3-__-.-.-.------------ 8.3 8.3 8.2

1 Specimen of greater than average size.
2 Approximate.

CORYNORHINUS ALLEGANIENSIS Gidley and Gazin!

Figure 1

Type —Skull, U.S.N.M. no. 12412, with incomplete dentition and
lacking bullae and zygomatic arches.

Specific characters —Skull close in size to that of C. rafinesquia.
Frontal region not so highly inflated above dorsal plane of rostrum.
Median depression in the dorsal surface of muzzle not so deep. Depth
of posterior portion of skull from occipital condyles to top of inter-
parietal less. Occipital condyles slightly farther forward. Temporal
ridges do not unite posteriorly in a median ridge.

Material—Representing Corynorhinus alleganiensis are 5 skull
portions and 29 lower jaw fragments. The most complete specimen
is the type, which is well preserved and clearly shows the proportions
of the rostrum and cranium (fig. la and 6). However, the dentition
is not entire and the specimen lacks the zygomatic arches and tym-
panic bullae. U.S.N.M. no 12413 is an incomplete rostral portion
including all the cheek teeth posterior to the canine in the right
maxillary (fig. 1c) and three cheek teeth in the left. A fragmentary
third skull, U.S.N.M. no 12415, consists of the left portion of the
rostrum with the teeth, P* to M’, and a part of the frontal region.
The two remaining skull portions consist only of maxillary fragments
and teeth. The majority of mandibles are remarkably well pre-
served and include a greater part of the dentition (see fig. 1d), although
only two specimens retain the incisors.

Description —The skull of Corynorhinus alleganiensis is about the
size of that in the living species C. macrotis and C. rafinesquir. How-
ever, several characters are observed that clearly separate it from the

8 Gidley and Gazin, 1933, pp. 345-347, fig. 1.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 13

nearest living forms. The most important of these are concerned
with the less inflated condition of the brain case and appear not to be
due to crushing. The frontal region in C. alleganiensis does not pro-
ject so markedly above the dorsal plane of the rostrum; consequently
the dorsal profile from the nasal opening to the top of the frontals
more nearly approaches a straight line. Also, the median pit or
depression in the dorsal surface of the rostrum is not so deep as in
CO. rafinesquu and is not separated from the anterior nasal notch by
so conspicuous a ridge. The height of the posterior portion of the
cranium as measured from the occipital condyles to the top of the
interparietal is noticeably less in the fossil. Furthermore, the con-
dyles are slightly farther forward in position as indicated by the dis-
tance between the preserved condyle on the left side and the glenoid
fossa. This suggests that the bullae, though not preserved in the
fossil, may have been of smaller anteroposterior diameter or possibly
not so well separated from the condyles. The temporal ridges at the
anterodorsal margins of the temporal fossae are distinct and converge
backward but instead of uniting to form a sagittal crest continue
posteriorly a short distance as nearly parallel ridges. This character
was also clearly observed in a second, more incomplete skull portion of
CO. alleganiensis (no. 12415). In all specimens of C. macrotis and C.
rafinesquii examined these lateral ridges unite posteriorly to form a
short median ridge. Furthermore, these ridges in C. alleganiensis
are somewhat better defined anteriorly than in modern forms and at
the termination of each is the very slightest suggestion of a postorbital
process.

Corynorhinus phyllotis Allen (1916, pp. 352-353) from the region of
San Luis Potosi in Mexico, though defined as having a flattened
brain case, is apparently a larger and broader-skulled type than
C. alleganiensis. The rostrum of C. phyllotis is described as more
depressed and having a more marked median excavation than other
living species of Corynorhinus. Furthermore, the audital bullae are
even larger than in the more northern species.

No important or consistent differences were observed between the
dentition of C. allegamiensis and that of the living forms of Coryno-
rhinus. Also, the lower jaws are apparently not distinguishable from
those of modern forms. Although the dental formula for the lower
jaw of Corynorhinus is the same as in Myotis, not one of the many
specimens from Cumberland Cave was found to represent the latter
genus. It is probable that Gidley had reference to this Corynorhinus
lower jaw material when he included Myotis in his faunal list of 1913.
The mandible of Corynorhinus has a relatively shorter premolar series
and longer molar series than does that of Myotis.

The presence of Corynorhinus in the fauna is of particular interest
inasmuch as it is the first record of the genus from the Pleistocene and

14 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

it adds a new and interesting species to this group of bats. Further-
more, its occurrence in western Maryland is apparently somewhat
outside the recorded distribution (Allen, 1916, pp. 338, 349-350) of the
living species. The range of C. macrotis extends northward into
North Carolina, and the species has been recorded from the Dismal
Swamp of Virginia. C. rafinesquii is known in southwestern Virginia,
Kentucky, and southern Indiana, as well as farther west.

FIGURE 1.—Corynorhinus alleganiensis Gidley and Gazin: a, Skuil, type specimen (U.S.N.M. no. 12412),
dorsal view; 6, same, lateral view; c, upper dentition, paratype (U.S.N.M. no. 12413), occlusal view;
d, left ramus of mandible, paratype (U.S.N.M. no. 12414), lateral view. X< 3. Cumberland Cave Pleisto-
cene, Maryland.

TABLE 2.— Measurements (in millimeters) of skulls and mandibles of Corynorhinus
alleganiensis and C. rafinesquii pallescens

to Be Corunorhinus
Measurement ununor hints rafinesquii
g a pallescens
U.S.N.M. no.
U.S.N.M. no.| 213367 (Biol.
12412 (type) Surv.)
Length of skull from anterior margin of canine alveolus to supraoccipital____- 15.75 15.9
Widthtofpostorbitsl! constrictions 9-2. ees ee ee 3.8 3. 75
Depth of posterior portion of skull from inferior margin of occipital condyles
to top’ of interparietal...-. =.=)... bet Tee TERE eee 5. 25 6.3
Distance from postglenoid process to posterior margin of occipital condyle _ _ 5. 65 6. 4
U.S.N.M. no.
12418 |
Length of upper premolar-molar series___.....--__._.___.---_---_-________- 4.4 | 4.5
U.S.N.M. no.
12414
Length of lower dentition from C to Ma, inclusive.___..______........____ 5.9 | 5.8
|

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE = 15
Order CARNIVORA
Family CANIDAE

CANIS ARMBRUSTERI Gidley ¢

Figures 2-10

The type of this species is a portion of a left lower jaw, U.S.N.M.
no. 7662, containing teeth, P, to My, and the alveolus for M, (fig. 6a
and 6). Two paratypes were designated: A portion of a right lower
jaw with teeth, P, to M,, U.S.N.M. no. 7661 (fig. 7a and 6), and por-
tions of the right and left lower jaws of another individual, U.S.N.M.
no. 7482, containing teeth, which include carnassials of both sides and
M, and the posterior half of P, of the left side.

The material in the National Museum representing this species
consists of a number of well-preserved specimens (figs. 2-10), includ-
ing nine skull portions of which five are more than half complete.
There are also 17 lower jaws and jaw portions, one nearly complete
fore limb and foot, and several unassociated limb bones and vertebrae.
Most of this material was collected after Gidley published his deserip-
tion of this form.

The chief characters given by Gidley to distinguish Canis arm-
brusteri are as follows:

Size slightly less than that of C. occidentalis, as that species has been defined by
Miller [1912, p. 2], but tooth characters indicate an animal quite distinct from
any of the true wolves. Its principal differences are seen in the greater relative
depth of jaw, smaller canine, more simple p2 and ps3, the presence of a posterior basal
tubercle on p;, and in the relatively larger heel of the carnassial. The paraconid
also is less expanded at base, with more perpendicular anterior face. The meta-
conid is larger and higher placed, while the protoconid is less broad and full, as
seen from the inner side. The carnassials as a whole suggest those of a jackal, fox,
or coyote rather than those ofa wolf. * * *

In the newer material is a portion of a lower jaw associated with a
broken and crushed skull, U.S.N.M. no. 11881, in which most of the
cheek teeth are preserved. The very close resemblance of the lower
teeth in this specimen (fig. 8) with the corresponding ones of the type
make its reference to C. armbrusteri logical. However, the depth of
the ramus in no. 11881 is not so great. By comparison with no. 11881,
other skull portions in the collection are referred to this species.

Considerable individual variation exists in the series of canid spec-
imens, sufficient to suggest the recognition of more than one distinct
species were only the extremes represented. However, the degree of
variation, particularly in size and proportions of skull and teeth, is
not greater than that which was observed in a series of 35 Recent
canid skulls in the collections of the Biological Survey from Gila Na-
tional Forest in New Mexico.

4 Gidley, 1913b, pp. 98-102.

16 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

The skull of Canis armbrusteri as indicated by specimens no. 11886
(fig. 2a and 5), no. 11883 (fig. 3a, b, and c), and no. 11885 (fig. 4a, 6,
and c) approximates in size that of Canis occidentalis. The rostrum
is slender, but the frontals are relatively broad and heavy. The
sagittal crest, though variable, is more prominent in no. 11886 and
no. 11883 than in old individuals of C. occidentalis. In no. 11886 the
inion projects backward to a marked degree, but not nearly so much as
in the Rancho La Brea wolves.

The superior first three premolars are comparatively simple and
slender. The anterointernal portion of the posterior root of P? and
P’ does not project lingually so much as in C. occidentalis. The upper
carnassial and molar teeth are relatively large and massive.

TaBLe 3.—Measurements (in millimeters) of skulls and superior dentition of
Canis armbrusteri

U5). | UsS: US. USS. U.S. | U.S.
N.M. | N.M. | N.M. | N.M. | N.M. | N.M.

Meusurement no. no. no. no. no. no.
11886 11881 11883 11885 11887 7994
Skull
Length from anterior end of premaxillary to condyles_ | V270 5 Bes eee Ne aS PAO 2 a kee oa See ee
Length from anterior end of premaxillary to posterior
Nasalionening = 4.22.2 8 lees Se ee 140; ae Eee 1130 a oe es ee ee
Length from glenoid fossae to condyles.__....--.--.--- 55) cbaes aces eases 53. 5 47 45.5
Distance from line between postorbital processes to

NOT Sh Se rae ee ee ee a ee eat ae L3G [ese csoee | tener 1120 1105 100
Width of muzzle in front of infraorbital foramina...--| 151 j_._-__-- 147 eg Ate ll | a a A ls 44.2

Least width between superior borders of orbits_..._--| 156 |__------ 52.6 44.7 44.1 42
Width across postorbital processes of frontals_-__------ 167.7 | 187 167 61.5 53.9 59.6
Greatest width across zygomatic arches__......-..--.- DAGO. Fes 8225 1150 L130) 0 eee eee
Width of palate between alveoli of M!__.------.-__-_- Page pe ea ae 144.5 39 38. 6 36. 5

Supertor dentition |

Length of cheek-tooth series, P! to M2, inclusive_.__._| 1101 1101.5 96 06:(5) |2enescnetoeee aces
C, anteroposterior diameter at base of enamel_________|..---___|-------- W3s3i [2s oe ee | SS
O, transverse diameter at base of enamel_____.-...-.-_|_-------]_------- Oust sc ae SS aeeee | Le
PA Vanteroposterior Gia Meterse se 5 ele ee ee ee ee | eee 8.4 Sf 6c lec accres eee
Pl greatest transverse dlameters:c: 29. 45-5512 2293 oe | ee 5.5 4: Si \weres [poe see
Pi anteroposterior diaméter-.--.-.---- 2-5 - e |S P44 cc oe |S ee eee ere
PS greatest: transverse diameter. s2- 2 =-2 ose 5 eee DSS) | Sear ee Peet oN eee se eee eos cceae
ips; anteroposterior diameters] =: 2.) = )* ese a ee eee 17. 4)) lara ites 1a 6 jes
P3-oreatest. transverse diameter... - 5-2 ses eae ee 6.9 Ce ols ccet ee eee ele ee
P4, anteroposterior diameter____....__-__-_-_- pane eee St 28.8 28.3 28.1} 129.4 26.7
P4, greatest transverse diameter including protocone-- 14.9 14.5 13.6 14.5 | 113.2 12.9
P4, transverse diameter across paracone__.-._._.------ 11.5 12 11.9 12 11.9 11.5
M!, anteroposterior diameter parallel to outer wall. __- 18.1 18 18, 2 18 19.1 17.2
M1, transverse diameter perpendicular to outer wall__| 21.8 22.5 22.7 21.7 23. 4 20.9
M1, greatest transverse diameter__...._.----.-.------- 24.4 25. 4 24.9 24.5 25. 5 23.4
M2, anteroposterior diameter-___....-.___.-.----_----- 10.3 10.3 O50. eae 10.4 8.3
M?, greatest transverse diameter_.._..__.-.-.--.-_---- 15.7 15 1557 pss dees 17.5 14,2

1 Approximate.

Two isolated upper molars of Canis armbrusteri were described
recently by Bryan Patterson (1932). He noted the large size of these

teeth and the relatively large M? as compared with M!. The Field
Museum M! possesses a well-developed hypocone, as do several of the
National Museum specimens, but, as anticipated by Patterson, this
character is variable and in some dentitions where the heel of M! is
narrower the hypocone is not so prominent. However, the reduction
of the hypocone in these cases does not reach the stage common in C.
dirus of Rancho La Brea.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE

TaBLE 4.—Measurements (in millimeters) of mandible and inferior dentition of

Canis armbrusteri

woe | sh | Wa, | | NS | NS

Measurement wOB? no. no. no. no. no.

(type) 11881 8168 8169 11887 8172

Mandible |

Depth of jaw below anterior part of P3_.....---------- eee 27 dea eaeSe 25. 5 21.4

Depth of jaw below heel of M.----..----------------- 36.3 32. 6 31 31.3 29 125
Thickness of jaw below heel of Mj-------------------- | 16.3 15.7 15.6 15 14.6 12.9

Inferior dentition
Length of tooth series from anterior margin of canine

alveolus to posterior margin of Ma-_-----------------|-------- 2127 S130 hee [a eee | pees 1106

C, anteroposterior diameter at base of enamel-_--------|-~------ WIGS6: | BA: - oes Soe 142

C, transverse diameter at base of enamel_-__---.-------|-------- 141.3 10) = Ses cca leoseea 13
Pi anteroposterior diameter-. 22. 2-=2==-5- 22s ee | eee 7 6 Gi eae ea etree eee ese
Pi sereatest transverse diameter..--.---------—22-- eae] a nee 5.2 4587 eee aioe aioe oe
Ps anteroposterior diaiielerae se-=- <2 <5 ae eee oa eee 13.8 loses sae 13.9 12.4
Po, greatest trans verse) diameter-_.-_...--.----225--.2_}_-. <= --|_- =e 6:4 | esses 5.4 5.6
Ps,-anteroposterior Ginmeter=—2--+-------- oe a— ne eee |e=—onn 15.4 15.8 16 14.6 13.4
Es, greatest. transverse: diamoeter--...-=-..-.s2-2n22-2a|P oo 6.7 6.9 TAS eauaee 5.8

Pi; anteroposterior diameter222--=._... 2-22-28 17.9 17.7 17.8} 118 18.3 16
P,, greatest transverse diameter__-----.--------------- 9.1 9.2 8.8 19.5 9.6 Ponti
My, anteroposterior diameter--_------------------------ 31 31 131 1294 eae 27.9
Mi, greatest transverse width of trigonid__---.-.-.---- 12.5 13.2 12.1 122) Soke 10.6
M1), greatest transverse width of talonid---.-.-------- 12.5 PAO LOS eel ae ey te ae eee oe
Mo, anteroposterior diameter--.-_---------------------- 13.6 13.3 | 113.5 LSS Sl oes 12.4
Ms, greatest transverse diameter-_--.-.-.-------------- 10. 4 9.8 19,9} 110 10. 2 8.8
Mawanteropostenion Giame tener —- == ee a a ee ee a nee ee 7.2 6:5 }=22-82 5
Mis greatest transverse diameter--------=--------s2-=5|aeaa-n |= - =n en |= —-— = 5.6 PS Sele

1 Approximate.

C. (Aenocyon) dirus is distinctly more robust than C. armbrusteri.

The specimens from Rancho La Brea have a much wider rostrum and
the carnassial teeth are more powerfully built. The larger limb bones
in the Cumberland Cave collection, presumably referable to C. arm-
brusteri, are equally as long but slenderer than corresponding elements
of various individuals of the Brea wolf in the National Museum col-
lections.

The coyotelike characters in the dentition of C. armbrusteri pointed
out by Gidley do not necessarily require that this form be grouped
with the coyotes in the subgenus Lyciscus or Thos but emphasize the
specific separation of C. armbrusteri from the living wolves of North

18 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

‘SS Ci SU
Dy: 2

——
SS
Saag

Seats sce
SSS =
SS

FIGURE 2.—Canis armbrusteri Gidley, skull (U.S.N.M. no. 11886): a, Lateral view; 6, ventral view of left half of palate.

Cumberland Cave Pleistocene, Maryland.

Two-thirds natural size.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE

‘pUBLAIBIA ‘OUGD0ISI9[qg OALO PUL[EQUIND ‘02/5 [BINIVU SPATYI-OMJ, “[EISSBUIBO IOAOT JO SAOIA [BSNJOO PUL [BI10qRT '9
‘oynyed Jo Jey IO JO MOTA [BIJUOA ‘q ‘[[MAS JO MTA [B1OYB'T ‘D :(ESRIT “OU “IN N'S'Q) [BIsseurEO Jomo[ pue [[Nys ‘AO[PID Mapsniguisy suDQ—'g AMAA

; ;
Z re SS
oS . 2 Te y:

> = = Sp
Wp, Uy, =— > --, ay =
qin,” St EO OS
Gi

SEN) qj <<
oe ME /) I
‘ (ye ss WD KE. ~
fi a

ANd
U,

amy Wy 7
ee Or

HS Ae
[i tay = oe
Dy, Lies ese
nF

D 4
MIS ie f
— Westtanier=

C®) :

BULLETIN 171, UNITED STATES NATIONAL MUSEUM

, i) )

we Ry, Me
) SRDS

FIGURE 4.—Canis armbrusteri Gidley, skull (U.S.N.M. no. 11885): a, Dorsal view; 6, lateral view; ¢, ventral
view. Two-fifths natural size. Cumberland Cave Pleistocene, Maryland.

SS y, >
Lh

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 21

FIGURE 5.—Canis armbrusteri Gidley, skull (U.S.N.M. no. 7994): a, Lateral view; 6, ventral view.
One-half natural size. Cumberland Cave Pleistocene, Maryland.

FIGURE 6.—Canis armbrusteri Gidley, left ramus of mandible, type specimen (U.S.N.M. no. 7662):
a, Lateral view, two-thirds natural size; 6, occlusal view, natural size. Cumberland Cave Pleistocene,
Maryland.

22 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Si Dui : j Ly;

FIGURE 7.—Canis armbrusteri Gidley, right ramus of mandible, paratype (U.S.N.M. no. 7661): a, Lateral
view, two-thirds natural size; b, occlusal view, natural size. Cumberland Cave Pleistocene, Maryland.

FIGURE 8.—Canis armbrusteri Gidiey: Left ramus of mandible (U.S.N.M. no. 11881), lateral view.
Two-thirds natural size. Cumberland Cave Pleistocene, Maryland.

FIGURE 9.—Canis armbrusteri Gidley: Right ramus of mandible (U.S.N.M. no. 11882), lateral view.
One-half natural size. Cumberland Cave Pleistocene, Maryland.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 23.

America. In proportions of the skull and lower jaw the fossil form is
distinctly more wolflike than C. rufus (=C. frustror), which also shows
points of resemblance to the coyotes.

-
SS
one.
7 X
2S
<\
\
“ va
1 s 1
N x
ee ut
Ape eetn Ne
Be SENG ‘
CRY ee
AS \ Y
Tre. \
4 we \
\ \
4 \

\, ~~
‘.

"S ook ee
x =~
Sere eee or
Sere :

Ficuke 10.—Canis armbrusteri Gidley: Right ramus of mandible (U.S.N.M. no. 8172), lateral and occlusal
views. One-half natural size. Cumberland Cave Pleistocene, Maryland.

CANIS cf. PRISCOLATRANS Cope

A skull fragment with part of P® and P* of the right side and the
root portions of M' and M? on the left shows the presence in the
fauna of a coyote near Canis latrans. The anterior portion and right
side of the muzzle are incomplete, but the cranium and left zygoma
are well preserved. The skull is very close in size to that of C. latrans
but is slightly more robust, particularly in the region of the frontals.
The small, very slender, sharp-cusped character of P? and the frag-
ment of P* further substantiates reference of the form to the coyote
group. Direct comparison with C. priscolatrans from the Port Ken-
nedy deposit in Pennsylvania or with C. riviveronis Hay from Florida
is not satisfactory, because of the different nature of the materials.
The reference of the Cumberland Cave form to C. priscolatrans is
based on the somewhat more robust character of the Maryland
form as compared with living coyotes.

Measurements of the skull fragment (U.S.N.M. no. 7660) are as
follows: Distance from line between postorbital processes of frontals
to inion, 88 mm; least width between anterior margins of orbits,
33.2; width across postorbital processes of frontals, 51 mm; greatest
width across zygomatic arches, 110 mm (last three measurements
approximate).

Family URSIDAE

EUARCTOS VITABILIS (Gidley)
Figures 11-14; Puiatr 8

The Cumberland Cave collection includes an unusually large
amount of well-preserved bear material. There are skull portions of
at least 25 individuals. Four of the skulls are nearly complete
(figs. 12-14), and nine more are represented by rostral portions

24 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

including most of the teeth on both sides. The remainder consists of
maxillary fragments with teeth. About 35 individuals are indicated
by lower jaws. Six specimens include both rami and most of the
teeth. A large portion of the separate jaws are well preserved and
have nearly all the teeth. In two cases jaws and skull were associated.

A composite skeleton (see pl. 8) has been mounted from the better-
preserved remains and placed on exhibition at the National Museum.
This mount is composed of parts belonging to at least six individuals,
but a small portion of the skeleton was necessarily restored. The
scapulae, left ulna, fibulae, ribs, sternum, and some of the vertebrae
were not represented in the collection. The composite skeleton has
received the catalog number 10304, and is composed of the following
catalog portions: Skull, no. 8188; mandible, no. 10302; three posterior
dorsal and three anterior lumbar vertebrae, no. 10303; left radius,
no. 8126; parts of both hind limbs and right hind foot, no. 8179;
and complete left hind foot, no. 10301.

Ursus (Huarctos) vitabilis was described by Gidley (1913b, pp. 96-97)
from a nearly complete pair of lower jaws (fig. lla and b), U.S.N.M.
no. 7665. The description is as follows:

About the size of U. (Huarctos) americanus, but differs from that species in (1)
comparatively larger canines; (2) wider space between the anterior cheek-teeth,
combined with a less wide branching of the horizontal rami in general; (3) a
relatively larger symphysis, which is more sharply constricted and more flattened
latterly behind the canines; and (4) longer diastema between canines and cheek-
teeth.

Most of the ursid material in the collection was obtained since the
above description was published. A restudy of the group with this
added material at hand has brought to light new characters and has
furnished further information on the constancy or variability of
others. In view of these conclusions it has seemed advisable to
redescribe the species on the basis of characters shown by the type,
U.S.N.M. no. 7665, supplemented by a series of skull portions and
lower jaws.

Specific characters.—Size equaling that of the living black bear,
Euarctos americanus (Pallas). Upper and lower cheek teeth relatively
narrow. P* large and well developed, with prominent protocone.
M? large and usually does not narrow so abruptly posterior to the
metacone as does the corresponding tooth of £. americanus. Anterior
lower premolars more commonly present and better developed than
in EF. americanus. P, well developed.

All the Euarctos material in the collection is referred to E. vitabilis,
which indicates considerable variation in size for this species. Several
jaws in the collection compare favorably with those of small individuals
of EL. americanus or with the glacial bear, L. emmonsti. The largest
specimen of F. vitabilis, a skull, U.S.N.M. no. 12247, equals in size
large specimens of F. luteolus from Louisiana.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 25

The skull of ZL. vitabilis is not so robust as that of Ursus horribilis,
nor is the muzzle so elongate. In the relative proportions of the skull
and lower jaw the Cumberland Cave form compares favorably with
living black bears of North America. The shape as well as the size
of the fossil skulls varies somewhat, but no important or consistent
differences can be cited to establish a division within the fossil collec-
tion, or to distinguish them from Recent forms on the osseous parts
alone.

In the dentition EH. vitabilis shows points of resemblance to both
E. americanus and E. luteolus, but in some characters it seems clearly
distinguished from them. £. luteolus possesses teeth averaging some-
what greater size than does H. americanus. Also, in E. luteolus the
cheek teeth appear relatively wider and P* is better developed. The
upper and lower teeth of E. vitabilis are relatively much narrower
than in H. luteolus and average somewhat narrower than comparable
teeth in EL. americanus. Upper and lower canines in the fossil form
are usually large as in £. luteolus.

FP! in £. vitabilts is noticeably well developed, with a prominent
internal cusp placed opposite the notch between paracone and meta-
cone. This tooth is much larger than in /. americanus, and in several
fossil specimens it is appreciably larger than in LF. luteolus. The
corresponding tooth in U. horribilis is in turn very much larger than
in L. vitabilis, and in the grizzly the protocone is accompanied by
several accessory cuspules.

M!, other than being relatively somewhat narrower, differs little
from this tooth in EZ. americanus. M7? is an elongate tooth that does
not usually taper so abruptly posterior to the metacone as is common
in E. americanus; however, this tooth does not reach the development
and proportions seen in U. horribilis.

The anterior lower premolars are more commonly present in
E. vitabilis than in EL. americanus and E. luteolus and are usually better
developed. P* averages larger than in the living black bears, but it
is not nearly so large as in U. horribilis, nor does it show the sulcate
heel, which characterizes this tooth in the grizzly.

M'is a narrow tooth having a trigonid cuspate as in the ursids not
trenchant as in the tremarctine forms. The succeeding molars are
also narrow but otherwise resemble corresponding teeth in Recent
black bears.

A large amount of limb and foot material of Huarctos vitabilis is
included in the collection. The various elements for the most part
were unassociated. A noticeable variation in size is seen and no
consistent differences were observed which would serve to distinguish
the fossil from the living bear.

The skull of EZ. vitabilis differs from that of Ursus procerus Miller
(1899) from the Pleistocene of Ohio in having a less concave dorsal

26 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

surface of the rostrum and most noticeably in the smaller size of
the teeth.

E. vitabilis is also distinct from U. amplidens Leidy (1856) in having
lower teeth of much smaller size.

Upper teeth from the Conard Fissure referred by Brown (1908,
pp. 183-184) to Ursus americanus may represent Huarctos vitabilis.
The characters of the upper dentition cited by Brown are also seen
in the Cumberland Cave bear, as the development of P*, the narrow-
ness of M!, and the character of the heel of M?. In the past most finds
of Pleistocene black bear have been referred to the living species
Euarctos or Ursus americanus, and it seems probable that with a very
limited amount of material it would be difficult to recognize as dis-
tinct a species approaching the living form so closely as does EF.
vitabilis.

Tasie 5.— Measurements (in millimeters) of skull and superior dentition of three
species of Kuarctos

Euarctos | Fuarctos

Fuarcios vitabili 3
5 americanus| luteolus

Measurement s yee | i
M.no. | M. no. | Meno. | U-S:N.M.| U.8.1.M.
8181 12247 8143 | 20. 35388 | no. 132546
Skull |
Length from anterior end of premaxillaries to posterior |
endiof condyles: 2-25.24 2-23 == ee ee ee OE ene esa ose eee 284 304
Distance from line between posterior surfaces of glenoid
fossae to posterior end of condyles-_--.-.----------------- Ole Fae 5243S. see ee 96 101
Distance between anterior margin of orbit and anterior
endsofipremaxillary 3222000 ener - ce sen eee eee 105 1 120 112 103 108
Distance from line between postorbital processes of fron-
Cait; MON aes a ae enn en eee nce Diy ($n Pepsi S| Fee See 168 184
Greatest width across muzzle from outer walls of canine
alveolis =< eee oes: Re SAS ee ee eae 162 74.4 | 164 62 64.8
Least width between anterior margins of orbits__-------- 74 18350 jleoc-ses= 75 71
Greatest width across postorbital processes- ------------- 102 108) 79= 2258273 105 101.5
Greatest width across zygomatic arches__--.----.-------- TZN a Bae ny) Ne ee 180 186
Width of palate between alveoli of M?_....-.------------ 47 145 45.5 50 47.2

Superior dentition

Length of tooth row from anterior margin of C alveolus to |

posterior surface of Ma. 2 ~~ sss ne en 1103 1118 1110 103 107
Length of tooth row P4 to M2, inclusive----------------- LOS 64 62 54. 2 59. 5
C, anteroposterior diameter at alveolus---_-.-..--------- 120 125 121.5 20.6 20. 2
C, greatest transverse diameter at alveolus_---_---------- 11.6 15 13 12 1257
Pt anteroposterior diameter: ---- + ------ == eames 13. 2 13.8 13.7 11.5 12.1
P4, greatest transverse diameter _.-.-------------------- 9. 4 10 11.3 8.2 9.2
Mi; anteroposterior diameter: -~-.-- #2 =e == eee 19.3 19.3 21 17.2 19
M!, greatest transverse diameter_....._------.----------. 13.2{ 14.4 14.6 13. 2 15
M?, anteroposterior diameter_._......--...-------------- OT Tel S3ie ty] R273 26.6 29
M3, greatest transverse diameter_....-.--.----------.---- 15.2 | 17a 15.8 15.7 17

} Approximate.

PLATE 8

BUEEEDRIN 171

U. S. NATIONAL MUSEUM

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7 PE Adie 9

BULLETIN

NATIONAL MUSEUM

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A) UL porqryxay

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PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 27

TABLE 6.—Measurements (in millimeters) of mandible and inferior dentition of
three species of Kuarctos

peas Euarctos | Euarctos
Euarctos vitabilis americanus| luteolus

Measurement U.S.N.
M. no. | V-S:N-|U-8.N.| v.s.N.M.| U.S.N.M.
7665 814 % 8143 “| no. 35388 | no. 132546
(type)
Mandible
Length of mandible from anterior end to posterior surface
Of cond yiletee ee oso = Re oe ose oe eee senacsseess|hohseeee 1197 1195 1193 210
Depth ofijdwibelow Mir: 2252222 == 222 22 aoe oe aos 41.3 41 38 36.6 40
Thickness: of taw, belowsli 22222. 22 eee we ee oe eee 16 14.6 15.5 1733
Least width across both rami at constriction posterior to
Canines 230 35_ 0. 2 epee SA eee seo tedeenesese 36 41 | oceeeee 35 37.4

Inferior dentition

Distance from anterior margin of canine alveolus to pos-

terior'surface, Of) Mig stores boosh Ree Soto os nas 117 121 115 118.5 123. 4
Length of tooth row, P4 to Ms, inclusive___-------------- 162 67 66.5 61 69. 2
Diastema between canine alveolus and P4__------------- 136.8 33.3 31 36. 4 32.8
C, anteroposterior diameter at alveolus..__-.-_---_------- VS. 1), 20 119 120 1 20. 5
C, transverse diameter at alveolus_---------------------- 112 12.8 11 11.6 11.9
PP, ‘anteroposterionidiameters-=o-2- eee oe oo 11 10.9 9.9 9.5 9.8
Pi; greatest transverseidianietens. .-s2ss--s--——-2------=-|-22=---- 6.7 5.8 5.0 5.7
Mi, anteroposteriondiameterses---=.25)=-sa5o- ean o 17.8 20. 5 19.6 18.6 20.5
Mi, transverse diameter of trigonid-...-----.------------ 6.9 8.3 7.3 7.9 8.6
Mi, transverse diameter of talonid_...-_----------------- 8.3 9. 4 8. 4 9.3 10.7
Ms; anteroposterior diameter. —---=--~-..-..-.2-.---=.<-- 17.7 20.8 20. 5 19.5 22.5
Ms, greatest transverse diameter-_......------------------ 11.4 12.1 WS 11.8 14.5
IMs) anteroposteriomdiametersss--)--o--4-2 = se ee 1551 15.1 17.7 13.8 17.6
Mz, greatest transverse diameter_.._...------------------ 11.9 12 12.4 11.5 13.8

1 Approximate.
ARCTODUS HAPLODON (Cope)

A crushed right lower jaw, with the third incisor, canine, and second
molar, and an isolated second molar from the left side of the same
mandible, U.S.N.M. no. 8005, show the presence of a rather large
tremarctine bear in the cave fauna. Several limb bones and parts of
fore and hind feet, greatly exceeding in size those of a black bear and
apparently belonging to one individual, are also believed to be those of
Cope’s eastern cave bear.

The lower jaw from Cumberland Cave has been badly distorted by
crushing but appears to be slightly deeper than the jaw figured by
Cope (1899, pl. 19, fig. 2) from the Port Kennedy bone deposit.
Although the posterior part of the ramus, U.S.N.M. no. 8005, is not
preserved, the configuration of the jaw below M, suggests that the
antemasseteric pit probably extended forward nearly to a point below
the posterior margin of M,. In Ursus the anterior margin of the
masseteric fossa is not so far forward.

16541—38——3

BULLETIN 171, UNITED STATES NATIONAL MUSEUM

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f0z]S [BINJVM ‘META [VSNJOIO ‘D :(999L ‘OU “JA"N’S'Q) Ueumpoeds odéy ‘ofqipueuI Jo snureI yYSII ‘(AO[PIH) syiqnz1a sopunNn—‘T] aun

sere

ee ere Zi} a
SSS A e
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SS — Bun = = ;
NG SSX ey ete naes
N SS . Seen <= : Coo ea
\ v3 > o'NSSS ? GF ees
LYASsg Poor Teo Kote. EES spins Z —- edhe : ae
Xs ESS ssa: : ite eS ; =
Mer, > =~ + ee —
: typ Vint

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 29

| ZY #£ ds

FIGURE 12.—Ewarctos vitabilis (Gidley): Skull (U.S.N.M. no. 8181) and mandible (U.S.N.M. no. 8141), lateral view, three-fifths natural size.
Cumberland Cave Pleistocene, Maryland

30 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

FIGuRE 13.—Huarctos vitabilis (Gidley): Skull (U.S.N.M. no. 8181), dorsal view, three-fifths natural size.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE

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Figure 14.—Euarctos vitabilis (Gidley): Skull (U.S.N.M. no. 8181), ventral view, three-fifths natural size.

32 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

The canine and second molar are nearly identical in size with these
teeth in the Port Kennedy jaw. ™M, has an occlusal surface as that in
Cope’s specimen, but not so tuberculate as in Arctodus pristinus Leidy ;
however, the external surface is deflected as in Leidy’s specimen
(Leidy, 1860, pl. 23, figs. 3,4). Arctodus floridanus Gidley (1928) is a
distinctly smaller species approaching closely the living Tremarctos
inornatus Gervais. The second molar from Cumberland Cave is
noticeably narrower than in the western forms of Tremarctotherium.

The Frankstown Cave form, referred by Peterson (1926) to Are-
totherium haplodon, compares closely in size with the lower jaw from
Port Kennedy. M, in our specimen is similar to that in the Carnegie
Museum specimen and shows only a small difference in the outline of
the external surface. In the general appearance of this tooth the
Cumberland Cave form is perhaps closer to the Port Kennedy form
than is the figured specimen from Frankstown Cave.

The astragalus of the partial skeleton, U.S.N.M. no. 8180, referred
to Arctodus haplodon corresponds very closely to the dimensions given
by Cope for the Port Kennedy A. haplodon, but the third metatarsal
is distinctly smaller than that measured by Cope. It seems probable
that the difference is to be accounted for by size variation between
individuals in the Port Kennedy fauna. The foot bones of the
Cumberland Cave specimen are noticeably small compared to Trem-
arctotherium simum (Cope), according to dimensions given by Merriam
and Stock (1925), although the tibia has nearly the proportions of that
in the Potter Creek Cave form. 1. californicum (J.C. Merriam) from
Rancho La Brea is very much larger (Merriam and Stock, 1925).

Family MUSTELIDAE
MARTES PARAPENNANTI Gidley and Gazin §
Ficure 15

Type.—Anterior portion of male skull (fig. 15a), U.S.N.M. no.
8010, carrying most of the cheek teeth of the left side.

Material—In addition to the type there are 10 specimens in the
collection, consisting of lower jaws and skull fragments with teeth.
One pair of maxillaries, U.S.N.M. no. 12336, with nearly all the cheek
teeth on both sides, has a fragmentary right ramus of the mandible
associated.

Specific characters.—Size intermediate between Martes pennanti and
M. americana. Skull construction, as far as can be determined, nearly
as in MZ. pennanti. Upper carnassial relatively shorter than in M.
pennants with protocone more forward in position as in M. americana.
Upper carnassial with uncovered and well-developed external median
rootlet as observed in M. pennanti. Mi! relatively short anteropos-
teriorly with metacone and cingulum posteroexternal to metacone

5 Gidley and Gazin, 1933, pp. 347-349, fig. 2.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 33

somewhat reduced. External roots of M' not so distinctly separated
as in M. pennanti. M, with metaconid much reduced. Mandible
with low but anteroposteriorly elongate coronoid process. Masseteric
fossa deeply impressed. Coronoid process with two well-defined
fossae, one on its external upper half, the other on its inner base.
The modifications of the coronoid region are more pronounced than
in living species.
TasLe 7.—Measurements (in millimeters) of skull and mandible of Martes
parapennanti

U.S.N.M. | U.S.N.M. | U.S.N.M.

Measurement no. 8010 no. 11877 no. 11878
(type) (male) (female)
Skull
Width across postorbital processes-_----.---.-------.---------------- WIS) |Scceussesase |bass-oseccee
. Least width between superior borders of orbits...------------------- 194) ee |S cocsecause
Width between outer surfaces of M!__..---.-.---------------------- N92" ac cccebosos|sbekesossecs

Superior dentition

Length of cheek-tooth row, P! to M!, inclusive.-..------------------ 805i pe seeee Siac eee oo ceee
P4, anteroposterior diameter--------..--...------------------------- Diu poccee cone aacomaan oe
P?, greatest transverse diameter-._.--------------------------------- BG oe ecceses|sosececsenoe
Pe anteroposterior GiaNleLen. = 22 ese sa neo eee eee nae Seema ena Gi Gale as Sock adee
P3, greatest transverse diameter_-_...-----.------------------------- Ott | oseecne sacs | P Sse ee conan
Pi anteroposterior diameter. = ---------- ae ana aan eee == BO oN wwo caetecc|-cosecceescae
P‘, transverse width across anterior part including protocone-------- Gee [bccn ascent leaeeecoeso oe
P4, transverse width across metacone------------------------------- Be see
M1, anteroposterior diameter of outer lobe_------------------------- Gi oenccce een tees cacecaae
MI, anteroposterior diameter of inner lobe_-.----------------------- 6* leeetccecee ws Eososeotoece
M1, greatest transverse diameter--..-------------------------------- OUR ease sacse oes |aeaneaccosne
Mandible

Length of lower jaw from anterior surface of canine at cingulum to

middle of posterior surface of condyle-.-.---------------------------|------------ 6825) |cceacaeaas—o
Distance between posterior surface of M: and posterior margin of

coronoid above condyle--------2--2---2--=--------- <<... =----=-|----=------- Bi 2 (| oe ee
Height of coronoid above base of angle__----------------------------|------------ BL eenensecsase
Depth of jaw below heel of M: on external side_---------------------|------------ 12.6 9.2

Inferior dentition

Length of tooth series, C to M; inclusive_.--_-----------------------|------------ 40.8 136.5
Py anteroposterior Giameteres se ess sesne nee aaa eee aeee | eae ee eee enna ene a
Ps, anteroposterior diameter-..-..----------------------------------|------------ 5 2.8
Ps, greatest transverse diameter--._.--------------------------------|------------ 2.6 2.1
Ps, anteroposterior diameter---------------+---=--=_-_-_=-2-<---0-|--------=__- 6.4 5.4
Ps, greatest transverse diameter-------------------------------------|------------ 2.8 2.3
P,, anteroposterior diameter_--_--.-_--------------=---..---_-------|------------ 7.9 6.4
Pi, greatest transverse diameter-__...--------------------------------|------------ 3.3 2.6
Mir aTiteropOStenlor Gia MeLeheessess sae eee ee ane enna an ee |e 12.7 110.5
Mi) transverse width of trigonide 22 oo enon eee eee eee eee ean == 4.6 3.6
My, transverse width of talonid.---_-.------------------------------|------------ Ay 7; || ee

1 Approximate.

In Martes americana and M. caurina the upper carnassial is rela-
tively short, as it is in M. parapennanti, but P* in these living forms

34 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

lacks the external median rootlet characteristic of the fishers. M1!
also is relatively short anteroposteriorly in the martens and the meta-
cone is reduced, more so than in M. parapennanti.

The condyle of the lower jaw in M. parapennantt is relatively narrow
transversely and shallower than in the living species examined. A
glenoid portion preserved with one of the specimens, U.S.N.M. no.
11959, shows a correspondingly narrow glenoid fossa and the post-
glenoid process curves well forward along its inner border to form
an interlocking joint as in Gulo. In the living species of martens
this joint is perfectly hinged but is not completely interlocking.

FIGURE 15.—Martes parapennanti Gidley and Gazin: a, Anterior portion of male skull, type specimen
(U.S.N.M. no. 8010), lateral view with occlusal view of dentition; b, right ramus of female mandible,
paratype (U.S.N.M. no. 11878), lateral and occlusal views; c, right ramus of female mandible (U.S.N.M.
no. 11876), lateral and occlusal views; d, left ramus of male mandible, paratype (U.S.N.M. no. 11877),
lateral and occlusal views. Natural size. Cumberland Cave Pleistocene, Maryland.

Martes diluviana (Cope, 1899) from the Port Kennedy deposit in
Pennsylvania has a much shallower jaw, as indicated in the two figured
specimens, than male specimens of M. parapennanti with teeth of com-
parable size. The depth of the jaw is near that of a female M/. para-
pennanti with teeth considerably larger. Moreover, the metaconid
of the lower carnassial is apparently better developed in M. diluviana.
This is noticeable when comparisons are made with U.S.N.M. no. 12352
from Cumberland Cave, which has relatively unworn teeth. There
is but a very slight development of the metaconid on Mj, and this

PLEISTOCENE VERTFBRATES FROM CUMBERLAND CAVE 35

specimen though exhibiting a youthful dentition has a relatively much
deeper ramus than in either of the youthful specimens figured by Cope.
Dr. E. R. Hall (1936), however, regards these differences as of only
age significance.

The modifications of the upper facial region in the Cumberland
Cave form resemble those seen in M. nigripes, but in the latter species
the postorbital processes are more prominently developed and the
convergence backward to the postorbital constriction is more extreme.
The dental characters throughout are decidedly more like those of
M. vison. In M. nigripes the protocone of the upper carnassial and
the protocone shelf of M! are more reduced, and the heel of the lower
carnassial is very narrow, with the talonid basin entirely lacking.

MUSTELA cf. VISON Schreber
Ficures 16-18

Included in the mustelid material from Cumberland Cave are six
skull portions and an equal number of lower jaws belonging to mink
(see figs. 16-18). The fossil material closely resembles the form
M. vison mink living in Maryland at the present time. A number of
differences were observed between the fossil and modern forms, but
the magnitude or importance of these differences is not greater than
that of the distinctions seen to exist between several of the subspecies
currently recognized.

TaBLE 8.—Measurements (in millimeters) of skull, mandible, and dentition of
Mustela vison

nn nee EEUU ESSE Sa

Must_la v. | Mustela ov.

Mustela cf. 7 re a eae
vison S.N.M. .S.N.M.
Measurement U.S.N.M. | no. 2242 | no, 241192
no. 8156 (Biol. (Biol.
Surv.) Surv.)
Skull and superior dentition
Distance anterior end of nasals to line between postorbital processes_ lant) 15 16.8
WidthubetweenlOnDitSee see esea anes ee een e nena seen a nena eee 14.5 14 14.7
Width at postorbital processes--_=-----.----...---------..------.--- 16.8 16 16.3
Width across outer margins of canine alveoli-____-------------------- 13.8 1255 13.6
Length of upper tooth series, anterior margin of canine alveolus to
posterior surface of carnassial-__----------------------------------- 19 17. 4 18.4
P4, anteroposterior diameter along outer wall_-_--------------------- 7.4 6.7 7.5
P4, transverse diameter including protocone-_------------------------ 4.7 3.8 4.4
P4, transverse diameter posterior to protocone----------------------- 2.6 2.2 2.4
Mandible and inferior dentition
Depth of lower jaw below trigonid of M1-.--------------------------- 6.5 6 6.7
Distance from apex of coronoid to base of angle--------------------- 17.7 17 18.4
My, anteroposterior diameter: ----------------------------=------=--=-= 8 7.5 8.2
Mi, transverse diameter of trigonid over anterior root--------------- 2.5 2.4 2.5
Mi, greatest transverse diameter------------------------------------ 3 Sal 3.2
Mi, transverse diameter of talonid...------------------------------- 2.8 2.9 3

36 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

WF
Mg
j

FIGURE 16.—Mustela cf. vison Schreber, anterior portion of skull and right ramus of male mandible
(U.S.N.M. no. 8156): a, dorsal view of skull; b, ventral view of skull; c, lateral view of skull; d, lateral view
of mandible; e, occlusal view of interior dentition. x 144. Cumberland Cave Pleistocene, Maryland.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE on

In size the fossil form is close to M. vison mink. A comparison
with this subspecies shows the muzzle of the fossil skulls to be rela-
tively broad posteriorly with a slightly elongate face. Viewed from
above the lateral lines of the postorbital constricted portion of the

Figure 17.—Mustela cf. vison Schreber: Anterior portion of skull and mandible (U.S.N.M. no. 11880),
anterior and lateral views. Natural size. Cumberland Cave Pleistocene, Maryland.

skull (fig. 166) converge posteriorly, perhaps more so than in the
Alaskan form, M. vison ingens. The upper dentition is essentially
like that in M. vison mink, although the upper carnassial of U.S.N.M.
no. 8156 (male) (fig. 16c), is somewhat broader posteriorly than in the
living mink. The lower jaw (fig. 16d and e) is less robust than in

FiGure 18.—Mustela cf. vison Schreber: Right ramus of female mandible (U.S.N.M. no. 8212), lateral and
occlusal views. X114. Cumberland Cave Pleistocene, Maryland.

M. vison mink, but the lower teeth, with the exception of M’, are not
different. The heel and middle portion of the lower carnassial are
narrower in the fossil jaws, and the hypoconid appears to be more
medially placed. The relative breadth of this tooth does not vary
greatly within subspecies, but between such forms as M. v. vison and
M. »v. ingens the proportions differ appreciably. Incidentally, the
junior author does not follow the procedure adopted by Dr. E. R.
Hall (1936, pp. 114-115) in referring the fossil material to the living
subspecies, M. vison mink, occupying the Maryland region today.

38 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

GULO GIDLEYI Hall ®

Figures 19, 20; Pats 9

The normal habitat of the wolverine today is the far north, and in
the Eastern United States it has not been known to range farther
south than northern New York. This animal is represented in the
Cumberland Cave collection by several specimens including an
unusually well preserved skeleton.

The wolverine skeleton, U.S.N.M. no. 8175, has been mounted and
is on exhibition in the hall of fossil vertebrates at the National Museum

AS y
Ly),

FiaurE 19.—Gulo gidleyi Hall: Skull and mandible (U.S.N.M. no. 8175), lateral view. Two-thirds natural
size. Cumberland Cave Pleistocene, Maryland.

(see pl. 9). The skull and mandible belonging to this specimen are
nearly complete and most of the teeth are intact. The ribs and the
anterior portion of the vertebral column are largely restored, and the
sacrum and most of the caudal vertebrae were not preserved but the
lumbar and posterior dorsal vertebrae are in good condition. The
right scapula is only half complete but has been restored from the one
on the left side which has very little missing. The fore limbs and right
fore foot are entire. Several carpals, phalanges, and claws of the
left manus were not preserved but have been modeled from those on
the right. Of the pelvic girdle only the tip of the left ilium and the
symphysial portion of both ischia are restored. The hind limbs are
complete except for the fibular and one patella. The left pes includes
most of the elements and served to model the right hind foot, which
had only the astragalus, cuboid, and third metatarsal preserved.

In proportions of the skull a noticeable degree of variation is seen
between specimens in the cave collection. The skull (figs. 19, 20)
belonging to the preserved skeleton, U.S.N.M. no. 8175, is character-
ized by a broad frontal region and well-expanded zygomatic arches;
moreover, the posterior narial passage is noticeably constricted, and

6 Hall, 1936, pp. 83-86.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 39

the pterygoids approach each other somewhat more than in the living
species. However, skull U.S.N.M. no. 8176, the type as designated
by Hall, in the cave collection, is not so broad through the muzzle, and
the posterior narial aperture is not different from that in an average

Z va Mi
Miwa
loss

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MWA i XG
' ‘ LN) Ge
ANA asttling tT Se
N\ We i)

Fiaure 20.—Guwlo gidleyi Hall: Skull (U.S.N.M. no. 8175), dorsal and ventral view. Two-thirds natural
$128.

modern specimen of Gulo luscus. The occipital condyles in no. 8175
are close together, and the foramen magnum is correspondingly small.
Unfortunately the occipital region is missing in other skull portions
from Cumberland Cave, but the distance between the condyles in
living specimens of G. luscus varies sufficiently to include the condition
seen in no. 8175.

40 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

TABLE 9.—Measurements (in mallimeters) of skeleton of Gulo gidleyi

Measurement a aire :
Skull
Total length of skull from anterior end of premaxillaries to occipital condyles__.._----------_- 136
Widthiat:mastoids_02 "2. =!: 202) 502 eee ence ee eek ee ee ee 77.6
Widthiacross'zygomatic' arches... 2222-22-22 5 ee eee 98. 8
Width'ofimuzzleiat' canines= +=. - 2-250 =e See SS ee es ee ee 37.5
Width of muzzle atjinfraorbitalforamensss. "loos foe 2S ee ee ee 41
Width: between orbits: ==> 2S ue. Sot See eS Se ee 43.3
Width/across: postorbitaliprocesses==e2-- soo a ee ee ee ee 50. 6
Width:at postorbitalconstriction_2-— 22-2. Ao ee ee 37.2
Superior dentition
Length of upper tooth series, canine to M!, inclusive. ..---....-....-.---------.---.----.----- 53.5
Ps anteroposterior diameter. - asa--2osee oon nonne nen eeate= sane an eas ee ee eee a 18.9
P4, transverse'diameter including protocone-j.=2=- 22 = 2s conc nce nn see ees cce sna eee eee ll
P4, transverse diameter (greatest) posterior to protocone..-.-.-.......-.....------------_.-.-- 8
M1, anteroposterior diameter of external portione.2.- 22... 2-2 ooo 22a see ee ee 7
M1 anteroposterior diameter’ of internal) portion==22- 2202s 2ee oe seccccesaceeceeneso eae 1.21
Mi, transverse: diaméter=..- 322.2220) o es edn das eae cee atnesas anceaneeeee ee eee ee 12.5
Mandible and inferior dentition

Depth of lower jaw: below trigonid of; carnassial_—-----2. _-... ees eee 19.5
Length of lower tooth series, canine to Ma, inclusive._..--.-..2.-----<_--.-.----=..<-+-------- 60. 5
Mi:;, anteroposterior diameters. 2.25282 Senet 8S soese cae so oee tow sae atccescenea ee ees 19.5
Mi, greatest transverse diameter of anterior portion. --_-.......--.-.----.-------------------- 8.4
Mi, greatest transverse diameter of posterior portion. —_-.......-.-...--...------------------- Ton

Anterior limb
Greatest length of scapula from coracoid process to suprascapular border___-.--.-..---------- 191
Greatest width of articulating end of scapula measured across glenoid cavity___-.--------.-_- 26.5
Distance from inner border of glenoid cavity of scapula to outer surface of acromion_____-__-- 26. 2
Greatest lengethofshumoerus #2525 226 2s eae es a eee Sone ee een 128
Greatest transverse diameter of proximal extremity of humerus__._--...__-___-------_--_______ 27.2
Greatest anteroposterior diameter of proximal extremity of humerus_......---.-.---_-------__ 20.3
Greatest width of distal extremity: ofishnmerns2...-2 == 2-2 22~ 5 aoe ne ne ee 34.6
Greatest length: of ulna 2e2: = 22. 2 See os a oa ee ee 129
Greatest transverse width across sigmoid cavity of ulna.._.-.-........-------------_-- 22 15.3
Distance from distal margin of sigmoid cavity to distal end of ulnma__.___--__----.-______.____ 104
Greatest length: ofraditis’..25 2225 iv oso ee a es 107
Long diameterrofproximaliemd: Of radi sso see es ae a a 15.8
Greatestiwidthrofdistaliend offradius sos see ease ee ee ee ee 20
Length of first moetacarpal:: sick See ates 2s Re ee ee ee 25.9
Lenethiof/second;metacarpal ses ses. see re ee oe ee ee es 35.9
Length of'third:metacarpal sss can cs aoe tee ee ee ee ee ee eee 44.5
Length:.of fourth metacarpali 2.2 "si s<200 eee ee pn ee eee ee ae 42.8
Length of fifth metacarpal: 2-2! 6.. Sarco! es ee ee ee 37

Posterior limb
Length from anterior end of ilium to posterior border of ischium__-_---.-.--...-----.---------_ 112
Width ofilitnm2-0) 102 eres aren SMe 8 a os SO a a ee ee ee ee ee 30
Width of ischium measured from ischial tuberosity to posterior end of ischial symphysis_----- 45
Diameter of acetabulum measured at right angles to long axis of internal notch______________- 18.5
Length of femur from top of head to distal margin of inner condyle_-_----------------------__ 134
Width of proximal end of femur from inner surface of head to outer surface of greater trochanter_ 34.4
Anteroposterior diameter of head offers sss ee oe eee 16.4
Widthiofidistal‘end of femur‘across cond yles¥4- 2. ae ee Se 30
Greatest length of tibia measured parallel to long axis.___.--.......-..----.-.---------------- 133
Greatest anteroposterior diameter of proximal end of tibia_____..-......___._-.--__---________- 27.8
Greatest, widthiof proximal end Jofitibia 22 te Ee ee eae 29.4
Greatest transverse width of distal end of tibia measured perpendicular to long axis of shaft __ 21
Length oftfirst metatarsal’ 32.2. a). Vd 0 eee ee ee eee ee ee 34.4
Length/of'sécond metatarsal-2) 15 Soft Baird Vat hi eee bi Ot Pi ea 1 44.6
Length of third metatarsal 20 =-- 28>). Le een ee ee ee ee 52. 5
Lengthiofifourthimetatarsal vaste. ose a a eae a eg 54.2
Lengthiof fitthi metatarsals ii1f 44) visti BOP Bam See Ge eG 51.5

1 Approximate,

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE Al

The teeth average somewhat smaller than in Gulo luscus, as noted
by Hall, although a few specimens were found in which the teeth
were definitely smaller than in the fossil form. The relatively wide
P? in the fossil material was matched in some recent individuals but
this condition is uncommon in CGulo luscus.

The other bones of the skeleton, no. 8175, show but few peculiarities,
the principal ones being that the anterosuperior border of the scapula
is more rounded than in the living species, and the shaft of the humerus
is slightly less curved.

The record of Gulo in the Pleistocene of North America is very
incomplete, and the material for the most part has been referred to
the living species, Gulo luscus. Several lower jaws of wolverine in
the Port Kennedy occurrence, believed by Cope (1899, pp. 229-230)
to be specifically identical with Gulo luscus, are referred to Gulo
gidleyi by Hall.

LUTRA PARVICUSPIS Gidley and Gazin’
FiaurE 21

Type—Portion of a broken and crushed skull (fig. 21), U.S.N.M.
no. 8213, in which the principal cheek teeth of both sides are present.
P!, the canines and incisors represented by their alveoli.

Specific characters—Size somewhat larger than Lutra canadensis.
Main outer cusps of M! relatively lower and less robust than in L.
canadensis. P2 and P® large with conspicuous posterointernal basin.
P‘ triangulate, with its base relatively wide anteriorly and with its
anterior border forming a right angle with the outer border of the
tooth. P*long and outer wall of M' relatively short anteroposteriorly
as compared with L. canadensis. M’ with paracone and metacone
subequal and both cusps with relatively narrow extension of the
basal portions of their outer walls.

Comparison.—The Cumberland Cave specimen represents an
extinct species of otter intermediate in size between L. canadensis of
the Eastern United States and L. felina or L. paraensis of Chile and
Honduras, respectively. Judged by their alveoli the canines of the
Cumberland Cave specimen are relatively robust, as in the living
species of Central and South America. The backward extension of
the protocone shelf of the carnassial and the posterointernal shelf of
the other premolars is somewhat more expanded than in L. canadensis.
The posterior margin of the upper molar, as in some South American
specimens, is not so deeply notched posterointernal to the metacone,
as noted in L. canadensis. Perhaps the most noticeable distinction in
the dentition of L. parvicuspis is the distinctly narrow proportions of
the external cusp row of M’.

Although L. parvicuspis is distinct from both North American and
South American otters, it approaches living species of South America

7 Gidley and Gazin, 1933, pp. 349-351, fig. 3.

42 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

in several of those characters in which it differs from northern forms.
Most noticeable among these are the size of the skull and canines, the
development of the upper premolars, and the somewhat smaller
relative size of the molar.

FIGURE 21.—Lutra parricuspis Gidley and Gazin: Maxillary portion of skull with premolar and molar
teeth, type specimen (U.S.N.M. no. 8213), lateral and occlusal views. Natural size. Cumberland
Cave Pleistocene, Maryland.

A comparison of L. parvicuspis with L. rhoadsii Cope (1899, pp.
238-239) from Port Kennedy, Pa., is difficult on account of the
different nature of the materials preserved. L. rhoadsii was based on
lower jaws and a single upper molar. The dimensions given by Cope
for the anteroposterior diameter of the lingual portion of M! shows
this tooth to be much smaller than in LZ. parvicuspis.

TaBLEe 10.—Measurements (in millimeters) of superior dentition of three species

of Lutra
Lutra Lutra Lutra
parricuspis canadensis felina
Measurement U.S.N.M. U.S.N.M. U.S.N.M.
no. 8213 no. 197943 no. 35377
(type) (Div. Mamm.)| (Div. Mamm.)
Length of cheek-tooth series, P! to M!, inclusive_ _-.-.__--- 134.5 30 33.6
C, anteroposterior diameter of alveolus__._--_-------------- 17.7 6.5 7.6
‘Pi anteroposterior, dismoeter=_ ---. 2-22 =---= =— a ee 5 5 5
P2Noreatest transverse: Glameter-.so-- se. eee 4 a3 4.3
Ps anteroposterior diameter. <225 2s 22 ee eee qed 7.5 8.2
P3: greatest; transverse diameter=.2...--.os2<5---5-8-—-=-25- 5.3 5 6
Pt anteroposterior @iameten- =. =e eee ee 12.8 11.5 14.5
P4, greatest transverse diameter perpendicular to outer wall _ 10. 2 9.6 11.5
M!, anteroposterior diameter across outer cusps__---------- 8 8.7 8.3
M!, anteroposterior diameter across inner portion____-_____ 8.7 8 8.5
M!, transverse diameter across paracone and protocone- --- 10.5 ll 11.8

1 Approximate.

Satherium piscinaria (Leidy, 1873, pp. 230-231) from the late
Pliocene or early Pleistocene of Idaho is a considerably larger form as
indicated by the size of the tibia. (See also Gazin, 1934, pp. 143-149).

E. R. Hall (1936, pp. 75-77) regards Lutra parvicuspis as a synonym
of L. canadensis lataxina. In examining all the material of this recent
subspecies in the National Museum the junior author finds that the

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 43

fossil species is probably distinct from the living form and that several
of the characters cited above for the fossil type are distinctive and
others are present only in unusual specimens or extreme conditions
of the modern form.

BRACHYPROTOMA PRISTINA Brown #
Figures 22-24
Included in the musteline material from Cumberland Cave are

three skull portions, one of which is nearly complete (fig. 22), and
four mandibular rami representing an extinct form of skunk.

FIGURE 22.—Brachyprotoma pristina Brown: Skull (U.S.N.M. no. 8155), dorsal and lateral views with
occlusal view of dentition. 144. Cumberland Cave Pleistocene, Maryland.

= The skull material corresponds closely in size and other characters
with the type of Brachyprotoma pristina, which was described by
Brown from the Conard Fissure in Arkansas. Skull U.S.N.M. no.
8155 (fig. 22) is rather short and broad, more so than in the living
species Spilogale ambarvalis. The width is most noticeable through
the frontal region and is accompanied by a reduction in size of the
postorbital processes. Prominent dorsolateral swellings in the
vicinity of the postorbital constriction emphasize the breadth of the

8 Brown, 1908, 176-179.
16541—-38—-4

44 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

skull in this region. The cranial portion is conspicuously shortened
anteroposteriorly and is convex dorsally in a longitudinal profile
somewhat as in the Florida S. ambarvalis. Some of the modern
western skunks, as S. gracilis and S. phenaz, have relatively flat dorsal
profiles and are also more elongate than the cave specimen. Further-
more, the longitudinal profile of rostrum is slightly concave anterior
to the frontals, as noted in several specimens of S. putorius.

The dental formula of Brachyprotoma is as in Conepatus, and the
characters of the upper teeth of the Cumberland Cave specimen corre-
spond rather closely with those of B. pristina. M1! has a broader heel
than in the type, but the difference may not be greater than can be
attributed to individual variation or possibly to difference of sex.

FIGURE 23.—Brachyprotoma pristina Brown: FIGURE 24.—Brachyprotoma pristina Brown:
Left ramus of male mandible (U.S.N.M. no. Right ramus of female or young male mandi-
12045), lateral and occlusal views. X 1}. ble (U.S.N.M. no, 8214), lateral view. X 1%.
Cumberland Cave Pleistocene, Maryland. Cumberland Cave Pleistocene, Maryland.

A comparison with Spilogale shows P* in our specimen to be similar
to that in the living species but with protocone relatively smaller
and the anterior margin of the paracone slightly more backward slop-
ing. M1! is relatively short anteroposteriorly and wide transversely,
and the median constriction is not nearly so pronounced as in Spilogale.
The external cingulum of this tooth is very well developed and termi-
nates anteriorly in a prominent cusp external to the paracone.

Of the four jaws referred to Brachyprotoma pristina, one (fig. 23),
U.S.N.M. no. 12045, is evidently of an adult male and corresponds
very closely to the lower jaws Brown recognized as belonging to B.
pristina. Jaw no. 12046 is apparently of an adult female and corre-
sponds favorably with the type of B. spelaca. The remaining two
jaws (fig. 24) seem to represent relatively young males. The adult
male specimen, U.S.N.M. no. 12045, is characterized by a moderately
robust mandible with a rather deere abrupt symphysis, and a well-
developed masseteric fossa as compared to Spilogale. The premolars
are crowded, overlapping, and the lower carnassial is narrow and short
heeled and exhibits a weak metaconid, differing markedly in these
respects from living forms. The female specimen, no. 12046, differs
from the male in having a weaker mandible, particularly Srallene be-
neath the posterior portion of My, a shallower symphysis, and a more
gently sloping ascending ramus.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 45

It seems evident from a consideration of the skeletal characters
distinguishing males from females among living mustelines related to
this fossil form that those characters that have been cited to separate
B. spelaea from B. pristina should be considered sexual.

B. obtusata from Frankstown Cave in Pennsylvania, as recorded by
Peterson (1926, p. 285), possibly differs from the type of that species,
which is from Port Kennedy, in the size of the lower carnassial. The
lower jaw illustrated by Peterson, however, does not resemble closely
the specimens of B. pristina from Cumberland Cave. The difference
is most noticeable in the greater downward projection of the symphy-
sis, the lower position of the ventral surface of the angle, and in the
differently shaped coronoid process of the Frankstown Cave jaw. If
the material from Frankstown Cave should be regarded as belonging
to the same species as that from Conard Fissure and Cumberland
Cave, it seems preferable to refer it to B. pristina, since no comparisons
are possible with the type of B. obtusata, which is lost. Cope’s meas-
urements for the latter, though possibly in error, indicate a smaller
species than B. pristina. Moreover, there is no assurance that the
Port Kennedy deposit is the same age as that at Frankstown Cave.

TasiE 11.—Measurements (in millimeters) of skull, mandible, and dentition of
Brachyprotoma pristina

Wi. tho) S046 | Wat | SAE | Wise | SE
: .M. .H. n0. -H. .M. SHS -.M. BEES
Measurement no. 12426 no. no. no. no. no.

8155 | (type) | 11722 12045 11773 12046 12399

Skull and superior dentition

Length of skull from anterior end of pre-

maxillaries to occipital condyles_------------ Bark YS ARSE | Seana ss | Sets ea ee eo ee
Width betweenerbits_ .-.--.----------=.---== DRT S52] pe 8 ann eee oe eee ee | ee ee ee ol eee tated
Outside measurement across upper molars_---} 20.1 DOS Bis eco = ee es ee SO lags ese ee ee
Length of tooth series, P3 to M!, inclusive----- 1S 2tleeeee = = QR Looe eee eee ee eee eee
P4, anteroposterior diameter___-----.--------- 6. 2 6.9 Gi 2)y | eee sees [Leese aCe 2 oes ree es
P4, transverse diameter including protocone-- 3.9 3.8 BEO) esate | eee |oaee en |aaeeceas
M1, anteroposierior diameter of outer portion

of tooth parallel to outer wall_-_------------- 4 3.8 oe ee ee) pecs = el |S eee ed ee
M1}, anteroposterior diameter of heel_---_------ 3.8 3 BoB u eee seers eae | ee ee eee
M1, greatest transverse diameter------------- 6.3 6.7 Tee | Senn eae ee een | sen ne soem

Mandible and inferior dentition

Length of lower jaw from anterior end to con-

iy One eae eo nee ene ee aa eat eae ae omens |[Pmmee 130 8055) |zeacees 26.6
Depth of lower jaw below P2-----------------|--------|--------|-------- 6.8 6.8 5.9 16
Depth of lower jaw below talonid of Mi------|--------|--------|-------- 5.1 5.3 4.1 4
Length of premolar series_...-----------------|--=-----|--------]-------- 6.3 6.5 633) |e See
My, anteroposterior diameter-__--------------|--------]--------|-------- “se tok 7 6.8
My, greatest transverse diameter--.-.----------|--------|--------|-------- 3 3 3 2.7
My, transverse diameter of talonid_.---.------|--------|--------|-------- 2.7 2.8 2.7 2.5

1 Approximate.

46 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

SPILOGALE MARYLANDENSIS Gidley and Gazin ®
Figure 25

Type.—Right ramus of mandible (fig. 25), U.S.N.M. no. 12048,
with canine and P; to M, preserved.

Specific characters—Mandible about the size of that in Spilogale
putorius. Symphysis abrupt and inferior margin of mandible less
convex longitudinally. Canine slender and long. Third and fourth
premolars and molar slender and not overlapping. P3 small. Basal
portion of P* nearly oval in cross section as seen in dorsal view.
Trigonid of M, relatively short and narrow. Metaconid moderately
well developed but not so distinctly separated from protoconid as in

FIGURE 25.—Spilogale marylandensis Gidley and QGazin: Right ramus of mandible, type specimen.
(U.S.N.M. no. 12048), lateral and occlusal views. 144. Cumberland Cave Pleistocene, Maryland.

S. putorius. Buccal surface of talonid not offset inwardly to so great
an extent as in living species.

Comparison.—Spilogale marylandensis compares favorably in size
and depth of jaw with female specimens of S. putorius. However,
the symphysis of the mandible is somewhat more abrupt than in either
male or female specimens and projects downward to a noticeable extent
giving the lower margin of the ramus a less convex and more irregular
longitudinal profile. The premolars are narrower and even less
crowded than is common in the living species. The trigonid of M,
is slightly shorter and narrower than in either S. putorius or S. in-
terrupta, and the external wall of the heel is not offset inward so
abruptly from that of the trigonid as in the living species. Further-
more, the metaconid of M, is more solidly united with the protoconid;
however, this cusp is not reduced as in Brachyprotoma.

Spilogale marylandensis does not differ greatly in size from Brachy-
protoma pristina but has smaller, more slender premolars and a larger
carnassial. Other characters separating the two forms are those which
distinguish Brachyprotoma from Spilogale, such as the crowding or
overlapping of the premolars and smaller size of heel and more reduced
metaconid of the lower carnassial.

9 Gidley and Gazin, 1933, pp. 351-352, fig. 4.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 47

Taste 12—Measurements (in millimeters) of mandible of Spilogale marylan-
densis and S. putorius

. Spilogale Spilogale

rinraldnateceta (panes, 0 patente

Measurement U.S.N.M. no. 42892 ewe 43719.

no. 12048 s E
(type) (male) (Biol. (female)
urv.) (Biol. Surv.)
NB 8

Depth of ramus below Ps---.------------------------------- 5.5 6 5.5
Depth of ramus below posterior portion of My-------------- 4.4 5 4.4
Length of dentition, C to Mu, inclusive-------------------- 18.3 18.5 18.1
P3, anteroposterior diameter ------------------------------- 2.6 2.8 2.9
P3, greatest transverse diameter---------------------------- ere 2.1 1.9
P,, anteroposterior diameter ------------------------------- 3.4 3.6 3.3
Pu, greatest transverse diameter---------------------------- 2a 2.4 2nd
Mj, anteroposterior diameter_------------------------------ | 7.4 7.4 7.6
My, anteroposterior diameter of trigonid ------------------- 4.4 4.7 4.8
Mi, greatest transverse diameter of tHigonidsese =e ne One 3.5 3.4
M1), transverse diameter of heel ---------------------------- 3 ool Sal

Ne eee ee
TAXIDEA MARYLANDICA Gidley and Gazin 1°

Ficures 26, 27

Type.—A complete skull and lower jaws (figs. 26, 27), an articulated
series of 11 vertebrae in which all the cervicals and four dorsal are
present, and the proximal half of the night humerus of one individual,
U.S.N.M. no. 7990.

Specifie characters.—Size large, about equaling the largest living
race of Taxidea taxus. Cheek teeth relatively large. Upper carnas-
sial with both protocone and hypocone having low regularly cone-
shaped contours with circular bases; hypocone relatively large. M1?
complex in structure, with the five principal cusps (paracone, meta-
cone, protocone, hypocone, and metaconule) low, regularly cone-
shaped and subequal. This tooth in consequence has a flatter appear-
ance than is usual in the living species. It is also more expanded
posterointernally, so that the general outline of this tooth is not
triangular, as is usual in the living species, but subquadrangular with a
relatively straight posterolateral margin and a posteromedially directed
lingual margin. Also, the posterior portion of the tooth is distinctly
wide transversely. Lower P, is relatively long with well-developed
heel, and the cusps of M; are distinctly robust.

The skull, in general appearance, is much like that of 7. taxus, but
it presents a few peculiarities that seem to distinguish it from any
living species. Viewed from the side, the region above the orbit is
high-arched; the occiput is relatively depressed ; and the postorbital
zygomatic process is relatively low. In the basicranial portion of the
skull the bullae appear to be more swollen and more evenly rounded
than is usual in the living species. In superior view the principal
differences to be noted are the slight development of the postorbital

10 Gidley and Gazin, 1933, pp. 352-354, figs. 5, 6.

48 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

processes of the frontals as compared with recent skulls in the same
stage of maturity and especially the form and size of the nasals.
These are broad in midregion so that the outer sides run nearly
parallel for a considerable distance instead of converging \-shaped.

ay ae 3
W\\) UT rs.
Wy W ij )

AS S\N

YY
1 HY

FIGURE 26.—Taridea marylandica Gidley and Gazin: Skull and mandible, type specimen (U.S.N.M. no.
7990), lateral view. Three-fourths natural size. Cumberland Cave Pleistocene, Maryland.

FIGURE 27,— Taridea marylandica Gidley and Gazin: Skull, type specimen (U.S.N.M. no. 7990), dorsal view...
Three-fourths natural size. Cumberland Cave Pleistocene, Maryland.

backward as is usual in the living species. Thus the points of the

anterior extension of the frontals are very definitely farther apart

than in any specimen of the living species observed.

The junior anthor dees not regard this specimen as sufficiently
typical of Taxidea taxus taxus to be placed in the modern species and
subspecies, as does Hall (1936, pp. 79-82).

Badgers today are considered strictly western in habitat, and in
this country they are now not found in any numbers east of the
Mississippi River.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 49

Badgers from the Pleistocene of North America are relatively little
known. Tavzidea robusta, described by Hay (1922, pp. 631-632) from
Arizona, is known only from an ulna and part of an innominate bone.
This material is not that in which one would expect to find significant
characters of specific importance; hence there is little justification in
referring it to a living subspecies, as does Hall (1936, pp. 82-83). The
Anita, Ariz., material should have been cited as Tazidea sp.

A specimen of 7’. tarus, originally described by Cope (1878, p. 227)
as T. sulcata, is recorded from the Pleistocene of Washington. Cope
(1899, p. 239) also recognized the modern species in the Port Kennedy
fauna.

TapLE 13.—Measurements (in millimeters) of skull and dentition of Taxidea
marylandica and T. taxus

Taridea
marylandica fe
Measurement U.S.N.M. US an

Taxidea tarus

no. 7990 (Div. Mamm.)

(type)
Lee eS SS SS
Skull
Total length of skull including condyles-.--------------------------------- 129.3 130
Width across mastoids.--- = <= oo eee = ne ene === = 77 73. 6
Width across zygomatic arches-------------------------------------------- 81.2 78.7
Width of muzzle across canines__.....-...----------..-----------------_-_- 36.3 35. 7
Width of muzzle at infraorbital foramina-.------.------------------------- 34. 2 34.3
Widths between O©bDitS sa seeeseseanee esa ooo eae e ne ween 30 27.5
Width across postorbital processes- ---------------------------------------- 33 33.8
Width across postorbital constriction _------------------------------------- 30 30
Depth of skull at bullae._-_--.-------------------------------------------- 51.5 56.5
Depth of occiput ---------------------------------------------------------- 42.5 45.5
Width of nasals at posterior points of premaxillaries_-_--------------------- 10.7 10.7
Width of nasals at anterior points of frontals_...--------------------------- 10.7 6.5
Superior dentition
P4, anteroposterior diameter along outer wall------------------------------ 112.5 12.2
P4, transverse diameter perpendicular to outer wall_----------------------- 110.5 10.8
M1, greatest diameter-___.-------------------------------------------------- 114 14
M1, transverse diameter perpendicular to postero-external walle s2sscs 2. 110 9.4
Inferior dentition

Length of lower tooth series, C to M2, inclusive._-------------------------- 60.5 52.5
Length of lower cheek tooth series P2 to M3-------------------------------- 40 41.2
Pu, anteroposterior diameter_---------------------------------------------- 9 7.9
Mi, anteroposterior diameter---------------------------------------------- 14.3 15
Mi, anteroposterior diameter of heel --------------------------------------- 5.2 5.2

1 Approximate.

Family FELIDAE

FELIS cf. INEXPECTATA (Cope)
Figure 28
A relatively small quantity of cat material is included in the col-

lection. An upper carnassial and a fragmentary left mandibular
ramus (fig. 28a and 6), U.S.N.M. no. 11890, with P, and M, preserved

50 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

are referred questionably to Felis inexrpectata (Cope, 1899, pp. 247-
249). The proportions of the upper carnassial are such that the tooth
may be from the same individual as the lower jaw."

Cope described F’. inexpectata with an upper carnassial from the
Port Kennedy deposit as the type. To this species he referred a lower
jaw fragment with M, and part of P,, some separate upper and lower
teeth, foot bones, and limb fragments. The Cumberland Cave upper
carnassial is slightly larger and higher crowned than the type of F.

FIGURE 28.— Felis cf. inexpectata (Cope), upper carnassial and left ramus of mandible (U.S.N.M. no. 11890)?
a, Lateral and occlusal views of upper carnassial; 6, lateral and occlusal views of mandible. Two-thirds
natural size. Cumberland Cave Pleistocene, Maryland.

inexpectata, but it has its protocone noticeably reduced as in that
species. However, the Port Kennedy lower jaw is much deeper and
has larger teeth than the Cumberland Cave jaw portion and the notch
between the paracone and protocone of the lower carnassial is deeper
and more open than in M, of the latter specimen. The Port Kennedy
lower jaw is out of proportion with the type upper tooth, which sug-
gests considerable individual variation or possibly that another species
is represented.

The Cumberland Cave upper carnassial is nearly equal in size to the
type upper carnassial of Felis longicrus Brown (1908, p. 178). The
principal differences to be noted include somewhat narrower propor-
tions, a more backward sloping parastyle, relatively shorter antero-
posterior length of paracone, and a more reduced and less forward
projecting protocone than in F. longicrus. The P, referred by Brown
to F. longicrus is similar to that tooth in the Cumberland Cave man-
dible but is distinctly larger. Brown noted the resemblance of this
tooth to I’. onca and to the P, in the jaw referred by Cope to F.
inexpectaia.

The mandible from Cumberland Cave is distinctly pumalike in
character, particularly in the relative proportions of P, and Mj.
Though smaller than an average size jaguar, it is apparently larger

1t The specimens were associated, according to Dr. Gidley’s notes.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 5l

than the living pumas. It approximates in size F. daggetti Merriam
(1918) from the Pleistocene of Rancho La Brea but possesses notice-
ably slenderer teeth.

TaBLE 14.—Measurements (in millimeters) of mandible and dentition of Felis
cf. inexpectata

Measurement aN
Superior dentition

P4 greatest anteroposterior diameter--—------.---2_----2-__-__--2 == -- en 26. 6
P4, anteroposterior diameter of paracone between notches------------------------------------ 10.1
P‘, anteroposterior diameter of metacone-----.....------------------------------------------- 10.9
P4, transverse diameter including protocone -_--_--.----------------------------------------- 12,2
P4, greatest transverse diameter posterior to protocone---------.----------------------------- 10

P4, height of paracone above base of cingulum externally --..----.---------------------------- 13.5

Mandible
Distance between posterior margin of M: and posterior surface of condyle-------------------- 64.8
Depth of lower jaw below cingulum of M, externally-__-------------------------------------- 28
Inferior dentition

PoP eaTILCEOPOSUCTIOG AIRING GIs eae se a ee 17.2
P,, transverse diameter of’anterior portion— —--------_-=_---=--_---._-_--_-_ =e 7.6
Pi greatesm@trans verse) GiamMeter a= ene e eo cere nee enn ee a ee 9.1
P4, height of principal cusp above base of cingulum externally-----.-------------------------- 12.5
IVEs sin LELOWOSLELION Ci aln Chek see an ee a ee er ee a ea 20.9
Mh, greatest transverse diameter at cingulum___.._-_._____-_.__--.-----_--_---------_------- 9.6
Miz, lenethoficutting edge of paraconid_-_-----------_- <<< ooo en _— 7.5
IMiplenet rot Cuttin ed CO lO LOL OL CCTM ee eee er eee 9.5
Mj, height of paraconid above base of cingulum anteriorly ----------------------------------- 9

M,, height of protoconid above base of cingulum posteriorly--_------------------------------- LGA

FELIS near ATROX Leidy

FicurE 29

Several feline foot bones in the collection appear to be much too
large and robust to belong with the upper tooth and jaw portion re-
ferred to Felis inexpectata and may represent the large North American
Pleistocene cat Felis atrox Leidy (1853). Included in this foot material
are a scapholunar and second metacarpal (fig. 29b) representing the
manus and an astragalus, caleaneum (fig. 29a), and the third (fig. 29c)
and fifth (fig. 29d) metatarsals from the pes. There is no evidence to
show that these bones are from the same individual.

These elements are much too big to belong to a large jaguar or
even an unusually large puma. The proportions are comparable to
those in the foot of a specimen of Felis tigris or F. leo. ‘The scapho-
lunar is distinctly larger than in any of the tiger or lion carpi examined
and shows a relatively deeper groove for its articulation with the
magnum. However, the second metacarpal, though equally as long
as in these large Asiatic and African cats, is somewhat slenderer, and
the proximal end does not have so large an articular facet for the
trapezoid.

52 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

The Cumberland Cave astragalus and calcaneum are of about the
same size as in an adult Felis tugris, although the posterior portion of
the calcaneum is relatively short and the proximal end is noticeably
heavy. The posterior dorsal surface of the calcaneum is not so sharply

WS SSS
a

FIGURE 29.—Felis near atror Leidy, foot bones (U.S.N.M. 12840): a, Astragalus and caleaneum, anterior
view; 6, second metacarpal, dorsal and proximal views; c, third metatarsal, dorsal and proximal views;
d, fifth metatarsal, dorsal and proximal views. One-half natural size. Cumberland Cave Pleistocene,
Maryland.

keeled as in available specimens examined of either F. tigris or F. leo.

The third metatarsal is as large and robust as in specimens of F.

tigris, but the fifth metatarsal though fully as long as in the tiger is

distinctly slenderer.
A comparison of the foot elements from Cumberland Cave with
those of Felis atroz from Rancho La Brea figured by Merriam and

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 53

Stock (1932, figs. 74, 92, 103, 106, 126, 130) shows the scapholunar, as-
tragalus, caleaneum, and third metatarsal to be markedly similar to the
corresponding elements in the California material but nearly one-third
smaller. However, their dimensions are only slightly less than those
given for a small individual in each case. The second metacarpal
and fifth metatarsal in addition to being smaller are relatively slen-
derer than in the Rancho La Brea form.

A third metacarpal in the Port Kennedy collection, referred by
Cope (1899, pp. 247-249) to F. inexpectata, compares favorably in
length with the corresponding bone of a large puma but belongs to a
foot much smaller than that indicated by the second metacarpal
from Cumberland Cave. A fourth metacarpal from Conard Fissure
bearing the number A.M.N.H. 11787, as does the type of FP. longicrus,
is as robust as in F. tigris but markedly shorter, resembling that of
a saber-tooth form.

TaspLe 15.—Measurements (in millimeters) of foot bones of Felis cf. atrox

See ee

Measurement ve Ae

Transverse width of scapholunar measured across medial portion of radial faceti ae s2s6552 44
‘Greatest length of second metacarpal-_-_.------------------------------------------------------ 94
Greatest diameter of proximal end of second metacarpal-------------------------------------- 21
Greatest transverse diameter of proximal end of second metacarpal--------------------------- 15.2
Length of caleaneum between groove for achilles tendon and CubOlGliacet=se ee eee anne 99. 5
Width of caleaneum across sustentacular and ectal facets------------------------------------- 38
Length of astragalus between center of navicular facet and most remote portion of tibial facet

on inner condyle..__.---=-------=--------------=--------------------=-=--<-=-=---------=--- 52
Greatest length of third metatarsal_-_--------------------------------------------------------- 113
Dorsoventral diameter of proximal end of third metatarsal__._.------------------------------- 29
Transverse diameter of proximal end of third metatarsal___.---------------------------------- 24
Greatest length of fifth metatarsal__---.------------------------------------------------------ 111.5
Greatest diameter of proximal end of fifth metatarsal _--_..------------------------------------ Vad

Order RODENTIA

Family SCIURIDAE

MARMOTA MONAX (Linnaeus)

In the collection from the Cumberland Cave deposit there are a
number of specimens referable to a single species of woodchuck.
They include seven nearly complete skulls, several lower jaws, and
other bones of the skeleton. The better-preserved carry the National
Museum catalog numbers 7997, 8127, 8157, 8158, and 8183.

The cheek teeth in the cave specimens average somewhat less in
size and the skulls average somewhat larger compared with a large
series of the living species, but in these and all other differences noted
they fall well within the range of individual variation of Marmota
monaz. For this reason they are referred to the living species.

54 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

CITELLUS cf. TRIDECEMLINEATUS (Mitchill)

A single specimen, an incomplete left ramus of a mandible, U.S.N.M.
no. 12054, containing the incisor (broken at alveolus), M,, and Ma,
represents the ground squirrels in the Cumberland Cave collection.
This specimen is about the size of Citellus tridecemlineatus, to which
it is provisionally referred. The two lower cheek teeth preserved
are slightly less compressed anteroposteriorly relative to their width
than is common in living specimens. ‘The absence of more complete
material prohibits a satisfactory comparison with the Recent form.

TAMIAS cf. STRIATUS (Linnaeus)

A fragmentary lower jaw, U.S.N.M. no. 12367, included in the
rodent material is recognized as belonging to a chipmunk. The
lower cheek teeth are present though badly worn, and the incisor is
broken away at the alveolar border. The specimen is not complete
posterior to the last molar. The jaw is comparable in size and pro-
portions to that of the living species, Tamas striatus. Nocharacters
were observed that would serve to distinguish the fossil from the
form now living in the same region.

SCIURUS (TAMIASCIURUS) TENUIDENS Hay
FIGuRE 30

The species Sciurus tenuidens was described by Dr. Hay (1920,
pp. 104-105, pl. 4, fig. 20) on what he believed to be an upper incisor
in a part of a skull from a cave deposit near Cavetown, Md. It was
found upon removing the matrix that the specimen is a right lower
jaw portion with the lower incisor. The cheek teeth were not pre-
served. As an inferior jaw the specimen does not possess such
marked characters distinguishing it from Sciurus hudsomcus. The
portion of the jaw preserved is rather robust, and the incisor is rela-
tively narrow transversely and of greater anteroposterior diameter
than in S. hudsonicus. Also, the mental foramen appears to be
placed lower on the side of the ramus than in the recent specimens
examined.

Sciurid material in the Cumberland Cave collection includes three
skull portions and four mandibular rami. The form represented
clearly belongs to the group Tamiasciurus but is apparently distinct
from S. hudsonicus. The mandible is more robust and possesses a
lower incisor of relatively great anteroposterior diameter and hence
presumably represents the Cavetown form, S. tenwidens. On the
basis of skull U.S.N.M. no. 8164 (fig. 30a and 6) and a nearly com-
plete left ramus of mandible U.S.N.M. no. 12050 (fig. 30c) from
Cumberland Cave, the species is redescribed.

Specific characters.—Size about that of Sciurus fremonti, larger
than the average of the S. hudsonicus group. Skull with notably
broad frontal region and with short, deep muzzle. Viewed from the

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 55

side, occiput more elevated than in the living species, giving a less
convex contour to the cranium. Anterior zygomatic region widely
expanded, with its outer rim strongly turned downward and ex-
tending forward well beyond the maxillary-premaxillary suture,
thus giving an unusually broad surface for the attachment of the
anterior portion of the masseter-lateralis. Jugal relatively deep,

FIGURE 30.—Sciurus (Tamiasciurus) tenuidens Hay, skull (U.S.N.M. no. 8164), and mandible (U.S.N.M.
no. 12050): a, Dorsal view of skull; 6, lateral view of mandible. X 14. Cumberland Cave Pleistocene,
Maryland.

about one-fourth deeper than the average in the living species. Bullae
much as in the living members of the group but more inflated back-
ward and outward, in posterior portion, giving a decidedly flatter
appearance to the posterior wall of its inner face and to the outer
face directly below the external auditory meatus. External audi-
tory meatus smaller than in living species.

Incisors more robust, with greater anteroposterior diameter than
in the living species. Cheek-tooth row of equal length witb that of
Sciurus fremonti, and of S. hudsonicus from the Fort Yukon district,
but teeth slightly narrower.

Lower jaw relatively deep, short, and massive, with broadly ex-
panded angle. Anterior portion of masseteric fossa deeper and

56 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

broader and with more robust border ridge than in specimens of
equivalent maturity.

The red-squirrel material from the Conard Fissure was referred by
Brown (1908, pp. 192-193) to Sciurus hudsonicus. Without a criti-
cal comparison of actual material it seems that the Conard Fissure
material represents a species close to Sciurus tenuidens, although
certain features, as the slightly less robust incisors, the less extreme
depth of the lower jaw angle, the apparently larger external auditory
meatus, and the more convex longitudinal profile of the posterior
portion of the cranium ally it perhaps more nearly to the living
S. fremont of the Rocky Mountain region.

TaBLE 16.—Measurements (in millimeters) of skull, mandible, and dentition of
Sciurus tenuidens and S. hudsonicus

Sciurus tenuidens Sciurus
hudsonicus
Measurement eae U.S.N.M.
no. 9197. U.S.N.M.| U.S.N.M.|} no. 151654
(type) no. 8164 | no. 12054 (Div.
Mamm.)
Length, posterior margin of incisor alveolus to condyles-_-__|_-_.----__- AQND| aes 39.5
Depth of rostrum below nasofrontal suture__----...--.-.--]----------- AZjG Alea son ae 12
Width of skull across postorbital constriction_____........_]_-------_.- 45S eee. ee 13.5
Length of diastema between alveolus for I and P4...______|_-_--__--_- ADA NE ee eae ll
Length of tooth series, P4 to M3, inclusive__..-..-...------|_...-.-___- Lael ee sae 78
Transverse: wid thiof M4. 222. 022 cos se es ee | eee ce ee asa (booed Pie
Depth ofijawbelowsPas*=22seeeese ons. Seen en cose eee aa eee ee ee | eee 8.7 8.5
Length of diastema between alveolus for I and alveolus for
Pea seae ncn eth ona sone ee ene seaen esac eoeee es eee 457 asoseeee ee 4.8 5.4
Length of tooth series from P, to M3 measured at alveolar
DOR ers == 23225 ieee See te Soe ee ee ee 1 Sloe see seo 18.1 8.2
Anteroposterior diameter of lower incisor-_.-_------------- 8:9" |eenaccc ses 3.5 3.2
Transverse diameter of lower incisor_......----.----------- 1G) |ooccecwsces 1.5 1.2
‘Transverse. width) ofiMy: <2. =t22 2 ct ae Seeeeie a. cae |e eee ee ere | eee ee as 2.2 2.2
1 Approximate.
GLAUCOMYS species

The flying squirrels are represented in the Cumberland Cave col-
lection by a single specimen, a broken lower jaw, U.S.N.M. no. 7687,
carrying three teeth, P, to M,. This fragment of jaw indicates a
species larger than Glaucomys volans, not quite so large as G. yukonen-
sis, and about the size of G. sabinus. The teeth combine characters of
these three living species, with features of their own that suggest the
specimen may represent a new species, but without more adequate
material by which to determine possible ranges of variation it is
preferred not to give it a definite assignment.

The general characters noted in the Cumberland Cave specimen are
as follows: Length of cheek teeth series P, to M.=5.3 mm; greatest
width of M,=2 mm. (G. volans length of cheek teeth P, to M.=4.2;
width of M,=1.6 mm.) P, relatively large, with deep talonid valley

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 57

that opens obliquely outward and backward through a notch that
distinctly separates the hypoconid from the outward extension of the
posterior rim of the basin. Paraconid, hypostylid, and entostylid of
P, well developed. Basins of the molars broad and evenly concave.
Space between protoconid and hypoconid relatively wide with very
distinct valleys on each side of the well developed hypostyle. Inner
rim of the cheek tooth crowns interrupted by a median, sharply
defined notch.
Family GEOMYIDAE

PLESIOTHOMOMYS POTOMACENSIS Gidley and Gazin

Ficure 31

Type.—Portion of left ramus of mandible (fig. 31), U.S.N.M. no,
8166, with all cheek teeth. Incisor not preserved beyond margin of
alveolus.

Generic characters—Anterior margin of symphysis of mandible
very abrupt. Pit on lingual side and at base of ascending ramus
shallow, as in Thomomys. Masseteric crest nearly straight and ex-
tending high on mandible anteriorly. Lower molars oval in cross
section and with enamel plates on anterior and posterior surfaces.

Specific characters.—Size near that in average adult specimens of
Thomomys fulvus. Mandible robust with deep symphysis. Incisor
relatively wide with anterior surface smooth and rounded. Lower
cheek teeth relatively narrow transversely.

The pocket-gopher material in the Cumberland Cave collection
referred to this species includes four other lower jaws, one of which,
U.S.N.M. no. 12049, possesses a complete lower dentition.

Comparison.—These specimens present several interesting charac-
ters that seem to separate them clearly from both Thomomys and
Geomys. The mandibles are remarkably deep and the symphysial
portion is turned upward rather abruptly, causing the nearly straight
anterior margin of the symphysis to form a sharp angle with the
inferior margin of the ramus. Also, the noticeably straight masseteric
crest rises anteriorly at a higher angle from the lower border of the
jaw than in living forms, making the distance between the anterior
extremity of the masseteric crest and the ventral limit of the symphysis
relatively great.

The dentition of these specimens is unique in combining characters
of both Thomomys and Geomys. The incisor has its anterior surface
rounded as in Thomomys but is relatively wide. P, has its enamel
plates as in both Thomomys and Geomys, but the columns of this
tooth are well rounded and the internal reentrant fold is compressed,
suggesting Geomys. The external fold is more open. The molars
are widely oval in shape, perhaps even less compressed antero-

13 Gidley and Gazin, 1933, pp. 354-356, fig. 7.

58 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

posteriorly than is common in Geomys. In Thomomys these teeth
have their lingual portion tightly compressed and their buccal surface
distinctly flattened and usually grooved longitudinally. However,

OMG

Wy) WY iy
fi MW

FIGURE 31.—Plesiothomomys potomacensis Gidley and Gazin: Left ramus of mandible, type specimen
(U.8.N.M. no. 8166), lateral and occlusal views. 2. Cumberland Cave Pleistocene, Maryland.

the molars in the fossil specimens have enamel developed on both
anterior and posterior surfaces as in Thomomys. In Geomys only the
posterior surface of these teeth has an enamel plate.

TABLE 17.—Measurements (in millimeters) of mandible and dentition of Plesiotho-
momys potomacensis and P. orientalis

Plesiothomomys potomacensis Fe ere
Measurement Pn at pee
no. 8166 a= tt tuo zatads
(type) (type)
Superior dentition
Length of diastema between alveoli of I and Py----.-------|----------------|---------------- 12
Anteroposterior diameter of upper incisor_-----------------|----------------|---------------- 11.8
Transverse diameter of upper incisor: - | -2-~---s-- 3 - ennnn| en nec ew ene e na | aan ae
P4, anteroposterior diameter at occlusal surface---.---------|----------------|---------------- 22
P4, transverse diameter of anterior lobe_-_------------------|----------------|---------------- 1.6
P‘, transverse diameter of posterior lobe._--..--------------|----------------|---------------- 1.8
M!, anteroposterior diameter at occlusal surface.-.---------|----------------|---------------- aba
Miitransverse diameter +2 02-2222 ate nanan a ease sens s secs | ewen tenn eae ane ae een 1.9
Mandible and inferior dentition
Distance between lowest point on dorsal surface of mandi-
ble anterior to Py and ventral extremity of symphysis-_--- 9 Sab Cassese
Length of diastema between I and Py---------------------- 18.1 S..Gh| 22 assesses
Length of lower cheek-tooth series measured at occlusal
BUTIACO Ns ne eee eee eae nee nese nea aea eee 7 136 eos sensesoeee
J anteroposterior: dlaMeterseses ses ae sane eee a eas | ae enna en 23 sees Sa soce
Ve transverse diameter. 2-22 en. Se eee eae ae yal 2.2 Neonates
Py, anteroposterior diameter at occlusal surface.----.------- 2.6 229 Seo 2o se eee
P,, transverse diameter of anterior lobe_-_-.---------------- 1.3 Taio oees see es
P4, transverse diameter of posterior lobe.------------------- 2 ae eh a
Mi, anteroposterior diameter at occlusal surface--_---------- 1.4 155) || eee
Mi, fransverseidiameter: -....-: =.) £2622. S222e- set se 2 Dr Dice. ee Saez ee
Mi, anteroposterior diameter at occlusal surface__.--------- 1.5 136i [See eW seco
Ms; transverse diameter. __-- 24-2022 ceaenconcese sees ae 1.9 2) aa ee See
Ms, anteroposterior diameter at occlusal surface-_----------- 1.5 1.6) |be doses los
Mis; transverse diameter=-----o-s.54-) ae ee leg, WSO) eee eaees

1 Approximate.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 59

Thomomys orientalis Simpson (1928, pp. 6-7) from Saber-tooth
Cave in Florida, although possessing characters that ally it to Tho-
momys, apparently should be included in the genus Plesiothomomys.
The upper incisor in the Florida specimen shows a single shallow
eroove placed well toward the inner margin of the enamel, as in some
modern species of Thomomys. Also, both P* and M? have the enamel
complete across the posterior surface as in the western pocket gopher.
However, the columns of these teeth are oval in cross section as in
Geomys and the reentrant folds in P* are not so widely open as they
are in Thomomys. Another interesting character observed in the
type of P. orientalis is that the rostrum appears to be somewhat
depressed anteriorly. This may be due in part to crushing, but the
portion of the palate between the cheek teeth and incisors does not
seem to ascend so steeply as in living species.

P. orientalis is a distinctly smaller species with smaller teeth than
P. potomacensis, corresponding more nearly in size to specimens of
Thomomys fossor.

Family CASTORIDAE

CASTOR CANADENSIS Kuhl

A single left upper molar in the Cumberland Cave collection,
U.S.N.M. no. 12056, may be so exactly matched in specimens of the
living species Castor canadensis that it must for the present be referred
to this species. Height of crown (apparently not much worn) 23 mm;
anteroposterior length, triturating surface, 6.5 mm; actual antero-
posterior diameter 6 mm, width 6.5 mm.

Family CRICETIDAE
PEROMYSCUS cf. LEUCOPUS (Rafinesque)

The deer mice are represented in the cave collection by eight frag-
mentary mandibles with incisors. Seven of the specimens have one
or more cheek teeth each. A comparison of this material with jaws of
Peromyscus leucopus from the Recent fauna of Maryland failed to
reveal any characters of diagnostic importance separating the fossil
from the living form. Although more complete material including
skulls may show important specific differences, this is not suggested
in the mandibles. Several modern species of Peromyscus, as P. mani-
culatus and P. leucopus, have remarkable geographic distribution, and
it seems not at all unlikely that their geologic range through Quater-
nary time may have been equally extensive.

NEOTOMA MAGISTER Baird #
FicurE 32

Three wood-rat mandibles, U.S.N.M. nos. 12033 (fig. 32), 12034,

and 12035, in the collection apparently represent the large Neotoma

13 Baird, 1857, p. 498.
16541—38——5

60 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

magister. The specimens compare favorably in size with the cotype
material from Carlisle Cave, Pa., being somewhat larger and deeper
jawed than in an average living specimen of N. pennsylvanica.* The
Cumberland Cave specimens are characterized by a relatively long
mandibular symphysis, robust incisors, and cheek teeth with postero-
external reentrant folds widely U-shaped and deeply pocketed at base.

FIGURE 32.—Neotoma magister Baird: Left ramus of mandible (U.S.N.M. no. 12033), lateral and occlusal
views. 2. Cumberland Cave Pleistocene, Maryland.

The Cumberland form is readily distinguished from N. floridana by
its greater size, larger incisors, and relatively longer diastema between
incisor and cheek teeth.

N. ozarkensis Brown (1908, pp. 196-197) is apparently somewhat
smaller than the Cumberland Cave wood rat, presumably nearer in
size to N. pennsylvanica.

TaBLE 18.—Measurements (in millimeters) of mandible and inferior dentition of
Neotoma magister

U.S.N.M. | U.S.N.M.

Measurement no. 12033 | no. 12034
Length of lower molar series measured at occlusal surface__-_------.-------------- 8.8 9.4
Length of lower molar series measured along alveoli-_----------------------------- 9.7 10
Mi anteroposterior length a3: 2222. = So ee ee ee 3.4 Sov
My itransverseswidth-e= = 225 2 2 he ee ee se ee eee senaee eee aee 1.9 2.1
Ms, ‘anteroposterior lensths soo 8 ea oa ee See eene ene oeasecee 2.8 3
Mo, transverse’width. : 2i0i1222252 622228 oe os case csastaaeccteeosee 2.2 a
M3; anteroposterior length =.----2--<2=-< 222s sss3— a Rot serene dee ee oscetaessas 252) 2.3
Msitransverse: width. 22.50. 222 es es ee ence en cseeseeee 1.9 1.9
Depth ofiramus'at junction of; May and "Migs ~ o= = es ee esa se neon ean 5.6 5.8

PARAHODOMYS SPELAEUS Gidley and Gazin '5

FicurREs 33, 34

Type.—Portion of the right ramus of a mandible (young individual),
U.S.N.M. no. 12040 (fig. 33), with all cheek teeth and part of alveolus
for incisor.

Generic characters —Cheek teeth in general structure as in Hodomys,
but with reentrant valleys, especially the external ones, shallower and

44 The possibility of N. pennsylvanica Stone being asynonym of N. magister Baird was noted by Rhoads,

1894, pp. 213-221.
18 Gidley and Gazin, 1933, pp. 356-357, figs. 8, 9.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 61

directed nearly at right angles to the line of the tooth row. [They are
directed more obliquely forward in Hodomys.] Anteroexternal valley
of M, and M; very shallow and absent in M;. [In Hodomys M; has a
shallow but well-defined anterior reentrant in young specimens.] Mz,
with a well-defined posterointernal reentrant valley. [This fold is
absent in Hodomys.]

As seen in mandibles U.S.N.M. nos. 12037 (fig. 34) and 12041, the
following additional characters appear as of generic importance:

FIGURE 33.—Parahodomys spelaeus Gidley and Gazin: Right ramus of mandible, type specimen (U.S.N.M,
no. 12040), lateral and occlusal views. <2. Cumberland Cave Pleistocene, Maryland.

Symphysis relatively deep. Lower branch of masseteric ridge very
prominent and sharply defined. Condyle relatively large and de-
pressed so that condylar bar is directed backward to a much greater
extent than in Hodomys and the notch between it and the coronoid
process is much more widely open.

Specific characters—Size about that of Hodomys alleni. Incisors
relatively robust, causing a more prominent swelling of the lower

FicuRE 34.—Parahodomys epelaeus Gidley and Gazin: Right ramus of mandible (U.S.N.M. no. 12037),
lateral and occlusal views. X 2. Cumberland Cave Pleistocene, Maryland.

border of the ramus than in H. allent. Diastema between incisor and
cheek teeth long. Posteroexternal reentrant folds in all molars
shallow and pocketed at base, wearing at occlusal surface in wide
open U-shaped pattern from summit to base of crown. Postero-
internal reentrant fold well developed in M3, extending somewhat
below middle of crown but disappearing in old age.

The material referred to this form includes seven lower jaws in
addition to the type, representing individuals in several stages of
maturity.

Comparison.—The genus here described is known only from lower
jaws but fortunately these reveal diagnostic characters that clearly
distinguish it from the nearest related living genera, Neotoma and

62 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Hodomys. Of these the most readily determinable characters are to
be found in the last lower cheek tooth. In old individuals, showing
well-worn teeth, this tooth, in fossil as well as in living genera, has but
two reentrant angles, one on the inner, the other on the outer face of
the crown. In Neotoma these reentrant valleys are oppositely placed
and tend to divide the crown surface into two subequal areas. In
Hodomys and Parahodomys the reentrants alternate, the inner being
well in advance of the outer one, tending to form a more or less S-
shaped enamel pattern on the triturating surface. However, the
latter genera are readily distinguished from each other by the fact
that in Parahodomys the reentrant valleys are directed almost at right
angles to the longitudinal plane of the tooth row, while in Hodomys
they slant well forward. In young individuals with slightly worn teeth,
additional distinctions are apparent in that there is present a third
reentrant fold in this posterior molar in all three genera, but this fold
is peculiarly situated in the fossil form. In Parahodomys it occurs on
the posterior internal lobe opposite the large external fold and extends
downward to a point slightly below the middle of the crown. In
Neotoma and Hodomys the third reentrant fold is on the anterior lobe
of the outer side. It occurs only very near the summit of the tooth in
Neotoma, but it extends down to about the middle of the tooth crown in
Hodomys.

TapLE 19.—Measurements (in millimeters) of mandible and inferior dentition of
Parahodomys spelacus and Hodomys alleni

Parahodomys spelaeus Hodomys allenit
a USMM. | ews. | Won
no. 12040 Yona no. 91525
(type) no. 12037 | (Biol. Surv.)
See eee ee
Total length of jaw, condyle to incisor border-.------------- eee ae ee 30 28.7
Depth of symphysis-.--------------------------------------]--------------- 5.4 4
Depth of ramus at point between M2 and Ms3--------------|--------------- 7.3 6.5
Height of condyle above level of cheek tooth row-----------|--------------- 7 10
Length of cheek tooth row measured at alveolar borders---- 9.5 9.5 10.8
Mi, anteroposterior diameter---...------------------------- SON Eee eee 3.6
M,, transverse diameter ----------------------------------- Oe ee eae | 2.3
Mz, anteroposterior diameter ---...------------------------- 2.8 3 3.5
M:,, transverse diameter ---.-------------------------------- 2.1 2.4 2.5
M3, anteroposterior diameter - --.-------------------------- 2.4 2.6 2.9
Ms, transverse diameter- --------.------------------------- 2 2 2.1

eee eee —

SYNAPTOMYS cf. COOPERI Baird

The anterior portion of a single lower jaw, U.S.N.M. no. 7774, in
the Cumberland Cave collection, carrying the incisor and first molar,
represents a species of Synaptomys that corresponds in size and is not
otherwise clearly distinguishable from S. coopert. In the many speci-
mens of the living species of this genus examined the triangles of the

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 63

molars are more tightly closed than in the fossil specimen. This is
the only difference observed, but if it could be verified in a large num-
ber of specimens it might constitute a species distinction.

SYNAPTOMYS (MICTOMYS) species

The subgenus Mictomys is also represented in the Cumberland Cave
collection by a single specimen, a broken lower jaw, U.S.N.M. no.
7773, with complete dentition. No characters were observed by
which the fossil form could be distinguished from the living species,
S. (M.) borealis. This species is currently recognized to contain all
the races within the subgenus Mictomys. However, all the lemming
mice of this group are at present Canadian and Alaskan in distribu-
tion (Howell, 1927, p. 19), extending down into northern Washington
and northern New Hampshire in the United States. Although the
fossil appears to demonstrate the presence of this modern boreal spe-
cies in the Pleistocene of Maryland, its occurrence so far outside of the
present distribution suggests that a distinct species may be represented.
Additional importance is to be attached to this occurrence in furnishing
the first Pleistocene record of Mictomys (Gidley, 1913b, p. 96).

MICROTUS (or PITYMYS?) cf. INVOLUTUS (Cope)

FiGurE 35

The voles are very sparsely represented in the Cumberland Cave
collection. Seven fragmentary specimens, one a part of a palate with
the two anterior cheek teeth on the right side, the others broken
mandibles, comprise the lot.

The material is clearly microtine in character but is apparently
distinct from North American living species. The root portion of the
lower incisor in the Cumberland Cave form passes between the lower
portions of the penultimate and ultimate molars as in voles, and the
cheek teeth are without roots in those specimens where this character
can be observed. However, the pattern of the first lower cheek tooth is
somewhat simpler than in the living pine-mice of the region. This
tooth, presumably M,, is characterized by a posterior loop followed
forward by three closed triangles, a pair of confluent triangles opening
into an anterior loop. In these characters the fossil jaw resembles
Pitymys but unlike the form living in the same region at the present
time, P. pinetorum, the anterior loop of M;, in three of the six jaws is
without reentrant angles. Two specimens have shallow reentrant
angles on both inner and outer surfaces, nearly as in P. pinetorum,
and one has a shallow groove on the antero-internal surface of the
anterior loop (fig. 35). Furthermore, the posterior surfaces of the
triangles in the lower cheek teeth are slightly more convex, andisola-
tion of the triangles is not so marked as in specimens of P. pinetorum.

1¢ Cope, 1871, pp. 89-90, fig. 16.

64 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Also M3; is somewhat more robust than is common in the Recent
species.

Microtus pennsylvanicus, the typical microtine form living in Mary-
land today, exhibits a more complex pattern in the first lower tooth.

FicureE 35.—Microtus (or Pitymys?) cf. involutus (Cope): Right ramus of mandible (U.S.N.M. no. 12386),
lateral and occlusal views (pattern of M3 taken from mandible U.S.N.M. no. 7772). 3. Cumberland
Cave Pleistocene, Maryland.

Presumably M. pennsylvanicus is to be regarded as possessing a more
advanced type of dentition.

TaBLE 20.—Measurements (in millimeters) of dentition of Microtus(?) cf.
involutus and M. dideltus

Microtus(2) ef. involutus ee
Measurement T8s#| 0.8,°}Us8s (U8. | AcE ALM)
N.M.|N.M.|N.M.|N.M.| N.H. | N.H. Ol

no. no. no. no. no. no. 8694
7686 | 12602 | 7772 | 12368 | 8699a | 8695 (type)

Superior dentition
T,anteroposterior diameter.ss-.cas-2ees eer eeeeoces oe 12 Ssccce alte sie Se be a Eee oe | ban oe eeeeroe
i; transverse! diameter: =222-. = 2-2 252-522226 2. -ses2—52 Le [eect ot boos te Ss Sets ot sete ee ao ae
M!, anteroposterior diameter !__.-.....-.------------ 2. 2h Soc arene |ot eee | Se owes 24 jot oe
M1. transverse Giameters.s.2- 2 .2s222525-2-<-cesecces Veal eos elesce kan lan oese al oence se 143)'|Ss22255
M,, anteroposterior diameter__..........------------ D6) sees sce sok ces eeeececb ee 18s ota
M3" transverse: diameters. 2- sos ae aneeeoassasoeees Li: Abeseeesel bese dsehee sccu laste =e L2hfsssone -
M3 anteroposterior diameter-s-c2o2sco2- ee see ee sas | sao cee joc ae cre Coeeeee | peeeees loaeenes LO icone =
Ms; transverse diamoters22222.2csh soc acs cocoa nen c= lobes ces) sense eth cote aes ae ae |eoeeees P21 ccee, 2
Inferior dentition

Z,janteroposterior diameters 222 oe. ses eee ee een eee 1,2 1,2 1330 tosdsccs|souuc ols
I, transverse diameter. 2222-22 25-2 ses sk 2 ese e ee es se|s2ess=5 0.9 1 1 Lee eesee 91.5

My; anteroposterior diameter22-.1—- =. 22. sce coe anne |e ates 2.6 2.5 at 26 lssese 3
Mia transverse: diameters22-2 <2 ose ooo oe eee as |oe ee 1d 1 1 1.4 jAcescce 1.2
Ms; anteroposterior diameter-s2--2-saeenscanseessen|eeaaee 1.5 1.5 1G 48020 |e eee eeeeees
Ms, transverse diameteris:2 222-28 cece een wenn eet 0.9 0.9 1 Jabatss Clee Sete ee
M3; anteroposterior diamoter-c: 224. co- sas aa=- sos |o=ee see 1.5 1,43) css-5 173. Joo 33S] sees 3
IMizs trans Verse dismMetebesss=se sss ees ae eee eee anne peeee 0.8 OT Rencece ON2|Poaeceo|Sassees

Length of lower cheek-tooth series measured at

occlusal jsurface...% .< 22. 4-<..s22eeeo Seed ens on |Saeneee Bad Blleccenns 95.6) | scsc-es -tcesse

1 Anteroposterior dimensions taken at occlusal surface.

21 Dimension as given by Cope but not verified.

4A pproximate.

The Cumberland Cave form corresponds closely in size to the type
of Microtus? involutus (Cope) from the Pleistocene of Pennsylvania.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 65

The enamel pattern of the lower cheek teeth is markedly similar in
the two forms. It is important to note that the anterior triangles in
the first lower cheek tooth of the type are confluent as in Pitymys, not
as in Cope’s illustration. This error in the drawing tends to exagger-
ate the appearance of the anterior loop, which in reality is not greatly
different from that in some of our specimens. Cope noted the re-
semblance between his form and P. pinetorum, but apparently the
Pleistocene form is distinct from the living species in exhibiting a
slightly simpler enamel pattern.

The teeth in Microtus(?) speothen (Cope, 1871, pp. 87-88, fig. 13)
and M.(?) dideltus (Cope, 1871, p. 89, fig. 15) are distinctly larger
than in the Cumberland Cave form. In M.(?) speothen the enamel
pattern in the first lower cheek tooth is of a different type than that
in Pitymys. This tooth in M.(?) dideltus, while showing confluent
anterior triangles as in Pitymys, has an anterior loop that is narrow
and elongate, and the portion of the tooth posterior to the loop is
relatively wide.

As noted by G. S. Miller, Jr. (1896, p. 59), the anterior lower cheek
tooth of Pitymys is not different from that in Pedomys, or Microtus
ochrogaster, living in the Mississippi Valley. Hence, reference of the
Cumberland fossil to the genus Pitymys in the absence of satisfactory
skull material cannot be made with certainty.

ONDATRA cf. ANNECTENS (Brown)

FIGURE 36

A very small species of Ondatra is represented in the Cumberland
Cave material by a broken mandibular ramus (fig. 36), U.S.N.M.
no. 12044, containing all the teeth. In size this specimen agrees closely
with the type of the Conard Fissure species, O. annectens, being only
slightly smaller and possessing a somewhat narrower incisor. It
agrees with O. annectens in most other important details observed and
is therefore provisionally referred to this species.

It was suggested by Hollister (1911, pp. 33-34) that Brown had
wrongly interpreted some of the characters described. Brown con-
sidered the Conard Fissure species as ‘‘an intermediate type connecting
Fiber (=Ondatra) with Microtus through the subgenus Neofiber.””
The type of 0. annectens as observed by Hollister indicates “a species
close to existing forms in everything except size’, and he stated that
the “anterior loop of first lower molar is more deeply cut by the
reentrant angles than in any specimen of an existing species examined.”
If these anterior reentrants are to be considered within the loop, as
implied by Hollister, then the number of reentrants in M, posterior
to the loop on each side appears to be one less than in 0. zibethica,
approaching the condition seen in Neofiber alleni. In either case the

17 Brown, 1908, p. 197.

66 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

columnar portion anterior to the fifth reentrant, counting forward
along the inner side of the tooth, is not so well developed in O. annec-
tens as in O. zibethica. Advanced wear in M, of O. zibethica may show
this column or loop to be without reentrants although well developed.

In the description of O. annectens Brown called attention to the
markedly reduced condition of the anteroexternal column of M3.
This column is not more reduced in the Cumberland Cave specimen
than in certain specimens of O. zibethica. In Neofiber this column is
greatly reduced.

In both the Cumberland Cave and Conard Fissure jaws the surfaces
for the attachment of the masseter muscles are nearly alike but differ
slightly from those of the living species of Neofiber and Ondatra.

CRE ND

FIGURE 36.—Ondatra cf. annectens (Brown): Left ramus of mandible (U.S.N.M. no. 12044), lateral and

occlusal views. XX 2. Cumberland Cave Pleistocene, Maryland.
The masseteric crest in general is bulging and sharply keeled, and there
is a well-defined groove marking the posterior margin of the area for
attachment of the anterior masseter medialis. The distinguishing
feature, however, lies in the position of this sulcus, which in O. annec-
tens has its origin much farther back from the anterior end of the mas-
seteric ridge from where it curves sharply upward, leaving a definitely
wider margin between it and the anterior border of the ascending
ramus. It is further noted in the Conard Fissure jaw that the shaft
leading to the condyle carries a fairly well developed sulcus on its
anterior face nearly as in Ondatra zibethica. This sulcus is incipient
or absent in Neofiber.

TABLE 21.—Measurements (in millimeters) of inferior dentition of Ondatra ef.

annectens

A.M.N.H.

Measurement ue Perk no. 12424

ee (type)

Length of cheek-tooth series measured at alveolar margins_--_------------------- 111.5 12
Length of cheek-tooth series measured along occlusal surface____.---------------- 10.5 10.9
Mi, anteroposterior length at occlusal surface__......---.------------------------ 5.1 5.4
Mistransverse: width=. \- 2... $2 ee ee ee ek oe ee eee 2.1 2.2
M2, anteroposterior length at occlusal surface__...--...----.--------------------- 27 2.8

Mostransverse: widthisscs. 23. ote re ne en eR i ee ae 2 2
Ms, anteroposterior length at occlusal surface._-..---.--------------------------- ONT a7
Mis} transversetwidth:_<¢ Fs. - 2 seh be, re A eg ee eles peek eS 1.6 1.6

1 Approximate.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 67

O. annectens seems to stand as a well-defined species of Ondatra but
is presumably less advanced than the living species. Neofiber may
be considered as still less advanced in the development of the pattern
reached in M,, although more advanced in degree of hypsodonty.

Family ZAPODIDAE

ZAPUS species

The genus Zapus is represented in the Cumberland Cave material
by part of the right ramus of a mandible, U.S.N.M. no. 7777, carrying
all the cheek teeth. This jaw represents an old individual in which
the teeth are considerably worn. They nevertheless present the
characters peculiar to the genus to which the specimen is referred and
show certain peculiarities that prevent assigning the form to any
known species. The material, however, is not sufficiently complete
to warrant establishing a new species.

The tooth crowns of M, and M; are relatively shorter and the an-
terior main lobe of M, is relatively wide as compared with those
examined of a large series of specimens of the living species.

NAPAEOZAPUS cf. INSIGNIS (Miller)

A second lower jaw, U.S.N.M. no. 12366, of a jumping mouse in the
collection is recognized as belonging to the woodland form Napaeo-
zapus. This jaw fragment is from the left side and has the incisor and
last two molars preserved. M, is represented by the alveoli and root
portions. The jaw is distinctly larger and has larger teeth than the
specimen referred to Zapus. Moreover, the two molar teeth show a
tightly folded, complex pattern on the occlusal surface, which re-
sembles Napaeozapus more closely than Zapus.

The fossil compares favorably in proportions with specimens of the
living species, N. insignis, differing in having a slightly more robust
incisor, which appears to be somewhat less curved than is common in
the modern form. Also, the alveoli for M, suggest that this tooth may
have been a little longer anteroposteriorly than in N. insignis.

Family ERETHIZONTIDAE

ERETHIZON cf. DORSATUM (Linnaeus)

Figure 37

The porcupine is represented in the Cumberland Cave collection by
about 15 determinable specimens, including the greater part of 3
skulls and 12 broken lower jaws.

In North America, according to Miller’s list of Recent mammals
(Miller, 1924), there are two living species of Hrethizon now recog-
nized: E. dorsatum and E. epixanthum. 'The former comprises two
subspecies and the latter five subspecies. Apparently the distinction
between the two full species is based solely on color differences in the

68 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

pelage. In an examination of a series of Recent skulls and jaws in
the collections of the U. S. National Museum and U. S. Biological
Survey, it was found that they show a wide range of individual and
sex variations and that it was apparently not feasible to formulate a
series of definite characters that would satisfactorily separate the
two species.

The specimens from Cumberland Cave likewise show considerable
variation but apparently not exceeding in any important details the
limits of individual variation seen in modern specimens.

FIGURE 37.—Erethizon cf. dorsatum (Linnaeus): a, Skull (U.S.N.M. 7996), lateral view; 6, mandible (U.S.
N.M. no. 7672), lateral and occlusal views. Two-thirds natural size. Cumberland Cave Pleistocene,
Maryland.

A remarkably large skull (fig. 37a), U.S.N.M. no. 7996, in the cave
collection is characterized by a noticeably broad frontal region, wide
palate, and particularly broad incisors. The incisors appear to be
slightly wider than in the Recent specimens examined, although other
proportions characterizing this skull seem to be duplicated in some
of the larger of the modern specimens.

Order LAGOMORPHA
Family OCHOTONIDAE

OCHOTONA species

Six fragmentary mandibles and three maxillary portions, all carry-
ing teeth, are recognized as belonging to a species of Ochotona. With
this limited material no important differences can be cited separating

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 69

it from the living species of North America, nor can it be assigned to
any modern form with certainty.

The form corresponds favorably in size with specimens of both
O. princeps and O. schisticeps. Although the dentition is closely the
same as in these Recent species, the styles or columns on the inner and
outer surfaces of the lower molariform teeth in a couple of the fossil
mandibles are slightly closer together than is common in North
American Qchotona. This condition sharpens or narrows somewhat
the inner and outer reentrant angles.

The presence of Ochotona in the fauna is significant inasmuch as the
North American species are found living only in the western and
northwestern part of the country. The present occurrence shows the
pika to have been present in the Appalachian region in Pleistocene
time.

The fossil record of Ochotona is very scanty. A form described by
Cope (1871, pp. 93-94) as Praotherium palatinum from the Port
Kennedy deposit may possibly be Ochotona. The Cumberland Cave
specimens, however, do not conform to the description given by Cope.
The upper molariform teeth of the Maryland Ochotona show a pro-
nounced external groove as in the living species and an enamel ridge
divides the occlusal surface of each tooth. Furthermore, the first,
second, and last teeth of the cheek series are structurally distinct from
one another and from the two intermediate teeth, as in living pikas.
According to Cope the four upper teeth preserved in the tooth row of
P. palatinum are structurally alike; also, their external surfaces are
not markedly grooved and the occlusal surface of each does not show
a distinct median ridge.

Family LEPORIDAE

LEPUS cf. AMERICANUS Erxleben

Hares are well represented in the cave collection. At least 10 skull
portions and 19 lower jaws are included in the material. A noticeable
range in size and proportions of skulls, jaws, and teeth exists, though
apparently within the limits of individual variation. No persistent
characters of importance were noted distinguishing the fossil from the
living hare, L. americanus. However, it is possible that some of the
smaller, more incomplete lower jaws may represent Sylvilagus, but
none of the skull portions shows the distinct interparietal and postor-
bital processes formed as in Sylvilagus.

70 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

Order PROBOSCIDEA
Family MASTODONTIDAE

MAMMUT cf. AMERICANUM (Kerr)

A mastodon is represented in the Cumberland Cave collection by
four juvenile teeth and a tibia without epiphyses. The teeth are
apparently the second, third, and fourth left and the third right of the
lower-jaw series.

There is nothing to distinguish these scanty remains from the
typical Mammut americanum.

Order PERISSODACTYLA
Family EQUIDAE

EQUUS species

The genus Hquus is represented in the Cumberland Cave collection
by only three teeth and three toe bones. The teeth, an upper and a
lower of one individual and an upper of a second individual, are those
of juveniles; they represent the milk dentition. The toe bones are
two first phalanges and a metacarpal, representing adult individuals.
These specimens belong to one of the large species of eastern horse, as
E. complicatus or FE. pectinatus, but on this incomplete material an
attempt to determine the exact species represented would be no more
than guess work, based on size alone.

One peculiarity of the foot bones may be mentioned. Both the
metacarpal and phalanges are relatively wide for their length and
depth as compared with the western species of Pleistocene horses,
suggesting a species with short stocky limbs.

Family TAPIRIDAE

TAPIRUS species

Figure 38

Two last lower molars (see fig. 38) of the right side belonging to
adult individuals, and a fragment of a left upper premolar of a third
and much younger individual, represent the tapirs in the Cumberland
Cave collection.

These teeth indicate a species somewhat larger than the living tapirs
of South and Central America and the Florida Pleistocene tapir, 7’.
veroensis Sellards, but not quite so large as T. haysii Leidy. Also they
are of narrower proportions than the corresponding tooth in the Port
Kennedy specimen, which has been assigned to 7. haysii. In astudy of
lower teeth of a series of the living species of tapirs, the proportions

PLATE 10

BULLETIN 171

U. S. NATIONAL MUSEUM

‘wnosnyy [PUOHEN *S “f)
Ul poUqiyXy ‘sAVD puvjIoquINny Woy (Ad] pry SISUAPUDILIQUNI SN uUosKy] q) Areooad Jo uojaays poyunojy

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE Te

of length to width seem to run fairly constant; hence the extreme
narrowness of the molars from the Cumberland Cave suggest they

FIGURE 38.—Tapirus sp.: Last lower molar (U.S.N.M. no. 7667), lateral and occlusal views. Natural
size. Cumberland Cave Pleistocene, Maryland.

may represent an undescribed species. But a more complete dental:
series is necessary to confirm such a conclusion certainly.

Order ARTIODACTYLA
Family TAYASSUIDAE

PLATYGONUS CUMBERLANDENSIS Gidley
Fiaures 39-45; Puatre 10

Synonym: Platygonus intermedius GIDLEY.

The type of this species is a nearly complete skull (figs. 39-41),
U.S.N.M. no. 8146, including all the cheek teeth but without canines
or incisors, and the symphysial portion and left ramus of mandible
with inferior cheek teeth and both canines. A nearly complete skele-
ton of an adult female, U.S.N.M. no. 8200, was designated as a
paratype.

The peccary skeleton (see pl. 10) has been mounted and is on ex-
hibition with the bear and wolverine skeletons also from the cave.
The skull and mandible belonging to this specimen are slightly crushed
but required no restoration. Nearly all the elements of the vertebral
column are preserved; only the tips of the spines and processes needed.
modeling. Many of the ribs were preserved and much of the sternum.
The pectoral and pelvic girdles were badly fractured and required
some patching. The fore limbs and feet are complete except for the
distal portion of the right ulna and one left and two right carpals,
which are represented in plaster. The hind limbs are also nearly

18 Gidley, 1920b, pp. 659-673.

2 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

entire; only the left required restoration. All the parts of the hind
feet are represented, but in this case one or two elements have been
substituted from the general collection.

Peccaries were one of the commonest forms found in the cave, ex-
ceeding even the black bears in quantity of material. There are 22
skulls more than half complete, 12 of which are nearly entire. Seven-
teen mandibles including the greater part of both rami are preserved.
In addition to the skeleton described above, there is also an articulated
hind foot (fig. 45) and a large number of isolated vertebrae and limb
and foot bones.

The following characters were given by Gidley as distinguishing P.
cumberlandensis from other Pleistocene species of Platygonus:

A large species, nearly equalling P. vetus in size. Length of cheek-tooth series,
type (male) 94 mm.; paratype (female), 87 mm. Differs also from P. vetus in
having a greater relative length of the molars in animals of corresponding age,
and in the possession in the upper molars of well-developed intermediary cusps
and lophs, which connect at their bases the two principal cross lophs, as in P.
compressus and P. leptorhinus. It differs from both these last named species in
the much larger size of the skull, in which there is a relatively higher and more
backwardly produced inion and a more strongly developed expansion of the
zygoma, the latter being about one and one-third times the vertical diameter of
the orbit in the females, and two or more times this diameter in the males.

Apparently the most distinctive features of this form are the elon-
gate skull, the marked backward projection of the inion accompanied
by a decided backward-sloping occiput, and the unusual extension of
the zygoma downward and forward from the orbit. The develop-
ment of the zygoma varies considerably with the age of the individual
and between sexes, but it seems to have progressed farther in this
species than in others as known.

A second species described by Gidley as Platygonus intermedius (figs.
42-44) does not appear to possess sufficiently well defined characters
to justify distinct recognition. The relative sizes of teeth and skull
and the length of the diastema between canine and second premolar
are not persistent enough throughout the series of specimens to war-
rant this specific division.

A comparison of the limb and foot material of P. cumberlandensis
with that described by Leidy (1889a) as belonging to P. compressus
and that for P. leptorhinus by Williston (1894) disclosed important
characters in the Cumberland Cave form. P. cumberlandensis pos-
sesses longer and heavier limb bones in keeping with the larger and
more elongate skull in this form than in either P. compressus or P. lep-
torhinus. This may be noticed even in the relatively slightly built
female skeleton. On the other hand, the foot bones, particularly the
metacarpals and metatarsals, though usually as large and in some
instances larger than in the two earlier described species, are relatively
short in proportion to the limb elements. This is clearly illustrated
in the proportions of the fore and hind limbs of the articulated skeleton.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE

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PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE

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FIGURE 42.—Platygonus cumberlandensis Gidley: Skull, type of P. intermedius Gidley (U. S.N.M. no. 8148), lateral view.

Cumberland Cave Pleistocene, Maryland.

natural size.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE TE

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Figure 43.—Platygonus cumberlandensis Gidley: Skull, type of P. intermedius Gidley (U.S.N.M. no. 8148), ventral view. One-half natural size.

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PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 79

P. cumberlandensis presents even more contrasting features when
compared with living members of the Tayassuidae. In many charac-
ters of the skull the fossil form has progressed considerably beyond
species of Tayassu and Pecari, most notably in size and elongation
and in the exaggeration of the jugal. The dentition, however, as in

FiIGuRE 45.—Platygonus cumberlandensis Gidley: Left hind foot (U.S.N.M. no. 7690), anterior view.
One-half natural size. Cumberland Cave Pleistocene, Maryland.

other species of Platygonus has developed along a somewhat different
line, the molars being more nearly lophodont than in modern peccaries.
The premolars in Platygonus are less progressive than in either of the
living genera but approach more nearly the development in Pecari
angulatus. More adequate descriptions of Platygonus as compared to
other genera include those of Leidy (1889), Williston (1894), Hay
(1914, pp. 212-228), Gidley (1920), and Rusconi (1930).

80 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

TABLE 22.—Measuremenis (in millimeters) of skull, mandible, and dentition of
Platygonus cumberlandensis

U.S.N.M. | U.S.N.M.
Measurement no. 8146 no. 8200
(type) (paratype)

Skull
IDxtreme length Of Skt sae ee oe a ee oe re ee eee 420 382
Total basal length measured from condylar notch_----------.-------------------- 360 320
Posterior border of orbit to posterior border of inion----.------------------------ 125 92
Anteroposterior diameter Of OLDit=2=--— == s—neee see a= aan ae eee eee 140 38
Anterior border of orbit to extreme end of premaxillary._..-..------------------- 270 250
Depthotskulliaticond ylessascees aa ae ane ae eee ee eee cee aan ee 140 135
Pepthiof skull’atigienoid' fossas2 sss =e ono ste ann ane nne ene e eee 1130 125
Greatest breadth of zygomatic expansion below orbit--.-.-----.----------------- 84 55
Greatest: width of skull‘acrossizyzomace= --- 222-52" oie aio ae ane See 230 1168
Widthiofifaceatmiddla of orbits. eee ene eee een enone eee 135 114
Width ofiface at postorbital processéSs=- 9 = seen an aaa nena nen anaes on aeneee 140 125
Width of face'above :infraorbital foramen- <2--<=2-=-2--==2 2° o-oo cee i eee eens
Widthiofipalate:between canines <s2a2. 2.6 eee nnn eee een eeeaes 50 50
Widthrof}palateitustanterior tove?=--2--— asses ane nee oes ae ae ae 45 36
Width of palate between second premolars--.-.----.------------------------------ 29 21
‘Width of palate bet ween\second: molars-_----222==*=- 222-5 _-- <5 <=" eee 22 23
Mistance: across first Paik Ol MOlArS sae see re ee ee eee 61 59
Tength of:diastema between. © and tlt. 222-2 ete sen ne eee na nee anne neeee ee 26 26
Tenethofdiastemay behind\canine==*-2- -- saeco sec one aes naan eee eee 65 71

Superior dentition

‘Totaljlengthiofupper dental seriess.---e2=. == ean oa eee eae eee eee 214 209
Wength ofupper, cheek tooth: series 3-2 2 oe an een oe enaeee ean cee nae 94.5 86. &
Lengthiof troper: premolar serless- -22-2- 2----se- eset a sane e aan meee eee 35. 2 34
mM anteroposterior diamoter::..--o----s.cseenan oe - sn eta ene nas sen e ne eaeen | aameeeeanees 1l
Fl TANS Vers Gia eCCl see tes aes ea ee ne ee ee ee eee 837i le csceecceens
Ts anteroposterion Gia Mevel sos essa eases ane eee nee nee | eee eee 8
T2htransyerse GisMetel- 2 --2enes sae eae ae ee eee ae nee een eee een eee 6.3
C;anteroposterior diameter... <2 --2-2=-5-05- ose ans ear ee ae es coneneseesesenane== 120 15
CG; transverse: diameter = 2226-552. ean ae ee eae eee eaceteseneas 12 10.4
Ps anteroposterion diam eteha ose) sae ee a ae ae a a eee cenn eee anne acne Lb y/ ll
ip? transverse) ism Gtetes- seo) ace sone eee eee eee ee en oae seen eee 11.3 10.6
Paanteroposterior: dismeter=..2- sa. a5 2-2 ae ee ee nce neneeae aaa eenaee tea 11
‘Pi transverse diametersa-2a-cosce re case ne eee oe ae eee ee een anaes eennae 13.5 13.3
Ps anteroposterior diameter. .-- -ss-5 22-2 oo tee eee nn ae eee eee meen enseneeandane LRT, 11.6
‘Pitransverse diameter:=-- 2252-22-05 22-2 eee eee een ee oes 14.3 14
Mi anteroposterior Gismotere. = 222] ne osne eae eee ee ao eee eee 15.2 12. &
Mil transverse diameter: cca. -ac cosas a oat re eee a oe Sena eneea 15 16
M3; anteroposterior diameters s-.--.ac22-4 esas soso ee ee en eee en nan eeeecanes 19.6 18
M3; transverse;diameter.-ccse-scee eae ee eee eee ena nee ae ae eae eee ee eee 18 16
M3; anteroposterioridiameters.- ..- --22 2222 sacecaen noes sacs a eet oceeane noc eecem== 24.5 20. 6
IMs: transverseGiamM etensnnc. oe some ee ee ee Se ae ee nee eee 18, 2 17.
Mandible
Total length of lower jaw, condyle to'tip of-incisors 2-2 822222 —- <2 2 noe enon soeenen aaa ce 268
MepthroPlower iawiat Wis oess sense ee eee ane ee eee eee eee 52
Depthof jaw:at coronoids process = ase wee = sea ee ee ee | eee eee 115
Depthrofilower jawiat/ cond yle-tos< sooo ae ee ee ee ee ee een een eee 90
Width between'alveoli of: lower/canines=<- =. - 95. e sae eee eae 20 22
Depth'of'symphysis‘in median liners 2. = ee anes ne eee een nnn 31 30
Depth of symphysis,at mental foramen.<=-2..==--2-22----522-==---=-<--------. <= 48 45
Length of symphysis in front (in straight line).---==2------+=-------.-=---<---2-- 100 100
Length of diastema between Isiand’ ©2223 222 eset ee eee eae e ee oe 4 9
Length of diastema behind canine (straight line) -.------------------------------ 176 75

1 Approximate.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 81

TABLE 22.— Measurements (in millimeters) of skull, mandible, and dentition of
Platygonus cumberlandensis—Continued

U.S.N.M. | U.S.N.M.
Measurement no. 8146 no. 8200
(type ) | (paratype)

Inferior dentition

Totalilenethiot lower dentaliseries= 222-2. 22 oa Ska eno eee Nie oe eee ane 208 201
iRengthioiiower cheek tooth Series == 22-22. oo ence nee nec eeaeasseoeseoecee 96 90
enethiotlower premolar SOMess-- 222. 22 so ee eee e ease cece ace neces eee eaters 35. 2 34
Tre anteroposterion GIAMCLOr= = sas s2= sen son se tee abe poe ce eae an eee nee ee enn | Seececceeene 6
NW ETANS Verse GISMGlCl. o= sass ana a= canes sace ce ee een ace wer none e eee eee naceensn | poh ranennass (Aes
Ts vanteroposterior didimeter:.-522222222525- sss esc ee eset sce ane nee cee ce ese eas|anaecenee ene 6.4
aS LLATISVCISO) GIAINOLED = oe aan sa een ee emer e neem ene an ante rennemne | seeeeaen esas 6
@TAanteroposteriOn GiniNe tele ea === eee eae ae ne nee eae 16.5 13
Se LTOTSVeNnSe GisING LOD sane ese ea ae eee a ee ene eee 13.4 10.5
Pesanteroposterion GiaMeterss- = =22- 22+ ass 2 oon se nese ane aoe neces eenene ee 19,5 10
SPaitLANSV.CESG GiginOlel ans en sense nena eee ee an ene ena ne eee 8 7.5
PenariterOpostorior Glametel= 220-2225 atone naan nee nn eee anno cecenee 12.8 11.6
IPaitransverse Gis Metoknss sss eee ne eae re ee a eee eee een een eee 10 9
Pa ;anteroposterion GiaMeter..- 9 == -s--- ass snec coe aon ae ea ea ee eee een anon acenoneee 12.8 11.6
FE asi E SFA CES 1 ERT LG Nae 11.7 11.5
Mit an teropos terion Gis Metel= === 6-24. en oe ne a ae a een nen ecoeenee ae 114 12.6
Mi niTansVCrseld AWC lCh sass a eee ee eee eee ean 13 11.8
iar iti LCLODOSLET Ors (at OLOl = a eee ee tae ae en eee aoe ee 20 18
Ma ai rans Verses (Ole esae eae oe eee nan e eee aE en eenes ae eeee 15.5 14
iat An LeLODOSLELION CIS IO CON sae ee eee anne nee 25. 5 22. 8
IMs ntranis Verse GlametOne ase. a = onsen ene eee ee eee eee aaneeeoeens 16 15

1 Approximate.

TaBLE 23.— Measurements (in millimeters) of skeleton of Platygonus cumber-

landensis
U.S.N.M.
Measurement no. 8200
(paratype)
Vertebrae
Length of vertebral column from atlas to distal end of sacrum-_-.-.-------------------------- 920:
Length of cervical series of vertebrae_--...--.-.----------------------------------------------- 220:
length of/dorsal'series of vertebrae =~. 9 = an nn oe eee aan 420
Length of lumbar series of vertebrae._-.-------------------.._~---------------- = -=---=--=----- 130
MEO P EEN Of, SACEUI eee oe ee ne a nee eae 150
iGreatestawic taf Glas sete ae ee ee eee eae ee oe ean ee nena onaenaan a= 108
Width otcondy lor facets Of ables massa eee eee ee ee een een nee eaa eae nen mann 60
CUA CLE TOT ANT A LeLa COLSTON UL AS ee oe ee a ee een Ree ee ease 53
Length of axis, exclusive of odontoid process-_-_-_-.-------------------------------------------- 44
TETG TE HG tS OOO 2X08 oe ee ee ee ee ee 30
Heichtiof spine.of first dorsal eee eee nee eee o ae eee aan ana oe= 140
eight of spine of.forrth) dorsal =. — —- ee ee ce em 120
Heieht of spine of eighth dorsal]. a ee ee oe nes ane een 70
Height of spine of second Jumbar-_-_----------------------2-----------------=------- == =5--- 30
Height of spine of last lumbar-__----...------------------------------------------------------- 25
Leneth of nine caudal vertebrae-_--------------------_---__--___- 135
Anterior limb
BIAS TAN SPS (Cox OT a oes ee ere a 220
Greatest width of blade of'scapuls.. 222 e ee oe ee oer eet ee enh ean 110
NGG LOS twp AChE Fiucots uit Cun) ear face a ee ee ee eee ea oi 33
Height of spine of scapula: 22--=2--22 256225222 = kn ew nn wee nen a en nnn wn=== 35

Wength of humerus.-...---.-..22-22---2--+---2----2-------2-222--2---------+2---------=------ 220

82 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

TaBLE 23.—Measurements (in millimeters) of skeleton of Platygonus cumber-
landensis—Continued

U.S.N.M.
Measurement no. 8290
(paratype)
Anterior limb—Continued
Width: of proximal articular faceiof humerus! 22 = Sees ee ne eee 40
Width of distal articular face of humerus. 2---22--- 222 == ee ee 34
Anteroposterior diameter of shaft just below deltoid tubercle__-.....--..--------------------- 39
Greatest length of ulna___------------ pier ee ee te a2 § Sea eae aoe ee oe ee ee ee ae 240
Greatestleneth: of raditis22. 2<-- 2A 32 eet oon ateeue eee ee eee ee 170
Transyerseidlamoeter Of SifMOid! 10SSA=2= a a a oe he a a ee ee ee eee 35
Transverse Giameterof.distal face ofradius: ——-=- 222222222 28 oe ean ea eae eee esas eee 41
east: width: of conjoined! bOneCS = 5-2 a2 a ne ee Sere ee ee ee eee 33
‘Transverse diameter Of Canpusc2 2 = sc2s. = aaa ee ee ee ee eee eee 37
Teng th. of/carpus; insid@: 2222-2 <2. 5222-5 s ee eee eee nee one oo eee a eee eee 28
Tsength‘of; carpus, OUtsid@. 222. -- sa-e ee ee en ee ena Se cee eae ae eee eee 35
Width of proximaliend'ofimetacarpals iil WandehVize 2325-2 et ee en nee eee ae aee 37
mengthiof metacarpal «Ll, sinner Sid@i--- -s=-— se eee = eee ne ee eee nee eee ee QL
Teast transverse width ofmetacarpals) TL and! DVi— 22222 22see se oo ee eee eee 37
Tength of the:three: phalanges of digit Wle- 22 = 2-8 = 2 ae ree ee eee eae 95
Posterior limb
Greatest length of: innominate bone: 2222226 = == seo aa ana oe a ee ae ee = en eee 270
Center of acetabulum to anterior border of ilium) 222-222. 22-222 222-2 ee eee 140
Centeriof, acetabulum) toiposterior border of ischiuim==-22- -2-- ose nae ee eae 135
‘Diameter ofacetabulum: 20-25-22 Sa. 822 oe ee es Be ee ee ee 35
Leng throtipubic: Sym py SiSsa- a2. oso ee ee ne a ee oe es 90
ongest.diameteriofobturatonforamenks 26 = ae e nn eee eeeeee 52
Greatestlenethroftam nr Ses rer a er ee oe ne ee ne 226
Anteroposterior diamoeteriof head of femursss = se ere ee a eee 34
‘Transverserdiameter of head Offemure= oes an an er Se eae eee ee ee eee 30
Transverse diameter of condyles of femtUbias.5 se cacao = eee a eae es ae en eee ee 53
Transverse:aiamecer or. trochles Ob lemuns=--2 oes ee eee ae ee eee 25
Teneth .ottibia, innemside.2.- 22. -+ 222 se ee a ee ee oe ee eee 205
Width ofiproximaliond! ofitibia 222-2 e ss a ene 64
Wiidthiot distalsend: of tibiais-<2 2 se he er ee eee ee 35
Width: of:proximaliend ofiastragalus==---2---- =---- 3 === se ene eee ee een enenene 26
Greatestlenathofiastrsg al iis sae re we a eee 42
Greatest length’ of calceanetm se 2a e a os aa as Se er eee 75
Motaltlengthiottarsussccce ako e saa ee a ee ee ee ee eee 62
Wiidth‘of;proximaliend’ ofimetatarsalsossessne sons a a ee ee eee 30
Width ofidistal:endot metatarsals: se ne ee arn ee ees ae ee eee 34
Length of metatarsals, inner sid@=- 2.) 23 sae sso cn See ea nee een nena 97
Least widthioftused. metatarsal soe ae a ee le re ee ee eee eee 23
Totalilensthrof three: phalanges of Gititii ll sense = eee nn enero nae 86

PLATYGONUS VETUS(?) Leidy

A single palate and maxillary portion, U.S.N.M. no. 8917, with the
permanent and two deciduous premolars and the first two molars,
was believed by Gidley (1920b, p. 658) to represent Platygonus vetus.
The molars in this specimen are somewhat larger than the average in
P. cumberlandensis and unusually wide, approaching the relative
proportions seen in P. vetus. Although this specimen may represent
an individual of P. cumberlandensis showing a variation of characters
in the direction of P. vetus, its reference to the latter species is tenta-
tively retained. It is possible that were Platygonus vetus better known
the variation of characters within that species would be sufficiently
great to include those outlined for P. cumberlandensis. However, in

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 83

view of observed dental differences it seems preferable to regard the
latter as a distinct form.
MYLOHYUS EXORTIVUS Gidley 1°
Ficures 46-48

The type of this species is from Cumberland Cave and consists of a
nearly complete lower jaw (fig. 46), U.S.N.M. no. 8876, containing the

pe specimen (U.S.N.M.

Maryland.

no. 8876), lateral view. Two-thirds natural size. Cumberland Cave Pleistocene,

FIGURE 46.—Mylohyus erortious Gidley: Right ramus of mandible and upper cheek teeth, ty

entire dentition of both rami (fig. 476) and five upper teeth, P? to M3,
from the right side (fig. 47a). The original description given by
Gidley is as follows:

About the size of or perhaps somewhat smaller than M. nasutus (Leidy), and

apparently differing from that species in (1) the modifications of the canines,
which seem to be relatively wider in cross-section and almost entirely lacking the

18 Gidley, 1920b, pp. 676-678.

84 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

longitudinal ribbing so characteristic of these teeth in M. nasutus; (2) the form
of the third upper premolar (the only tooth which can at present be directly com-
pared) which is proportionately broader, while the secondary cuspules are much
less prominent; and (3) the relative distance between the cheek-tooth rows of the
opposite sides, which is much greater than in the type of Leidy’s species.

A second specimen (Cat. 8160 U.S.N.M. Coll.), consisting of three milk molars
and the first true molar of the right side [fig. 48], seems to belong to this species,
although the first molar, which is entirely unworn, is much narrower than the
corresponding tooth in the fully adult type specimen. This difference, however,

FIGURE 47.—Mylohyus erortivus Gidley, type specimen (U.S.N.M. no. 8876): a, Superior cheek teeth;
6, inferior cheek teeth. Two-thirds natural size. Cumberland Cave Pleistocene, Maryland.

may be due to the fact that in the young specimen this tooth had not reached the
stage where the roots had begun to form and is therefore so immature as not to
have yet acquired its maximum width.

This specimen is especially interesting, since it affords a comparison of the milk
dentition with that of the type of M. pennsylvanicus. The unworn condition of
the deciduous teeth and the incomplete development of the first permanent molar
in this species indicate a somewhat younger individual than Leidy’s type. It
differs from the latter in (1) the smaller size of the corresponding teeth (see table
of measurements), (2) in the narrower proportions of the milk-molars, and (3)

FIGURE 48.—Mylohyus exortivus Gidley: Superior deciduous premolars and first permanent molar of right
side (U.S.N.M. no. 8160), occlusal view. Natural size. Cumberland Cave Pleistocene, Maryland.

in the greater complication of secondary cuspules, especially in the anterior half
of each of these teeth. In this last feature they show a decided advance in com-
plexity of the milk-teeth over those of either of the living genera of peccaries. This
is less pronounced in M. pennsylvanicus but this species also has more complex
milk-molars than either Tayassu or Pecari.

A comparison of the lower jaw of the type of M. exortivus with
illustrations of the Frankstown Cave lower jaw referred by Peterson
(1926, p. 256, pl. 18) to Mylohyus pennsylvanicus shows the two to be
very nearly the same size, but a noticeable difference is seen in the
character of P,. In the Frankstown Cave form P, is a simple tooth
with two cusps longitudinally arranged. In M. exortivus the tooth is
completely molariform though smaller and relatively narrower than
the succeeding teeth.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 85

Mylohyus browni Gidley (Gidley, 1920b, pp. 675-676; Brown, 1908,
pp. 200-202, pl. 24) is a more slender and elongate jawed form than
M. exortivus, although according to the illustrations given by Brown
P, in the Conard Fissure specimen is molariform, or nearly so.

A pair of fused metatarsals in the Cumberland Cave peccary
material is referred to Mylohyus. These bones are distinctly longer
and slenderer than in Platygonus.

TABLE 24.— Measurements (in millimeters) of mandible and dentition of Mylohyus

exortivus
S.N.M.
Measurement "to sib Ue nee
ype i
Mandible
HPepthiotjaws belowafirstymolanrs —- - oe ae ee ee ee. eS ee eee 700) | © een
Thickness ofsjawapelow Lirst, molar. s2e. ona 2 ao aoe an pa eee oaSeeeeeeee 20 eh | pec cto
Waidthiacross|both jawsibelow first molar:=-—2-+ 2 22-232 5 =< 5 = ona ZOhy || Seaeaeaueeee
Width of laws aticanines= se". <6 22 ee too sees ee cee 2 ee eae BO: Weesdecesses
Width between\canine’alveolit #222352. 22! 2.29.52 5 hoes 5 ee 2b. Se seca. UGiyy ees ewes os
Widthiotsymphysis atmarrowest point... =~. --- 225.225 -- = s-<s5.- ewes 24. p\escsecsascas
Werticalidepth of symphysis, median lin@l2-.25-222—--+-+----e sees s- cee e ee - 196 "|e22ht 2 ee
Tengethiof diastems in’front of'caniness< 2222. 222... 2.222 222222 4252. 35222220 5.-25 Din 2 Wee og. ee
Irene thy otdiastemai behind canines: o. 2-22 2225 2 koe acta eae ose ceeeeesees Wile See ae eae
Superior dentition
Ps anteroposterior GinmMelers= 26. sa-— ae san Soe eee wen ne ee aa an ene ee oe cce TON Sulbos nse
Ps. transverse GinmMeler.seases5_2 620 sete sess Shee oe SS eee ee 17: St ee Et es
PAvantero posterior Gismeter=-).s5--2 <a -oe sacs cece as ow ees cn sno satan sence TORS Hae wea ke Soe
Pé; transverse.diameters.. + (fees ee eee See ses 2 ea ee ee ee oe 1Sh8ule cece uc
MVMlanteropostenion Ginmetenn =. 2-222 2— aaa asa oes snk oaee---cosSeneeaee ae ee
IMUStransverse disinmerer. so 2 net a SSE Sea sonnet EEE SE 14. By | ee
Ma anteropostenloniGi ameter ~ sa oo5— sae tee ee eee on oe eee eee 5h | | See
IVES trANS VerserG laine L Gls see a on ee aren eee oa aoa eseaenesaee 15 eee a
Wis*antoropostenior dismeter..2225.4=. 226-4. a ee 16%) eae ee eee
IMiehtransverse Gismeter.-+ 2 2ts8-ascst ae ae ne ee ee ee Soi | aes ne ace
Inferior deniition
Oe antoropOseniOr Gi ailelelsa =a see saa. een ee ee aera eee eee MOT tees
Op tranis Verse GibINO LOL = 22 seen ose ea nae ane e eee aoe eee een eee Oy ie eee
Peyvantereposterionr diameters an. 22s. ee on ese eee een eee 0p |S
Pa ULavS Verse) GIAMeLer==. seo acee eee ae ee oa nee ee ee eee 634: |esScnt settee
Ps Anteroposterior Gia Meter. oo. 2s e ese n a= ae ae eee en eae ee eee bE en |
Py ULanSVerse GiaMelerans ae an one ae eee ne eee e na saa cane na aanecoes LOND 2 Ss a8 eae
Pa ianteroposterior diameter==2--.- 5--24 06 2-2 yt Sa aes eee es 12 oil ae ee eee
PP transverse Gis Meter. -cs--- oc oee as] b eon sao ene aoc anassecn ase eas ace ee 13) 6 less oeeaseee
Vir, anteroposterior dismMetersec sense ese se aot ee een tena a oe eee LBiir |e Se Se
MMi LrANS Verse diamMeLern sa ae ae ce oe a ee ee oe ee eens 13,4. je2Ae2cs5220
Wis Anitero posterior did Molter. aos.s5 325 2o es se eee mee ee en Sone ee he 16S eae eee
Me atTans Verse \diaMetel. ae 2a aoe aaa 2 oe ee se a eben ea eo eee 14, “ess .teseeee
Wis; anteroposterior GisMOtere sa ese eae see ene nee eee anne nen ee 10) 7; |Saneccsens25
Vig LranisVOrse GianOter seas safe sao 8 se seeeene ere ceo somone ae = =e 13.3 2s seece-nss
Immature superior dentition

Dp anteroposteriOniGgiameten. =~ eso) - ee ae ean nee eee eae nena ae | anes 10

Tass UPA TISVGESO IG 1AINO LOL se see ee ae a ee eee ee 7.5
ID pe anteroposterion,digt 6lOhs se ses seen aaa ne ocean ee eeae eee sneaen nae | eae 12.5
Dp strans verse diameters eas ses a ee eee eee ee ee ee eee eee 9

Dian teroposteriOniginmMelehacee sca. s ee ea Bn ee a ee eres | 12

IDs AECASVerSelG sO COR se sae ea ee ee nee eee heen o eee eee 11.5
Mil, anteroposterior diameter. -..-.--.---------.--_.-...---.--~----------=-=--.---=|------.----- 14

IMI trenis vorse Gis tOlLers sone ence ee ee eee ane ean oeners paeaeeaeeee 11.5

86 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

MYLOHYUS cf. PENNSYLVANICUS (Leidy) *

A jaw fragment, U.S.N.M. no. 8162, with the last two molars was
referred by Gidley (1920b, p. 674) to M. pennsylvanicus. ‘These
two teeth are much longer and M? is actually narrower and M? rela-
tively narrower than in M. ezortivus. A comparison of the first
permanent molars in the types of M. exortivus and M. pennsylvanicus
shows the latter to have a longer and relatively narrower M', and by
inference the second molars may have been similarly proportioned.

The Frankstown Cave form (Peterson, 1926, pl. 18) apparently
also has relatively narrow molars as indicated by the illustrations.

The various species of Mylohyus recognized are known only on
rather fragmentary material, and the amount of individual variation
that exists in any of these species is not yet determinable.

Family CERVIDAE

CERVUS species

An immature pair of fused metatarsals, apparently belonging to
an elk, shows the presence of a moderately large cervid in the fauna.
The cannon bone, though immature, is much too large to represent
Odocoileus or Eucervus. There is at present no way of knowing
whether the form Sangamona is represented here, inasmuch as the only
described species, S. fugitiva Hay, has for a type an isolated tooth.

GDOCOILEUS cf. VIRGINIANUS (Boddaert)

Several upper and lower jaw fragments of deer are included in the
collection. No characters of diagnostic importance were observed
to distinguish the cave form from the living white-tailed deer,
Odocoileus virginianus. A comparison of the fossil dentitions with
those of the black-tailed deer, O. columbianus, shows a somewhat
narrower inner wall in the upper premolars of the Pleistocene form.
This anteroposterior compression of the inner lobe of the upper pre-
molars is noticeable in the subgenus Odocoileus.

Family BOVIDAE
EUCERATHERIUM(?) AMERICANUM (Gidley)
Ficures 49, 50

An upper dentition (fig. 49) U.S.N.M. no. 7622, of a bovid type was
described by Gidley (19132) as representing a new eland, Taurotragus
americanus. The living eland is African in distribution and belongs
to a group of antilopine forms foreign to the New World. Hence,
the recognition of Taurotragus in a North American cave deposit
merits a critical reexamination of the fossil evidence.

Comparison of the Cumberland Cave dentition with that of the
bovid types Luceratherium and Preptoceras was made by the junior

30 Leidy, 1889, pp. 8-12, pl. 2, figs. 3-6.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 87

author (Gazin, 1933). This comparison was greatly aided by addi-
tional material secured by Gidley subsequent to his report, which
included a lower jaw (fig. 50a), U.S.N.M. no. 7993, with teeth and
lower premolars (fig. 506), U.S.N.M. no. 8006, belonging to another
individual.

In a general way certain outstanding characters seen in the fossil
dentition, as large size, hypsodonty, and proportionally large pre-

SN
SS

FIGURE 49.—Euceratherium(?) americanum (Gidley): Superior cheek teeth, P3-M3, type specimen
(U.S.N.M. no. 7622), lateral and occlusal views. Three-fourths natural size. Cumberland Cave Pleisto-
cene, Maryland.

molars, are strongly suggestive of the eland and likewise contrast it
with the larger living bovids of North America. A more detailed
examination, however, with particular regard for the structure of the
premolars, warrants removing the fossil form from Yaurotragus.
The differences seen in the upper molars are not many and not of
great importance when we consider the variability of the pattern of
these teeth in bovid forms. The external styles are better developed
in the fossil, as noted by Dr. O. P. Hay (1920), and these teeth are of
somewhat greater size. The upper premolars are particularly robust,
are apparently more hypsodont, and have a less compressed postero-
external style.

In the lower molars M, is noticeably wide transversely and M;
relatively narrow as compared with the modern eland; differences in
the lower premolars appear to be more significant, however. The
premolars are distinctly more hypsodont. The crown of P, is more
complexly folded, resembling in development a P; in Ovibos. The

88 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

second premolar in the eland though large is markedly simple. The
fossil P; is particularly distinctive. The anterior portion of this tooth
is broader than in the eland, and the prominent lingual fold is deeply
pocketed. The extension of the inner wall encloses the fold for a

FIGURE 50.—Euceratherium(?) americanum (Gidley): a, Left ramus of mandible (U.S.N.M. no. 7993), lateral and occlusal views;
b, right lower premolars (U.S.N.M. no. 8006), occlusal view. One-half natural size. Cumberland Cave Pleistocene, Maryland

greater portion of the crown height. The prominent lingual fold in the
third premolar of the eland is open in all specimens that were examined.
The fourth premolar of the fossil jaw possesses a talonid column hav-
ing a somewhat greater anteroposterior development; apparently this
tooth is more nearly molariform.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 89

TaBLE 25.—Measurements (in millimeters) of superior and inferior dentition of
Euceratherium(?) americanum

US.N.M.| og nM.

Measurement no. 7622 no. 7993

(type)
Superior dentition
Length of upper tooth series, P3 to M3 inclusive--------------------------------- 140515} 2ee Se
Length of upper molar series, M! to M$ inclusive-------------------------------- TOU.) S58 oe Peace
P3, greatest anteroposterior diameter-.------------------------------------------- TOMA KL eee
P3, greatest transverse diameter at occlusal surface !__---------------------------- TSN sles nee eee et ee
P4, greatest anteroposterior diameter -------------------------------------------- 99°15 ee SEE
P4, greatest transverse diameter at occlusal] surface--_---------------------------- 18} Wests ee
M1}, greatest anteroposterior diameter-------------------------------------------- 30)0) jpscesee eee
M!, greatest transverse diameter at occlusal surface------------------------------ QE 7 (eas se
M2, greatest anteroposterior diameter-------------------------------------------- S45 Gul bwseeeeoe ee
M:?, greatest transverse diameter at occlusal surface_----------------------------- 22,2 Wace SNS ue ue
M3, greatest anteroposterior diameter-_------------------------------------------- BT Ti ane
M3, greatest transverse diameter at occlusal surface------------------------------ 20 Gee eee
Inferior dentition
Length of lower tooth series, P2 to P, inclusive-_---------------------------------|------------ 157
Length of lower premolar series, P2 to Ps inclusive_.-----------------------------|------------ 269
Length of lower molar series, M: to M3 inclusive_---_----------------------------|------------ 100
Po, greatest anteroposterior diameter----_----------------------------------------|------------ 3715
Ps, greatest transverse diameter at occlusal surface--_----------------------------|------------ 210.6
P3, greatest anteroposterior diameter----_----------------------------------------|------------ 19.2
Ps, greatest transverse diameter at occlusal surface-------------------------------|------------ 13
Pu, greatest anteroposterior diameter-__-------------------------------------------|------------ 25
Pu, greatest transverse diameter at occlusal surface--_----------------------------|------------ 13. 5.
Mi, greatest anteroposterior diameter--------------------------------------------|------------ 26
Mu, greatest transverse diameter at occlusal surface------------------------------|------------ 17.8.
Mz, greatest anteroposterior diameter--------------------------------------------|------------ 33. 8
Mz, greatest transverse diameter at occlusal surface------------------------------|------------ 16.1
Ms, greatest anteroposterior diameter--------------------------------------------|------------ 41.7
Ms, greatest transverse diameter at occlusal surface------------------------------|------------ 13.8

1 Width of occlusal surface of teeth increases with wear.
2 Approximate,

The dentition exhibited by specimens of Huceratherium collinum
Sinclair and Furlong (1904) and Preptoceras sinclairi Furlong (1905)?!
though of a bovid type is of a kind clearly distinguished from that
seen in the Bovinae, or even in Ovibos although to a less extent. The
hypsodonty reached in the specimens from the Pleistocene of Cali-
fornia is suggestive of an antilopine dentition, but the skull and horn
structures they possess show them to belong to a distinct group. The
development of the premolars in this western group has gone beyond
that in the eland. It is further noted that those dental characters
that appear most significant in distinguishing the Cumberland Cave
fossil from the African eland ally it to the Huceratherium-Preptoceras
group. Also, the marked increase in width and the shortening
anteroposteriorly of the upper molars in advanced wear noted by
Gidley (1913a, p. 1) for the Cumberland Cave type was observed by
Sinclair and Furlong (1904, p. 416) in Huceratherium collinum. The

21 See also Stock and Furlong (1927).

90 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

only outstanding difference noted is the greater size of the Maryland
type as compared with the California forms. In addition, the pre-
molars in the lower jaw, Univ. Calif. Pal. Coll. no. 8756, from Potter
Creek Cave, which the junior author examined, appear to be slightly
more hypsodont, and the isolation of the lingual fold in P; is more
complete.

In view of the disparity in size, it is possible that were more com-
plete material known exhibiting the skull and horn structure Hucera-
therium(?) americanum would be found to represent a distinct genus.
On the dental structure alone, however, no important differences can
be cited that would serve to distinguish this form generically from
Euceratherwum or Preptoceras.

Our form is referred tentatively to Huceratherium, as apparently
the differences that separate Preptoceras from Hucerathervum may not
be greater than could be accounted for by individual variation. No
important dental differences were seen between the two, although the
teeth in the type of Preptoceras are slightly larger than in Eucera-
thervum.

LITERATURE CITED

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1899. Vertebrate remains from Port Kennedy bone deposit. Journ. Acad.
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FENNEMAN, NEVIN MELANCTHON.
1930. Physiographic divisions of the United States. U.S. Geol. Surv. map,
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Furione, Eustace LEOpPoLp.
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Gazin, Cuartes Lewis.
1933. The status of the extinct American “eland.” Journ. Mamm., vol. 14,
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1934. Upper Pliocene mustelids from the Snake River Basin of Idaho.
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GIDLEY, JAMES WILLIAMS, and Gazin, CHARLES LEWIS.

1933. New Mammalia in the Pleistocene fauna from Cumberland Cave.

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1922. Descriptions of species of Pleistocene Vertebrata, types or specimens of
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1923. The Pleistocene of North America and its vertebrated animals from the
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Ho.uuister, Nrep.

1911. A systemstic synopsis of the muskrats. North Amer. Fauna, no. 32,

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1927. Revision of the American lemming mice (genus Synaptomys). North

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1853. Description of an extinct species of American lion, Felis atroz. Trans.
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1860. Description of fossil vertebrates. In F. S. Holmes’s ‘‘ Post-Pleiocene
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1873. Contributions to the extinct vertebrate fauna of the western territories.
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1889a. On Platygonus, an extinct genus allied to the peccaries. Trans.

Wagener Free Inst. Sci., vol. 2, pp. 41-50, 1 pl.

1889b. Notice and description of fossils in caves and crevices of the limestone
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Merriam, Ciintron Hart.

1900. The life zones and areas of Allegany County. Maryland Geol. Surv.

Allegany County, pp. 291-293.
MerriaM, JOHN CAMPBELL.

1918. New puma-like cat from Rancho La Brea. Univ. California Dept.

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Merriam, JoHN CAMPBELL, and Stock, CHESTER.

1925. Relationships and structure of the short-faced bear, Arctotherium, from
the Pleistocene of California. Carnegie Inst. Washington Publ. 347,
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1932. The Felidae of Rancho La Brea. Carnegie Inst. Washington Publ.
422, 231 pp., 152 figs., 42 pls.

Mi.urrR, GERRIT Situ, Jr.
1896. The genera and subgenera of voles and lemmings. North Amer.
Fauna, no. 12, 84 pp., 40 figs., 3 pls.

PLEISTOCENE VERTEBRATES FROM CUMBERLAND CAVE 98

1899. A new fossil bear from Ohio. Proc. Biol. Soc. Washington, vol. 13,
pp. 53-56.
1912. The names of the large wolves of northern and western North America.
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PATTERSON, BRYAN.
1932. Upper molars of Canis armbrusteri Gidley from Cumberland Cave,
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1894. A contribution to the life history of the Allegheny cave rat, Neotoma
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1928. Pleistocene mammals from a cave in Citrus County, Florida. Amer.
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Srock, Cuester, and Furtone, Eustace Lroronp.
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Stosz, Greorce WILxIs, and Miser, Hucn Dinsmore.
1922. Manganese deposits of western Virginia. Virginia Geol. Surv. Bull.
23, 206 pp., 39 figs., 31 pls.
Stose, Grorce WILLIS, and Swartz, CHARLES KEPHART.
1912. Description of the Pawpaw and Hancock quadrangles. U. 8S. Geol.
Surv. Geol. Atlas, Pawpaw-Hancock folio, no. 179, pp. 19-21.
WETMORE, ALEXANDER.
1927. A record of the ruffed grouse from the Pleistocene of Maryland. Auk,
vol. 44, p. 561.
WiLuisTon, SAMUEL WENDELL.
1894. Restoration of Platygonus. Kansas Univ. Quart., vol. 3, pp. 23-39,
6 figs., 2 pls.

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INDEX

(Principal references are given in boldface)

Abbe, Cleveland, Jr., 3, 4.
Aenocyon, 17.
dirus, 8, 17.
Alden, W. C., 1.
alleganiensis, Corynorhinus, 5, 8, 12 seq.
Allegany Mountain, 4
Allegheny County, 2.
alleni, Hodomys, 61, 62.
Neofiber, 65.
Alligators, 10.
ambarvalis, Spilogale, 43, 44.
americana, Martes, 32, 33.
americanum, Euceratherium, 5, 9, 8&6

seq.
Mammut, 5, 8, 70.
americanus, Euarctos, 8, 24, 25, 26, 27.
Lepus, 5, 8, 69.
Taurotragus, 86.
Ursus, 26.
amplidens, Ursus, 26.
Anaptogonia, 8.
hiatidens, 8.
angulatus, Pecari, 79.
angustidens, Mustela cicognanii, 8.
Anita, Ariz., material, 49.
annectens, Ondatra, 5, 8, 65 seq.
Appalachian Plateau, 3. 4
Appalachian region, 69.
aquaticus, Scalopus, 8.
Arctodus, 5, 8, 9, 27 seq.
floridanus, 32.
haplodon, 5, 8, 27 seq.
pristinus, 32.
Arctotherium, 32.
haplodon, 32.
Armbruster, Raymond, 2,
armbrusteri, Canis, 5, 8. 15 seq.
Artiodactyla, 5, 71 seq.
atrox, Felis, 5, 8, 51 seq.
australis, Symbos, 9.
Badgers, 6, 7, 48, 49.
Bats, 6, 11.
IBGnEs oe ti 2a peas aay 20; 2s hg al es
Beavers, 7
Bison, 7.
Bison sp., 9.
Blarina, 5, 8, 16 seq.
brevicauda, 5, 8, 10 seq.
brevicauda ozarkensis, 8, 10.
simplicidens, 8, 11.
Blue Ridge, 3.
Bog-lemmings, 7.
Bonasa, 5, 10.
umbellus, 5, 10.
borealis, Synaptomys (Mictomys), 63.
Bovidae, 5, 86 seq.
Bovinae, 89.
Brachyprotoma, 5, 8, 43 seq., 46.
obtusata, 8, 45.
pristina, 5, 8, 43 seq., 46.
spelaea, 44, 45.

brevicauda, Blarina, 5, 8, 10 seq.

brevicauda ozarkensis, Blarina, 8, 10.

Brown, Barnum, 3, 7, 10, 11, 26, 43, 44,
50, 56, 65, 66, 85.

browni, Mylohyus, 9, 85.

calcaratus, Lynx, 8.

californicum, Tremarctotherium, 32.

ealycinus, Sciurus, 8.

Camels, 7.

Camp, C. L., 2.

canadensis, Castor, 5, 8, 59.
Cervus, 9
Lutra, 41.

Canidae, 5, 15 seq.

Canis, 5, 8, 15 seq.
armbrusteri, 5, 8, 15 seq.
dirus, 8, 17.
latrans, 23.
occidentalis, 8, 15, 16.
priscolatrans, 5, 8, 23.
riviveronis, 23.
rufus (=frustror), 23.

Carlisle Cave, Pa., 60.

Carnivora, 5, 15 seq.

Castor, 5, 8, 59.
canadensis, 5, 8, 59.

Castoridae, 5, 59.

Cats, 6, 7, 49, 51.

caurina, Martes, 33.

Cavetown, Md., 54.

Cervidae, 5, 86.

Cervus, 5, 9, 86.
canadensis, 9.

Chiroptera, 5, 11 seq.

cicognanii angustidens, Mustela, 8.

cinereoargenteus, Urocyon, 8.

cinereus, Sorex, 10.

Citellus, 5, 8, 54.
tridecemlineatus, 5, 8, 54.

Clark, W. B., 3, 4.

cloacinum, Sycium, 8.

Coastal Plain, 3.

collinum, Euceratherium, 89.

columbianus, Odocoileus, 86.

complicatus, Equus, 9, 70.

compressus, Lynx, 8.

Platygonus, 72.
Conard Fissure, 3, 7, 8-11, 26, 43, 45,
53, 56, 65, 66, 85.

conardi, Smilodontopsis, 8.

Conepatus, 44.

cooperi, Synaptomys, 5, 8, 62 seq.

Cope, E. D., 7, 27, 32, 35, 41, 42, 45,

49, 50, 53, 65, 69.

Corriganville, 2.

Corynorhinus, 5, 8, 12 seq.
alleganiensis, 5, 8, 12 seq.
macrotis, 12, 13, 14.
phylotis, 13.
rafinesquii, 12, 13, 14.
rafinesquii pallescens, 14.

96 BULLETIN 171, UNITED STATES NATIONAL MUSEUM

couguar, Felis, 8.

Coyotes, 6, 7, 15, 17, 23.

Cricetidae, a 59 seq.

Crocodylid, 5, 6, 7, 10. ne
Cumberland, Md., Daa: a
cumberlandensis, BE aie 5; ae 71

seq.

daggetti, Felis, 51.

Deer, 86.

Devonian limestone, 2.

dideltus, Microtus or Pitymys, 8, 64,
65

diluviana, Martes, 8, 34.

diluvianus, Microtus, 8.

dirus, Canis (Aenocyon), 8, 17.

dorsatum, Erethizon, 5, 8, 67 seq.

Eland, 2, 86, 87, 88, 89

Elephants, 7.

Elk, 6, 86.

emmonsii, Euarctos, 24.

epixanthum, Erethizon, 67.

Eptesicus, 5, 8, 11 seq.
fuscus, 11, 12.
grandis, 5, 8, 11 seq.

Equidae, 5, 70.

Equus, 5, 9, 70.
complicatus, 9, 70.
pectinatus, 9, 70.
scotti, 9.

Erethizon, 5, 8, 67 seq.
dorsatum, 5, 8, 67 seq.
epixe inthum, 67.

Erethizontidae, 5, 67 seq.

Euarctos, 5, 8, 23 seq
americanus, 8, 24, 25, 26, 27.
emmonsii, 24.
luteolus, 24, 25, 26, 27.
vitabilis, 5, 8, 23 seq.

Euceratherium, 5, 9, 86 seq.
americanum, 5, 9, 86 seq.
collinum, 89.

Eucervus, 86.

exortivus, Mylohyus,"5, 9, 83’seq., 86.

eyra, Felis, 8.

Felidae, 5, 49 seq.

felina, Lutra, 41.

Felis, 5, 8, 49 seq.
atrox, 5, 8, 51 seq.
couguar (concolor), 8.
daggetti, 51.

eyra, 8.

inexpectata, 5, 8, 49 seq.
leo, 51, 52.

longicrus, 8, 50, 53.
onea, 50.

tigris, 51, 52, 53.
Fenneman, N. M., 3.
Fiber, 65.
Fishers, 6, 7, 34.
Florida, 1, 238, 44, 59, 70.
floridana, Neotoma, 60.
floridanus, Arctodus, 32.
Sylvilagus, 8.
Flying squirrels, 56.
fossidens, Sorex personatus, 8.
fossor, Thomomys, 59.
Fox, 15.
Frankstown Cave, 32, 45, 84, 86.

fremonti, Sciurus, 54, 55, 56.

frustror, Canis, 23.

fugitiva, Sangamona, 86.

fulva, Vulpes, 8.

fulvus, Thomomys, 57.

fumeus, Sorex, 8.

Furlong, E. L., 89.

fuscus, Eptesicus, 11, 12.

fuscus grandis, Eptesicus, 11.

Galliformes, 5, 10.

Gazin, C. L., 2, 87, 90.

Geomyidae, 5, 57 seq.

Geomys, 6, 8, 57, 58, 59.
parvidens, 8.

Gidley, J. W., 2, 3, 6,7, 18, 15, 17, 24,

72, 79, 82, 83, 86, 87, 89.
gidleyi, Gulo, 5, 8, 38 seq.
giganteus, Lepus, 8.

Gilmore, C. W., 1, 10.
Glaucomys, 5, 8, 56 seq.
sabinus, 56.
volans, 56.
yukonensis, 56.
Goldman, E. A., 1.
gracilis, Mustela, 8.
Smilodontopsis, 8.
Spilogale, 44.
grandis, Eptesicus, 5, 8, 11 seq.
Granger, Walter, 2
Grizzly bear, 25.
Grouse, 2, 10.
Gulo, 5, 6, 8, 34, 38 seq.
gidleyi, 5, 8, 38 seq.
luscus, 39, 41.
Hall, E. R., 35, 37, 39, 41, 42, 48, 49.
haplodon, Arctodus, 5, 8, 27 seq.
Arctotherium, 32.
Hares, 6, 7, 69.
harlani, Mylodon, 9.
Harrisburg surface, 4.
Hay, O. P., 4, 8, 9, 49, 54, 79, 87.
haysii, Tapirus, ’9, "70.
Helderberg formation, 3.
hemionus, Odocoileus, 9.
hiatidens, Anaptogonia, 8.
Hodomys, 60, 61, 62.
alleni, 61, 62.
Hollister, Ned, 65.
horribilis, Ursus, 25.
Horses, 7, 70.
Howell, A. H., 1.
hudsonicus, Sciurus, 8, 54, 55, 56.
hudsonius, Zapus, 8.
inexpectata, Felis, 5, 8, 49 seq.
ingens, Mustela vison, 37.
inornatus, Tremarctos, 32.
Insectivora, 5, 10 seq.
insignis, Napaeozapus, 5, 8, 67.
ened ie Platygonus, 7 it 72, 76, 77,
8

interrupta, Spilogale, 8, 46.

involutus, Microtus or Pitymys, 5, 8
63 seq.

Jackals, 15.

Jackson, H.)H. T.,.1,°10.

Jaguars, 50, 51.

Jennings Run, 2.

Jumping mice, 6, 67.

Kellogg, Remington, 1.
Keyser member, 3
Knobly Mountain, 4.
laevicornis, Odocoileus, 9.
Lagomorpha, 5, 68 seq.
lataxina, Lutra canadensis, 42.
latidentatus, Vulpes, 8.
latrans, Canis, 23.
Leidy, Joseph, 32, 72, 79, 84.
Lemmings, 6, 7, 63
leo, Felis, 51, 52.
Leporidae, 5, 69.
leptorhinus, Platygonus, 72.
Lepus, 5, 8, 69.
americanus, 5, 8, 69.
giganteus, 8
leucopus, Peromyscus, 5, 8, 59.
Lions, 51.
lobulatus, Pelycictis, 8.
longicrus, Felis, 8, 50, 53.
Loricata, 5, 10.
lotor, Procyon, 8.
loxodon, Megalonyx, 9.
luscus, Gulo, 39, 41.
luteolus, Euarctos, 24, 25, 26, 27.
Lutra, 5, 8, 41 seq.
canadensis, 41.
canadensis lataxina, 42.
felina, 41.
paraensis, 41.
parvicuspis, 5, 8, 41 seq.
rhoadsii, 8, 42.
Lyciscus, 17.
Lynx, 8.
calcaratus, 8.
compressus, 8.
rufus, 8
macrotis, Corynorhinus, 12, 13, 14.
magister, Neotoma, 5, 8, 59 seq.
Mammut, 5, 8, 70.
americanum, 5, 8, 70.
maniculatus, Peromyscus, 59.
Marmota, 5, 8, 53.
monax, 5, 8, 53.
Martens, 34.
Martes, 5, 32 seq.
americana, 32, 33.
caurina, 33.
diluviana, 8, 34.
parapennanti, 5, 8, 32 seq.
pennanti, 8, 32, 33.

marylandensis, Spilogale, 5, 8, 46 seq.

marylandica, Taxidea, 5, 8, 47 seq.
Mastodon, 9, 70.
Mastodontidae, 5, 70.
Mathews, E. B., 3, 4.
Megalonyx, 9.

loxodon, 9.

scalper, 9.

tortulus, 9.

wheatleyi, 9.
Mephitis, 8.

nigra, 8.

mesomelas newtonensis, 8.
mercerii, Smilodontopsis, 8.
Merriam, J. C., 32, 52.

INDEX O7

mesomelas newtonensis, Mephitis, 8.

Microsorex, 8.
minutus, 8.

Mierotus, 5, 8, 63 seq., 65.
dideltus, 8, 64, 65.
diluvianus, 8.
involutus, 5, 8, 63 seq.
ochrogaster, 8, 65.
pennsylvanicus, 64.
speothen, 8, 65.

Mictomys, 5, 6, 8, 63.
borealis, 63.

Miller, G. S., Jr., 1, 15, 65, 67.

Mink, 6, 7, 37.

mink, Mustela vison, 35, 37.

minutus, Microsorex, 8.

Miser, H. D., 3.

monax, Marmota, 5, 8, 53.

Musk-ox, 7.

Muskrat, 6, 7.

Mustela, 5, 8, 35 seq.
cicognanii angustidens, 8.
gracilis, 8.
nigripes, 35.
vison, 5, 8, 35 seq.
vison ingens, 37.
vison mink, 35, 37.
vison vison, 35, 37.

Mustelidae, 5, 32 seq.

Mylodon, 9.
harlani, 9.

Mylohyus, 5, 9, 83 seq.
browni, 9, 85.
exortivus, 5, 9, 83 seq., 86.
nasutus, 9, 83, 84.
pennsylvanicus, 5, 9, 84,86.

Myotis, 8, 13.
subulatus, 8.

Napaeozapus, 5, 8, 67.
insignis, 5, 8, 67.

masutus, Mylohyus, 9, 83, 84.
Tamias, 8.

Neofiber, 65, 66, 67.
alleni, 65.

Neotoma, 5, 8, 59 seq., 61, 62.
fioridana, 60.
magister, 5, 8, 59 seq.
ozarkensis, 8, 60.
pennsylvanica, 60.

newtonensis, Mephitis mesomelas, 8.

nigra, Mephitis, 8.

nigripes, Mustela, 35.

obseurus, Sorex, 8.

obtusata, Brachyprotoma, 8, 45.

occidentalis, Canis, 8, 15, 16.

Ochotona, 5, 8, 68 seq.
palatina, 8.
princeps, 69.
schisticeps, 69.

Ochotonidae, 5, 68 seq.

ochrogaster, Microtus (Pedomys), 8, 65.

Odocoileus, 5, 9, 86.
columbianus, 86.
hemionus, 9.
laevicornis, 9.
virginianus, 5, 9, 86.

onea, Felis, 50.

Ondatra, 5, 8, 65 seq.
annectens, 5, 8, 65 seq.
zibethica, 65, 66

Ophidian, 5, 10.

orientalis, Plesiothomomys, 58, 59.
Teleopternus, 9.
Thomomys, 59.

Oriskany formation, 3.

Osmotherium, 8.
spelaeum, 8.

Otters, 7, 41.

Ovibos, 87, 89.

ozarkensis, Blarina brevicauda, 8, 10.
Neotoma, 8, 60.

palatina, Ochotona, 8.

palatinum, Praotherium, 69.

BULLETIN 171, UNITED

pallescens, Corynorhinus rafinesquii, 14.

paraensis, Lutra, 41.
Parahodomys, 5, 8, 60 seq.
spelaeus, 5, 8, 69 seq.
parapennanti, Martes, 5, 8, 32 seq.
parvicuspis, Lutra, 5, 8, 41 seq.
parvidens, Geomys, 8.
Patterson, Bryan, 16, 17.
Pecari, 79, 84.
angulatus, 79.
Peccaries, 2, 6, 71, 72, 85.
pectinatus, Equus, 9, 70.
Pedomys, 65.
ochrogaster, 8, 65.
Pelycictis, 8.
lobulatus, 8.
pennanti, Martes, 8, 32, 33.
pennsylvanica, Neotoma, 60.
pennsylvanicus, Microtus, 64.
Mylohyus, 5, 9, 84, 86.
Perissodactyla, 5, 70 seq.
Peromyscus, 5, 8, 59.
leucopus, 5, 8, 59.
maniculatus, 59.
pena) Sorex, 8.
eterson, O. A., 32, 45, 84.
phenax, Spilogale, 44,
phyllotis, Corynorhinus, 13.
Piedmont Plateau, 3.
Pikas, 6, 69.
Pine-mice, 63.
pinetorum, Pitymys, 638, 65.
iscinaria, Satherium, 42.
itymys, 5, 8, 63 seq.
dideltus, 8, 65.
involutus, 5, 8, 63 seq.
pinetorum, 63, 65.
Platygonus, 5, 9, 71 seq., 85.
compressus, 72.
cumberlandensis, 5, 9, 71 seq.
intermedius, 71, 72, 76, 77, 78.
leptorhinus, 72.
tetragonus, 9.
vetus, 5, 9, 72, 82 seq.
Plesiothomomys, 5, 8, 57 seq.
orientalis, 58, 59.
potomacensis, 5, 8, 57 seq.
Pocket gophers, 6, 57, 59
Porcupines, 6, 67.

STATES

NATIONAL MUSEUM

Port Kennedy, 7, 8, 9, 23, 27, 32, 34;
41, 42, 45, 49, 50, 53, 69, "70.

potomacensis, Plesiothomomy 3). 10%) 105
57 seq.

Potter Creek Cave, 32, 90.

Praotherium palatinum, 69.

Prentice, Sydney, 2

Preptoceras, 86, 89, 90.

sinclairi, 89.

prineeps, Ochotona, 69.

priscolatrans, Canis, 5, 8, 23.

DESUne Brachyprotoma, 5, 8, 43 seq.,
4

pristinus, Arctodus, 32.
Proboscidea, 5, 70.
procerus, Ursus, 25.
Procyon, 8,
lotor, 8.
pumas, 6, 50, 51, 53.
putorius, Spilogale, 44, 46, 47.
rafinesquii, Corynorhinus, 12, 13, 14.
rafinesquii pallescens, Corynorhinus, 14,
Rancho La Brea, 16, 17, 32, 51, 52, 53;
Reeside, J. B., Jr., ie 2.
Reithrodontomys, 8.
simplicidens, 8.
Rhoads, S. N., 60.
rhoadsii, Lutra, 8, 42.
riviveronis, Canis, 23.
robusta, Taxidea, 49.
Rodentia, 5, 53 sed.
Roeder, George, 2 2.
rufus, Canis, 23.
Lynx, 8.
Rusconi, Carlos, 79.
Saber-tooth Cave, Fla., 59.
Saber-toothed cats, 7, 9, 53.
sabinus, Glaucomys, 56.
Sangamona, 86.
fugitiva, 86.
Satherium, 42.
piscinaria, 42.
Scalopus, 8.
aquaticus, 8.
scalper, Megalonyx, 9.
schisticeps, Ochotona, 69.
Schooley Plain, 4.
Sciuridae, 5, 53 seq.
Sciurus, 5, 8, 54 seq.
calycinus, 8.
fremonti, 54, 55, 56.
hudsonicus, 8, 54, 55, 56.
tenuidens, 5, 8, 54 seq.
scotti, Equus, 9.
Serpentes, 5, 10.
Shenandoah Plain, 4.
Short-tailed shrews, 10.
Shrews, 6, 10, 11.
Shriver Ridge, 4
simplicidens, Blarina, 8, 11.
Reithrodontomys, 8.
simum, Tremarctotherium, 32.
Sinclair, W. J., 89.
sinclairi, Preptoceras, 89.
Skunks, 43.
Sloths, 7, 9.

Smilodontopsis, 8.
conardi, 8
gracilis, 8.
mercerii, 8.
troglodytes, 8.
Snakes, 10.
Sorex, 5, 8, 10.
cinereus, 10.
fumeus, 8.
obscurus, 8.
personatus, 8.
personatus fossidens, 8.
Soricidae, 5, 10 seq.
spelaea, Brachyprotoma, 44, 45.
spelaeum, Osmotherium, 8.

spelaeus, Parahodomys, 5, 8, 60 seq.

speothen, Microtus, 8, 65

Spermophiles, 6.

Spilogale, 5, 8, 48, 44, 46 seq.
ambarvalis, 43, 44.
gracilis, 44.
interrupta, 8, 46.
marylandensis, 5, 8, 46 seq.
phenax, 44.
putorius, 44, 46, 47.

Squirrels, 6, 56.

Stirton, R. A., 2.

Stock, Chester, 32, 53.

Stose, G. W., 3.

striatus, Tamias, 5, 8, 54.

subulatus, Myotis, 8.

sulcata, Taxidea, 49.

Swartz, C. K., 3.

Sycium, 8.
cloacinum, 8. .

Sylvilagus, 8, 69.
floridanus, 8.

Symbos, 9.
australis, 9.

Synaptomys, 5, 8, 62 seq.
borealis, 63.
cooperi, 5, 8, 62 seq.

Tamias, 5, 8, 54.
nasutus, 8.
striatus, 5, 8, 54.

Tamiasciurus, 54 seq.

Tapiridae, 5.

Tapirs, 6, 9, 70.

Tapirus, 5, 9, 70 seq.
haysii, 9, 70.
veroensis, 70.

Taurotragus, 86, 87.
americanus, 86.

Taxidea, 5, 8, 47 seq.
marylandica, 5, 8, 47 seq.
robusta, 49.
sulecata, 49.
taxus, 8, 47, 49.
taxus taxus, 48.

taxus, Taxidea, 8, 47, 48.
Taxidea taxus, 49.

Tayassu, 79, 84.

INDEX 99

Tayassuidae, 5, 71 seq.

Teleopternus, 9.
orientalis, 9.

tenuidens, Sciurus, 5, 8, 54 seq.

tetragonus, Platygonus, 9.

Tetraonidae, 5, 10.

Thomomys, 6, 57, 58, 59.
fossor, 59.
fulvus, 57.
orientalis, 59.

Thos, 17.

Tigers, 51, 52.

tigris, Felis, 51, 52, 53.

tortulus, Megalonyx, 9.

Tremarctos, 32.
inornatus, 32.

Tremarctotherium, 32.
californicum, 32.
simum, 32.

tridecemlineatus, Citellus, 5, 8, 54.

troglodytes, Smilodontopsis, 8.

umbellus, Bonasa, 5, 10.

Urocyon, 8.
cinereoargenteus, 8.

Ursidae, 5, 23 seq.

Ursus, 24, 25, 26, 27.
americanus, 59.
amplidens, 26.
horribilis, 25.
procerus, 25.

Valley and Ridge Province, 3.

veroensis, Tapirus, 70.

Vespertilionidae, 5, 11 seq.

vetus, Platygonus, 5, 9, 72, 82 seq.

virginianus, Odocoileus, 5, 9, 86.

vison, Mustela, 5, 8, 35 seq.
Mustela vison, 35, 37.

vison ingens, Mustela, 37.

vison mink, Mustela, 35, 37.

vitabilis, Euarctos, 5, 8, 23 seq.

volans, Glaucomys, 56.

Voles, 63.

Vulpes, 8.
fulva, 8.
latidentatus, 8.

Weber, Rudolf, 2.

Wetmore, Alexander, 2, 10.

wheatleyi, Megalonyx, 9.

Williston, S. W., 72, 79.

Wills Creek, 2, 4.

Wills Creek Valley, 3, 4.

Wills Mountain, 4.

Wills Mountain anticline, 3.

Wolverines, 6, 38, 41, 71.

Wolves, 2, 15, 16, 17.

Woodchucks, 54.

Woodrats, 59, 60.

yukonensis, Glaucomys, 56.

Zapodidae, 5, 67.

Zapus, 5, 8, 67.
hudsonius, 8.

zibethica, Ondatra, 65, 66.

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