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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

OCCASIONAL PAPERS

of the '^^^UBR^y^^^

MUSEUM OF NATURAL HI^ORY

The University of KansasJAN 1 4 W74
Lawrence, Kansas Harvard

U^l:^ ,

NUMBER 21, PAGES 1-54 DECEMBER 2r, 1973

NEW NORTH AMERICAN ERINACEINE
HEDGEHOGS (MAMMALIA: INSECTIVORA)

Thomas H. V. Rich^ and Donald L. Rasmussen^

Modern hedgehogs, members of the subfamily Erinaceinae, first
appeared in the fossil record of Asia and Europe approximately 35
million years ago during the Oligocene. Toda\' the subfamily con-
tinues to thrixe on those two continents as well as Africa, where
they are known to have occurred first 20 million years ago near
the beginning of the Miocene.

Although now extinct in North America, members of the sub-
family first appeared on that continent approximately 23 million
years ago near the beginning of the Miocene. At that time, three
erinaceine species, belonging to as many different genera, made their
appearance in the fossil record. Two of the species are closely
related to forms knowTi in Oligocene deposits of Asia and Europe
and hence, apparcntlx- immigiated into North America shortly after
their appearance there in the fossil record. No knowTi erinaceids in
the pre-Miocene fossil record of either the Eastern or Western
Hemisphere are closely related to the third species. Therefore the
question of whether this third species is a newly arrived immigrant
into North America or a descendant from an Oligocene endemic
form remains unresolved.

The Erinaceinae persisted in North America until the latter part
of the Miocene about 10 million years ago. During the interval of
their presence on that continent, only one additional erinaceine

^ Department of Geolog>', Columbia University; Department of \'ertebrate
Paleontology, the American Museum of Natural History. Present address: The
Museum, Texas Technological University, Lubbock, Texas 79409.

" Department of Geology and Museum of Natural History, The University
of Kansas, Lawrence, Kansas 66045.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

genus and species appeared after the initial, nearly simultaneous
appearance of the three forms that marked the beginning of the
episode. Otherwise, the group did not undergo any noticeable
change in North America other than the extinction of its members.
Previously, the North American history of the Erinaceinae had
been documented by the description of fewer than two dozen speci-
mens of which less than half were correctly recognized as erina-
ceines. In an attempt to further elucidate the history of the Erina-
ceinae, this report will describe a similar number of new North
American erinaceine specimens and review those previously men-
tioned ones which are referred to the same taxa.

ACKNOWLEDGMENTS

For the loan of specimens we are deeply indebted to Mr. Morris
F. Skinner and Dr. Malcolm C. McKenna, Department of Vertebrate
Paleontology, and Dr. Sydney Anderson, Department of Mammal-
ogy, American Museum of Natural History; Dr. Larry D. Martin,
Museum of Natural History, University of Kansas; Dr. Robert W.
Wilson, Museum of Geology, South Dakota School of Mines and
Technology; Dr. Peter Robinson, University of Colorado Museum;
Dr. Robert W. Fields, Department of Geology, University of Mon-
tana; the late Dr. Claude W. Hibbard, Museum of Paleontology,
University of Michigan; and Dr. Elwyn L. Simons, Peabody Mu-
seum of Natural History, Yale University. We also thank Drs. Craig
C. Black, Malcolm C. McKenna, and Larry D. Martin for criticizing
the manuscript and making several suggestions for improvement.

Field work in Montana by Rasmussen was supported by: a)
National Science Foundation Grant GP-1553 to Dr. Robert W.
Fields, principal investigator; b) Division of Vertebrate Paleontol-
ogy, Museum of Natural History, University of Kansas; and c)
Watkins Fund, Museum of Natural History, University of Kansas.

Ranchers Dean Tavenner and Marvin Radtke gi-aciously gave
permission to collect specimens on their properties.

We thank Mrs. Mary Lee Vickers and Mrs. Geraldine J. Ras-
mussen who typed this manuscript.

METHODS AND ABBREVIATIONS

The primary geochronologic units employed in this report are
the North American and European Land-Mammal Ages. Also given
are the approximate epoch and radiometric age equivalents based
on data summarized in Berggren ( 1971) and Van Couvering ( 1972).

Figure 1 illustrates the method employed for making measure-
ments of teeth discussed in this report, and figure 11 illustrates how
angular measurements of lower jaws were taken.

Museum abbreviations used with catalogue numbers are as
follows :

NORTH AMERICAN ERINACEINE HEDGEHOGS 3

AMNH Department of Vertebrate Paleontology, American Museimn

of Natural History

AMNH (M) Department of Mammalogy, American Museum of Natural
History

F:AM Prick American Mammals, Department of Vertebrate Pa-

leontology, American Museum of Natural History

KU Museum of Natural History, University of Kansas

MPUM Department of Geology, University of Montana

SDSM South Dakota School of Mines and Technology

UCM University of Colorado Museum

UMMP Museum of Paleontolog>', University of Michigan

YPM Yale Peabody Museum

HISTORICAL RESUME

Established by Koerner in 1940, Parvericius montanus was the
first erinaceine species to be recognized and named in North Amer-
ica. The new species was based on a single specimen, consisting
of a maxillary fragment with hV -^ and the buccal regions of P^"^,
from the Miocene Deep River Formation of Meagher County, Mon-
tana. Koerner regarded the species as an erinaceine and considered
it to be more closely related to Erinaceus than to any other member
of that subfamily, devoting the generic diagnosis to an enumeration

,l.p2

l,-P2

Fig. 1. Occlusal view of erinaceine dentition showing points between
which tootli measurements were taken. Abbre\"iations: a, anterorposterior
diameter; t, transverse diameter; tl w, talonid width; tr I, trigonid length;
tr w, trigonid width. For P-P, \P, L-Pi, and Ms, the transverse diameter, /,
is defined as the greatest distance from the buccal to the lingual borders of
the tooth measured perpendicular to the anteroposterior diameter, a, but not
necessarily on a single line. For M", the anteroposterior diameter, a, is defined
as the greatest distance from the anterior to the posterior borders of the tooth
measured perpendicular to the transverse diameter, t, but not necessarily on a
single hne.

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

of eleven differences he considered significant between the two
(Koerner, 1940:841).

Although Butler (1948:490) suggested that Parvericius might
belong to the tribe Erinaceini (subfamily Erinaceinae), he tenta-
tively placed it in the Brachyericini (subfamily Neurogymnurinae)
in his classification of the Erinaceidae (1948:488). Butler (loc. cit. )
regarded the M^ as more transverse and the M^ more reduced in
Parvericius montonus than in Amphechinus edicardsi. Examination
of Koerner's illushation of the upper dentition of P. montanus (see
also Fig. 12a, this paper) and Hiirzeler's of A. edicardsi reveals that
the M^ transverse width is equal to the distance from the buccal
edge of the M- opposite the metacone to the buccal side of the
hypocone on both specimens (Koerner, 1940:P1. 1, Fig. la; Hiirze-
ler, 1944: Fig. 4). Direct examination of the type of P. montamis
(YPM 13956) confirms this relationship in that specimen. In the
same way, the anteroposterior dimensions of the M-^ on the two
specimens appear to be proportionally the same, and morphologi-
cally the two teeth are quite similar. Therefore, it cannot be said
generally that P. montanus has a more reduced M'^ than is found
on A. edicardsi.

In 1960, R. W. Wilson identified an isolated lower molar from
the Hemingfordian lower Pawnee Creek Formation of northeastern
Colorado as an Mo of Metechinus marsJandensis. This specimen
now appears to be an Mi of either Parvericius or Stenoechinus, new
genus.

Friant (1961) placed Parvericius in her broadly defined Erina-
ceinae.

Van Valen (1967:273) synonymized Parvericius with Amphech-
inus for three reasons. At the time, Val Valen regarded it likely
that material referred to Metechinus marslandensis was the other-
wise unknown lower dentition of Parvericius montanus. On this
basis, he assumed that Parvericius was similar to Amphechinus in
possessing an enlarged lower incisor. Subsequently, however, the
material referred to M. marslandensis has been shown to be the
lower dentition of Brachyerix macrotis, an erinaceid quite unlike
P. montanus (Rich and Rich, 1971:12-13). Secondly, Van Valen
thought that both Metechinus (a genus he thought to have been
derived from Parvericius) and Amphechinus had foreshortened
skulls. However, comparison of figures of the skulls of Amphechinus
and other erinaceines such as Erinaceus, reveals that the skull of
Amphechinus is as elongated, if not more so, than those of other
members of the Erinaceinae (cf. Viret, 1938: Fig. 1). Finally, Van
Valen regarded the teeth of Amphechinus and Parvericius as quite
similar. Although the available teeth of Parvericius do resemble
those of Amphechinus, they show equally strong resemblances to
other known erinaceines.

NORTH AMERICAN ERINACEINE HEDGEHOGS

Fig. 2. Stenoechinus tantalus, n. gen. and n. sp. A. Lateral view; B. Occlu-
sal view; C. Lingual view of holotype, KU 18001, right mandible, from the late
Arikareean, upper Cabbage Patch beds, Tavenner Ranch locality 2 (KLT-Mt-
21), NW ^4, NW J4, SW J4, SW ]i, sec. 11, T 8 N, R 10 W, Powell Co., Montana
( = locality 1, Wood and Konizeski, 1965:462). D. Occlusal view of KU
18406, left mandible, from the medial Arikareean, middle Cabbage Patch beds.
Cabbage Patch locality 13 (KU-Mt-46), CSW }i, SE Vi, sec. 1, T 10 N, R 12 W,
Granite Co., Montana (=MV6547, Rasmussen, 1969:132). X6.

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 3. Stenoechinus tantalus, n. gen. and n. sp., KU 18002, left mandible.
A. Lateral view. B. Occlusal view. C. Lingual view. From the late Arikareean,
upper Cabbage Patch beds, Tavenner Ranch locality 2 (KU-Mt-21), NW %,
NW }i, SW %, SW }i, sec. 11, T 8 N, R 10 W, Powell Co., Montana ( = locality
1, Wood and Konizeski, 1965:462). x6.

Mandibles of Parvericius montamis described here for the first
time support the placement of this species in a genus other than
Amphechinus. Although the two genera share the elongate I] that
has so impressed previous workers, this new material demonstrates
that the Mi trigonid of Parvericius is anteroposteriorly compressed
rather than being anteroposteriorly expanded as in Amphechinus.
Although but a single difference, in a group as highly uniform as
the Erinaceinae, we consider it of generic significance.

Palaeoerinaceus minimus Bohlin (1942) was described first from
Taben-buluk, Kansu, China; recently, additional material from
Nareen Bulak, Mongolia, has been assigned tentatively to that
species by Sulimski (1970) under the designation AmpJiecliinus
{Palaeoerinaceus) cf. 7ninimus. Both localities ha\'e yielded faunas
regarded as somewhat younger than that from the Hsanda Gol
Formation of Mongolia and medial or late Oligocene in age (Mel-
lett, 1968:9; Sulimski, 1970:69). Although P. mini7nus apparently
has the enlarged Ii characteristic of all members of Amphechinus,
the Ml trigonid is not anteroposteriorly expanded; therefore, this
species is transferred to Parvericius with which it agrees in both
characters. Furthermore, the few mandibles of Parvericius 7ninimus

NORTH AMERICAN ERINACEINE HEDGEHOGS 7

can be distinguished from the few of Parvericitis montanus only by
the greater depth of the latter, a difference probably due to onto-
genetic stage of dev^elopment. Therefore, P. minimus is regarded
here as a junior synonym of P. montanus.

Despite the removal of Parvericius montanus from Amphech-
inus, the latter genus still is kno\\ai in North America because
Palaeoerinaceus horncloudi J. R. Macdonald (1970) from the early
Arikareean Monroe Creek Formation of South Dakota may be as-
signed to Amphechinus following Butler's action synonymizing the
two genera (Butler, 1948:473). J. R. Macdonald (1970:21) dis-
tinguished A. horncJoudi from other species of the genus by the
talonid having a greater width than the trigonid on P4 and the
shorter trigonid on Mi. However, the latter character does not
appear valid when figures of other species of Amphechinus are com-
pared to the type specimen of A. horncJoudi. (For example, com-
pare figures of the following species of Aviphechinus: 1) A. horn-
cJoudi, Macdonald, 1970: Fig. 5; this paper: Fig. 9; 2) A. edwardsi,
Hiirzeler, 1944: Fig. 13; 3) A. intermedius, Viret, 193S: Fig. 9;
and 4) A. arvernensis, Viret, 1929: PI. 28, Fig. lb.) Relative to
either the width of the trigonid or the length of the tooth, the
anteroposterior length of the Mi trigonid is as great or greater on
A. horncJoudi as on any other species in the genus.

From the same lithic unit and general area that yielded the
type specimen of AmpJiecJiinus JiorncJoudi, L. J. Macdonald (1972)
recently has reported the presence of additional specimens of that
species (under the name MetecJnnus marsJandens^is) together with
specimens of Parvericius montanus.

SYSTEMATIC ACCOUNTS

Class Mammalia Linnaeus, 1758

Order Insectivora Illiger, 1811

Superfamily Erinaceoidea Fischer von Waldheim, 1817

Family Erinaceidae Fischer von Waldheim, 1817

Subfamily Erinaceinae Fischer von Waldheim, 1817

Stenoechinus new genus

Type Species. — StenoecJiinus tantaJus new species.

Known DistriJndion. — Late Arikareean (22.5-21 my, early Mio-
cene), North America.

Diagnosis. — Distinguished from AetJiecJiinus, AmpJiecJiinus,
AteJerix, DimijJecJunus, Erinaceus, GymmtrecJiinus, HemiecJiinus,
MioecJiinus, ParaecJiinus, Parvericius, and PostpaJerinaceus by the
greater width of the M^ relative to its length; from AmpJiecJiinus,
DimyJechinus, MioecJiinus, PaJaeoscaptor, and PostpaJerinaceus by
an anteroposteriorly compressed Mi trigonid; from AetJiecJiinus,
AmpJiecJiinus, AteJerix, Erinaceus, HemiecJiinus, MioecJiinus, Para-

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 4. Stenoecliinus tantalus, n. gen. and n. sp., occlusal \ie\v. A. KU
18359, right M\ B. KU 18098, right M". Same locality as in figure 3. xl2.

echinus, Parvericius, and Postpalerinacetis by the presence of a
prominent postcingulum on Ms; from PaJaeoscaptor by a talonid
on Mo reduced to a prominent postcingulum; and from DimijJech-
inus by presence of M.;.

EUjmolo'^ij. — Stenos, Greek, meaning narrow, referring to the
proportions of the M\ M-, Mi trigonid, and M3 talonid; echinos,
Greek, meaning hedgehog.

Stenoechinus tantalus new species
Figures 2-4

Holotype. — KU 18001, right manible fragment with Mi-.;, lacking
ascending ramus and horizontal ramus anterior to Mi except for
the medial wall of the horizontal ramus that has traces of four
alveoli immediately anterior to Mi.

Type Localifi/ and Sfratigrapliic Position. — Upper Cabbage
Patch beds, Tavenner Ranch locality 2 (KU-Mt-21), NW }i, NW 'A,
SWJi SW }i, sec. 11, T 8 N, R 10 W, Powell Co., Montana (= Lo-
cality no. 1, Wood and Konizeski, 1965:462).

Diagnosis. — Only known species of genus.

Etymology. — Tantalos, Greek, mythological character symbolic
of eternal torment.

Referred Material— KU 18002, left mandible fragment with P4,
Ml, and that part of the mandible immediately below these two
teeth. KU 18003, right mandible fragment with M-, alveolus for
Mo, and lacking horizontal ramus anterior to M^, ascending ramus,
and angle. KU 18004, isolated left M,. KU 18098, isolated right
M-, heavily worn. KU 18342, edentulous left mandible fragment
with alveoli for M2-:!, and lacking horizontal ramus anterior to M;.
alveoli, ascending ramus, and angle. KU 18354, isolated right M,.
KU 18356, left mandible fragment with M, and alveoli for M.-,„
lacking horizontal ramus anterior to M, and all of mandible behind
M, alveolus. KU 18359, isolated right M^. KU 18404, isolated right
M, trigonid. All the material referred to in this paragraph was
collected at the same locality' as the t\'pe specimen.

NORTH AMERICAN ERINACEINE HEDGEHOGS 9

KU 18406, left mandible fragment with M1-3, lacking ascending
ramus and horizontal ramus anterior to Mi. Found in the middle
Cabbage Patch beds, Cabbage Patch locaHty 13 (KU-Mt-46), CSW
)i, SE )i, sec. 1, T 10 N, R 12 W, Granite Co., Montana ( = MV6547,
Rasmussen, 1969:132).

Table 1. Measurements (mm) of the Upper Dentition of Stenoechinus tantalu

and Palaeoscantor acridens

S. tantalus S. tantalus P. acridens
KU 18098 KU 18359 AMNH 22080

M^ anterorposterior diameter — 1.7 2.2

M\ transverse diameter - 2.2 2.6

M", anteroposterior diameter 1.2 — . 1.6

M", transverse diameter 1.9 — . 2.2

Lower Dentition. — (Figs. 2 and 3) Only indirect data are avail-
able to give an indication of the dental condition anterior to P4. In
transverse section, the horizontal ramus of KU 18001 has a medial
and a lateral layer of compact bone with a space in between. On
the internal side of the medial wall of this specimen, immediately
anterior to the Mi, are preserved traces of the partitions between
the next four anterior alveoH ( Fig. 2a ) . The most posterior of these
alveoli were occupied by the roots of P4 (Fig. 3a, c). Although the
forward wall of the most anterior alveolus preser\ed is absent,
enough of that alveolus is preserved to indicate that it probably had
a significantly greater anteroposterior diameter than the one im-
mediately behind. A similar size relationship exists between the
same alveoli on a specimen of another small erinaceine, Paloeoscap-
tor cf. acridens (AMNH 22082). On that mandible, the more pos-
terior of these two aheoli was occupied by the rear root of P3 and
the more anteroposteriorly elongated anterior alveolus was occupied
by the forward root of P3 and the single root of P2 (Fig. 5). If
there was a bony partition between the forward root of P;{ and the
root of Pl>, it has been lost without a trace on both KU 18001 and
AMNH 22082 and must ha\e been much thinner than the other
alveolar walls, for these two roots are quite close to one another
on AMNH 22082.

The most anterior alveolar waW preserved on the medial wall of
the mandible of KU 18001 extends nearly to its ventral border. In
erinaceids such as AnipJieduniis? rectus (AMNH 22084), which
have an enlarged Ii root alveolus extending posteriorly to the region
immediately in front of the anterior root of P4, the alveolar borders
of the C1-P3 may be traced on the internal side of the medial wall
of the ramus from the dorsal lip of their respective alveoli only
halfway to the ventral border of the mandible. At that point, the
alveolar walls of C1-P3 are cut off bv the aheolar wall of Ii because

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Fig. 5. Palaeoscaptor cf. acridens, AMNH 22082, left mandible. A. Lateral
view. B. Occlusal view. From the medial Oligocene, Hsanda Gol Formation,
Tsagan Nor Basin, Mongolia. x6.

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

the root of Ii passed medial to the roots of C1-P3. Hence, the root
of Ii in Stenoecliinus tantalus did not extend as far posteriorly and
was probably closer in relative size to the more reduced I] found
in the living Erinaceus europaeus.

P.;. — Tallest cusp on the tooth, the slender protoconid is half
again as high as the paraconid above the base of the enamel, the
height of the protoconid being one and one-fourth times the length
of the tooth. On the single specimen in which the protoconid is
preserved (KU 18002), the lingual third of that cusp has been
lost; thus, the casual observer might believe that the metaconid is
placed well away from the protoconid base, whereas the bases of
the two cusps are actually quite close together. The paraconid is
only slightly lower than the metaconid and is situated on the mid-
line of the tooth at its anterior edge; the cusp lies so far forward
that its base projects forward beyond the anterior root of the tooth.
Along the rear margin of the tooth is a well developed posterior
cingulum that dips steeply ventrobuccally. Continuous with this
posterior cingulum is a weak buccal cingulum that is developed
along the entire length of the tooth.

Ml. — The length of the trigonid is slightly more than half that
of the tooth. The trigonid is slightly shorter than wide; the proto-
conid is the tallest of the trigonid cusps and has the greatest basal
dimensions. Intermediate in height, the metaconid is anterolingual
to the protoconid; the two cusps are linked by a protolophid that
has an acute V-shaped profile in posterior view. The paralophid
links the protoconid with the paraconid, the lowest cusp on the
trigonid. In lateral view the paralophid has a V-shaped profile; the
angle between the two segments of the V is obtuse but close to a
right angle. The lingual segment of the paralophid is nearly hori-
zontal. The paraconid lies at the anterolingual corner of the tooth.
In lingual view, the paraconid axis is either vertical and parallel
to the metaconid axis, or anterodorsally inclined and forms an angle
as great as 30 degrees with respect to the metaconid axis.

Width of the talonid is equal to, or slightly greater than that
of the trigonid, with the entoconid and hypoconid at the extreme
posterior corners of the tooth. The entoconid is the taller of the
talonid cusps and is intermediate between the paraconid and meta-
conid in height. The entoconid is posterior to the metaconid and
the hypoconid lies slightly more labial than the protoconid. In an-
terior view, the buccal margin of both the protoconid and hypoconid
are buccally convex. Extending anteriorly from the hypoconid, the
cristid obliqua abuts against the posterior wall of the trigonid at a
point slightly buccal to the tip of the protoconid. A low entocristid
closes the talonid basin lingually. A well developed but narrow
cingulum extends along the entire buccal side of the tooth from a
point below the paraconid to the base of the hypoconid where it
passes into the posterior cingulum; the latter is directed dorsomedi-

NORTH AMERICAN ERINACEINE HEDGEHOGS 13

ally so that it terminates lingually, midway between the hypoconid
and entoconid just below the crest of the postcristid. In the region
of transition between the buccal and posterior cingula, adjacent to
the hypoconid base, these narrow cingula are even narrower.

No specimen is available with the alveoli of this double-rooted
tooth exposed.

M>. — Because this tooth is quite similar to Mi in general mor-
pholog}', a detailed description is unnecessary, but a few points of
difference are noted.

In length and width, this tooth is about four-fifths as large as
the Ml. No distinct paraconid is present; the paralophid terminates
lingually without any noticeable swelling. The protoconid and meta-
conid are equal in height.

The entoconid is only slightK lower in height than the metaconid
and markedly taller than the lingual region of the paralophid where
the paraconid would be expected if developed. Despite the fact
that the talonid is slightly narrower, not equal to, or slightly wider
than the trigonid, the hypoconid position is still somewhat more
buccal than that of the protoconid. The buccal cingulum is devel-
oped as extensi\'ely as on the M], and terminates anteriorly at the
forward edge of the pre\ allid.

Although the al\ eoli of this double-rooted tooth are equivalent
in width, the posterior aKeolus is nearb- circular and about twice
as long anteroposteriorU' as the elliptical anterior alveolus.

M.i. — This double-rooted tooth is markedK' different than the
more anterior molars; it lacks a well developed talonid, having only
a wide postcingulum. The trigonid is anteroposteriorly expanded
to the extent that it is slightK- longer than wide. Of the two well
developed trigonid cus'os, the protoconid is the taller and has greater
basal dimensions. Its base and that of the lower, smaller metaconid
are in close juxtaposition; thus, the protolophid bet\veen the t\vo
cusps is short. A small, obtuse, V-shaped notch is present near the
middle of the protolophid when \iewed from the rear. The lingual
end of the paralophid lacks a swelling marking the presence of the
paraconid. As in the more anterior molars, the paralophid in lateral
\iew has a V-shaped profile; the two limbs of the \^ meet at nearly
right angles with the lingual segment nearly horizontal. A well-
developed, continuous, and narrow basal cingulum is present along
the buccal margin of the tooth; the basal cingulum passes into a
much wider postcingulum that lies along the entire posterior side
of the trigonid. This postcingulum is widest midway between the
protoconid and metaconid, and tapers to minima at both the lingual
and buccal margins of the tooth.

The M.-i is double-rooted in the holotvpe ( KU 18001 ) , but in two
other specimens (KU 18003 and KU 18356) where alveoli of this
tooth are known, onlv a single, anteronosteriorlv elongated root is
indicated; thus, suggesting the condition of the root of M^ may vary

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

from specimen to specimen. On one of these specimens where the
alveolus is exposed, a single elongated root is preserved just below
the alveolar lip (KU 18356).

Mandible. — (Figs. 2 and 3) The preserved fragments of the
mandible indicate that it is slightly deeper below Mi than M3. Be-
tween those two areas, the ventral border is nearly straight, but
behind the Ma the ventral border curves sharply upward. The pos-
terior border of the mental foramen is preserved on KU 18002
adjacent to the anterior root of P4, midway between the dorsal and
ventral margins of the mandible. Beneath the Mi of KU 18001
there is a slight ridge on the buccal side of the mandible near its
ventral border. Presumably, this is the lateral border of the dia-
gastric fossa. No diastema are developed between P^ and M^.

Upper Dentition. — Two isolated upper molars from Tavenner
Ranch locality 2 probably represent the otherwise unknown M^
and M2 of Stenoechinus tantalus (KU 18359 and KU 18098, re-
spectively ) . The relative dimensions of these two upper molars are
similar to those of the homologous teeth on a specimen of Palaeo-
scaptor acridens from the Hsanda Gol Formation of Mongolia in
which the left jaw was found in occlusion with the skull (AMNH

Fig. 6. Palaeoscaptor acridens, AMNH 22080, partial skull, occlusal view
of right upper dentition. From the medial Oligocene, Hsanda Gol Formation,
Tsagan Nor Basin, Mongolia. X6.

NORTH AMERICAN ERINACEINE HEDGEHOGS 15

22080), (Table 1). In addition, when the dimensions of the upper
and lower teeth on this specimen of P. acridens are compared, the
ratios are found to be similar to those between the lower teeth of
S. tantalus and these two upper molars (compare Tables 1 and 2).
In morphology, the M^ is similar to that of P. acridens (compare
Figs. 4a and 6). Extensive wear during life all but obliterated the
cusps and trigon basin on the M-; howe\'er, enough remains to show
that in outline this tooth is similar to the M- of P. acridens. A
further reason for associating these upper molars \\'ith the lowers
in the type specimen of S. tantalus is that no other erinaceid is
known from Tavenner Ranch locality 2 except the much larger
Amphechinus horncloudi.

M'. — (Fig. 4a) The length of this tooth is about three-fourths
its width. Of the four principal cusps on the tooth, the metacone
is the tallest, its height being slightly more than half the length of
the tooth. The protocone and paracone are equal to one another
in height and both are equal to the metacone in basal dimensions.
Lowest of the four principal cusps and smallest in basal dimensions
is the hypocone. The protocone is slight])' forward of a point di-
rectly lingual to the paracone; the hypocone is lingual to the meta-
cone. A line passing through the paracone and metacone is parallel
to one drawn through the protocone and hypocone. The parastylar
spur is directed anteriorly away from the main body of the tooth;
the metastylar spur is directed posterobuccally. In occlusal view,
the lingual, buccal, and posterior borders of the tooth are markedly
concave, whereas the anterior border is broadh' conxex. Close to
the convex anterior border are the preprotocrista and preprotoconule
crista which grade imperceptibly into one another, because no
paraconule is developed. Th's crest extends from the anterobuccal
side of the prolcccnc to the \ic-nit>^ of the anterior side of the para-
cone base from whch it is separated by a shallow notch. In anterior
view, this lor.h has a profile rem.i is?ent of an inverted, broad, trun-
cated letter V; the margins arc formed by the steeply sloping sides
of the two cusps an-l the horizontal midsection is formed by the
crest between tncni. The pcst^rctocr'sta is directed posterobuccally
from the protocone. Th's ciista divides at a point lingual to the
anterior edge of the melaconc; one branch extends posterolingually
to the hypocone and the ether extcn-^s buccally to the metacone.
No trace of a metaconule is present. Dc\ eloped along the anterior
edge of the tooth, ike narrcv/ precingulum curves apically at its
buccal end, there joining the anterior terminus of the short para-
crista. The poslcror Icrmnus of the paracrista is at the base of
the paracone. A short rcntrccrista hnks the bases of the paracone
and metacone. From the metacone, the metacrista extends in a
buccal and slightly posterior direction along the posterior edge of
the metastylar spur for a distance approximately equal to one-
fourth the length of the tooth. Along the buccal margin of the

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

tooth between the paracone and metacone is an extremely weak
ectocingukmi. Three roots are developed on this tooth — one above
the paracone-paracrista region; a second over the metacone-meta-
crista region; and the third lingual root over the protocone-hypocone
region. The anterior and posterior borders of the third lingual root
converge toward its dorsal tip in contrast to other erinaceines in
which the borders are parallel.

M~. — (Fig. 4b) In basic outline, the tooth is a right triangle
in which the hypotenuse represents the anterior margin. The nar-
row parastylar spur projects buccally and slightly anteriorly away
from the main body of the tooth. The metastylar spur projects
posteriorly and the base of the hypocone forms a third spur that
projects posterolingually away from the main body of the tooth.
A narrow ectocingulum lies buccal to the paracone; as it passes
posteriorly, buccal to the metacone, the ectocingulum narrows. A
short metacingulum is developed behind the metacone. A precingu-
lum is developed along the anterior side of the tooth. The pre-
cingulum extends from the buccal region of the protocone base to
a point slightly lingual to the paracone where a prominent wear
facet on the parastylar spur has obliterated it. Three roots are
developed on this tooth — one above the paraconc-paracristra region,
a second above the metacone-metacrista region, and a third over
the Drotocone.

Comparisons. — Sfenocchinus fanfahis differs from Polaeoscaptor
acridens found in the medial OHgocenc of Mongolia in five impor-
tant characters: M^ talonid reduced to a orominent postcingulum;
Ml trigonid less anteroposteriorly exDanded; P4 paroconid markedly
lower than the protoconid; P4 protoconid height less than tooth
length; and I, root not extending as far posteriorly. Sulimski (1970:
63) characterized FaJaeosca^iior as having a more reduced Ii than
Amphechinus (t= Palaeoerinaceus of Sulimski's usage). ^ However,
examination of an X-ray photograph of the onlv known mandible
of P. acridens containing an Ii ( AMNH 22080) shows that both the

1 Following Butler (1948), Sulimski (1970:63) placed Palaeoerinaceus
Filhol (1879) and PaJaeoscaptor Matthew and Granger (1924) in Amphech-
inus Aymard (1850). In addition, he formally divided Amphechinus into two
subgenera: Palaeoerinaceus and Palaeoscaptor. The basis for this division
was the relative size of the L and single-rooted and reduced talonid condition
of the M.;. Although a re-examination of specimens of Palaeoscaptor in the
American Museum of Natural History which were tmavailable to Sulimski indi-
cates that all species included by him in Amphechiruis shared an enlarged L,
the second criterion, condition of the M.-,, continues to appear valid. By this
valid criterion, the type species of Amphechinus, A. arvernensis, clearly belongs
in the subgenus Palaeoerinaceus for the M:; is single-rooted and possesses a
reduced talonid. According to article 44(a) of the International Code of
Zoological Nomenclature (Stoll et ah, 1961), if a genus is divided into sub-
genera, the subgenus containing the type species must be placed in a subgenus
with the same name as the genus. For this reason, the name for the subgenus
Palaeoerinaceus should be changed to Amphechinus.

NORTH AMERICAN ERINACEINE HEDGEHOGS

Fig. 7. X-rays of mandibles in lateral view. A. Palaeoscaptor
acridens, AMNH 22080, reversed left mandible, from the medial
Oligocene, Hsanda Gol Formation, Tsagan Nor Basin, Mon-
golia. x4. B. Amphechinus homdoudi, SDSM 62113, type,
right mandible, from tlie Arikareean, Monroe Creek Formation,
T 39 N, R 42 W, Shannon Co., South Dakota. x3. C. Amph-
echinus horncloudi, KU 18162, right mandible, from the medial
Arikareean, middle Cabbage Patch beds, Cabbage Patch lo-
cality 3 (KU-Mt-11), SE )i, NE )i, SE }i, SE )i, SW )i, sec. 7,
T 10 N, R 11 W, Powell Co., Montana (= MV6617, Rasmussen,
1969:140). x3. D. Parvericius montanus, UMMP \'56569,
reversed left mandible, from the late Barstovian, Egelhoff
Quarry, from an unnamed lithic unit beneath tlie Valentine
Formation and overlying tire Rosebud Fonnation, SW comer
of NE }i, SW %, sec. 29, T 33 N, R 23 W, Keya Paha Co.

Nebraska.
DDS.

X4. X-rays taken by Dr. Howard K. Bloomfield,

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

crown and root are as extensively developed in that species as in
the various species of Amphechinus (compare figures 7a and 7b,
this paper and Hiirzeler, 1944: Fig. 7). Except for the features
mentioned above, the molars of S. tantalus and P. acridens are quite
similar in morphology, relative proportions, and size.

The species of Amphechinus from the Oligocene and Miocene
of Europe, Asia, Africa, and North America differ from Steno-
echimis tantalus by having an anteroposteriorly expanded trigonid
on Ml. Together with Parvericius montanus, the species of Amph-
echinus differ further in having an enlarged Ii with a root that
extends posteriorly to the region immediately anterior to the forward
root of P4; P4 paraconid nearly as tall as the protoconid; Pi proto-
conid height subequal to the tooth length; complete absence of a
talonid on M3; and M^ and M- more expanded anteroposteriorly.

The species of Gijmnurechinus from the Miocene of East Africa
described by Butler ( 1956, 1969 ) arc approximately twice as large

Fig. 8. Amphechinus horncluudi, KU 18097, left maxilla fragment. A.
Occlusal view. B. Lateral view. From same locality as in figure 3. X3.

NORTH AMERICAN ERINACEINE HEDGEHOGS

Fig. 9. Amphechinus horncloudi, KU 18162, right mandible.
A. Lateral view. B. Occlusal view. C. Lingual view. From the
medial Arikareean, middle Cabbage Patch beds, Cabbage Patch
locality 3 (KU-Mt-11), SE J4, NE ]i, SE )i, SE ]',, SW )i, sec. 7,
T 10 N, R 11 W, Powell Co., Montana (= MV6617, Rasmus-
sen, 1969:140). X3.

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 10. Amphechinus horncloudi, KU 18163, skull frag-
ment. A. Left lateral view. B. Palatal view. C. Right lateral
view. From the medial Arikareean, middle Cabbage Patch
beds, Cabbage Patch locality 2 (KU-Mt-9), SE }i, NW %, SW %,
SE }i, sec. 1. T 10 N, R 12 W, Granite Co., Montana ( =
MV6504-5, Rasmussen, 1969:131-132). x3.

as Stenoechiyius tantalus. In addition, the African species differ in
that the paraconid is large and nearly as tall as the protoconid on
P4; and the M^ and M- are expanded more anteroposteriorly.

The anteroposteriorly expanded condition of the lower molar
trigonids indicates that Stenoechimis tantalus is too advanced or
derived to be considered a member of the Adapisoricidae as Van
Valen (1967) constituted that group. Among late Mesozoic and
early Tertiary insectivores including the adapisoricids, the lower

NORTH AMERICAN ERINACEINE HEDGEHOGS 21

molar trigonid is characteristically anteroposteriorly compressed as
compared with the condition commonly observed among several
geologically younger groups of insectivores; e.g. Erinaceidae, Tal-
pidae, and Soricidae.

Within the Erinaceidae, Stenoechinus tantalus is more closely
allied with the Erinaceinae than Galericinae due to the reduction
of the talonid on M3 to a prominent postcingulum, a condition
rarely encountered among other insectivores and hence considered
derived. However, if S. tantalus is assigned correctly to the Erina-
ceinae, it is the most primitixe member of that subfamily because
its P4 paraconid lacks the prominence that is the unique derived
character state which distinguishes every other member of the
subfamily from all other insectivores.

If Stenoechinus tantalus is the most primitive known member of
the Erinaceinae, the known forms most closely related to that
species' immediate ancestors are presumably members of the Adapi-
soricidae. Found in Europe, Asia, and North America, this family
ranges temporally from Paleocene to Oligocene. For this reason,
it is not possible to select between the hypotheses that the stock
that gave rise to S. tantalus was part of the general Arikareean
invasion of North America or that the lineage had a prior Oligocene
history in North America.

Amphechinus Aymard 1850

Type Species. — Amphechinus arvernensis ( deBlainville 1838).

Known Distribution. — Stampian? — Vindobonian ( ?35 — 14 my,
early? Oligocene — medial Miocene), Europe; Aquitanian — Vallesian
(20 — 11 my, early to late Miocene), Africa; 32 — 31 my (medial
Oligocene), Asia; medial Arikareean — medial Barstovian (23 — 14
my, early to medial Miocene), North America.

Diagnosis. — Distinguished from Palaeoscaptor by absence of a
metacone on M'; from Palaeoscaptor and Stenoechinus by the
greater length of the M^ relative to its width; from Aethechinus,
Atelerix, Erinaceus, Gijmnur echinus, Hemiechinus, Mioechinus,
Paraechinus, Postpalerinaceus, and Stenoechinus by presence of
an enlarged Ii; from Aethechinus, Atelerix, Erinaceus, Gymnur-
echinus, Hemiechinus, Paraechinus, Parvericius, and Stenoechinus
by an anteroposteriorly elongated trigonid on Mi; from Gymnur-
echinus, Palaeoscaptor, and Stenoechinus by absence of a talonid
or postcingulum on M3; and from Dimylechinus by presence of
M3 and M^.

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Amphechinus horncloudi (J. R. Macdonald, 1970)
New Combination
Figures 7b-c, 8-10
Palaeoerinaceus horncloudi J. R. Macdonald 1970:20.

Holotype. — SDSM 62113, fragment of right mandible with
Po-Mi, damaged Ci, roots of Ii-o, and anterior root of Mo.

Type Locality and Stratigraphic Position. — Collected from the
Monroe Creek Formation in T 39 N, R 42 W, Shannon Co., South
Dakota (SDSM V6229; more precise locality information is on file
at the South Dakota School of Mines and Technology, Museum of
Geology).

Diagnosis. — Distinguished from other species of Amphechinus
by the talonid being greater in width than the trigonid on P4.

Referred Material— K\J 18097, left maxilla fragment with P^ P^
(except metacrista), M^ and anterior alveoli of M^. KU 18405,
right P^ fragment. From the upper Cabbage Patch beds, Tavenner
Ranch locality 2 (KU-Mt-21), NW )i, NW K, SW )i, SW K, sec. 11,
T 8 N, R 10 W, Powell Co., Montana ( = locality 1, Wood and
Konizeski, 1965:462).

KU 18162, right mandible with partially erupted I], P4, fully
erupted M3, alveoli for I2, Ci, P2, and Mi-o, missing region posterior
to base of angle. Found in the middle Cabbage Patch beds, Cab-
bage Patch localitv 3 (KU-Mt-11), SE )i, NE 'A, SE )i, SE }i, SW
M, sec. 7, T 10 N, R 11 W, Powell Co., Montana ( = MV6617, Ras-
mussen, 1969:140).

KU 18163, anterior half of skull with broken right P; complete
right P-P-; missing lateral part of left palate and lateral part of

Table 3. Measurements ( mm ) of tlie Upper Dentition of
Amphechinus horncloudi

KU 18097 KU 18163

P, anteroposterior diameter 1.1

F, transverse diameter — . 0.9

P, anteroposterior diameter — _ 2.0

F, transverse diameter .— 1.1

C\ anteroposterior diameter — . 2.6

CS transverse diameter — _ 1.2

P^ anteroposterior diameter -— 1.7

P^, transverse diameter — _ 0.9

P^ anteroposterior diameter 2.1 —

P^ transverse diameter 1.7 —

P\ anteroposterior diameter 3.5" —

P\ transverse diameter 3.3 —

M\ anteroposterior diameter 3.4 —

M\ transverse diameter 3.7

' Posterior limit of tooth estimated by assuming contact with parastylar spur
onM\

NORTH AMERICAN ERINACEINE HEDGEHOGS 23

Table 4. Measurements (mm) of the Lower Dentition of
Amphechinus horncloudi

type, SDSM 62113 KU 18162

Mandible depth below Mi anterior root . - 4.0 3.3

Mandible depth below M2 posterior root — . — . 3.9

P4, anteroposterior diameter 2.5 2.7

P4, transverse diameter 2.0 1.9

Ms, trigonid length — - 1.1

Ms, trigonid width — - — - 0.9

right palate behind P-; casts of olfactory bulbs, chambers for ethmo-
turbinals, and left maxillary sinus exposed. Found in the middle
Cabbage Patch beds, Cabbage Patch locality 2 (KU-Mt-9), SE M,
NW )i, SW )i, SE )i, sec. 1, T 10 N, R 12 W, Granite Co., Montana
(= MV6504-5, Rasmussen, 1969:131-132).

Lower Dentition. — (Fig. 9) /;. — Near the tip of this tooth, its
cross section is crudely circular with noticeable flattening on the
medial and anterior sides. The crown is anterodorsally inclined.

I2, Ci. — In dorsal view, the outline of the single alveolus for
each of these teeth is an oval with the broad end posterior and the
major axis directed anteroposteriorly. The maximum length and
width of the alveolus for I2 are 1.0 and 0.5 mm respectively; similar
dimensions for Ci are 1.6 and 0.9 mm. Both alveoli are antero-
dorsally inclined.

Pg. — The single alveolus for this tooth is too damaged to allow
meaningful statements concerning its outline or dimensions. It
seems to have been about the size of the alveolus for L..

P;. — Of the trigonid cusps, the protoconid is tallest and largest
in basal dimensions. The paraconid is anteromedial to the proto-
conid, from which it is separated by a distinct notch. In height,
the paraconid is equal to the metaconid, but in basal dimensions,
the paraconid is much larger. The protolophid is directed lingually
from the posteromedial corner of the protoconid. At the site of
the metaconid at the lingual end of this lophid, there is no swelling
to mark the presence of the cusp. The talonid is a narrow ridge
along the posterior border of the tooth with its highest point behind
the protoconid. The maximum transverse diameter of this tooth
occurs in the talonid region.

A/7. — The forward alveolus for this double-rooted tooth is trap-
ezoidal in outline; the base of the trapezoid is formed b>' the medial
wall of the alveolus. The maximum anteroposterior measurement of
this alveolus is 1.3 mm and the maximum mediolateral measurement
is 1.1 mm. The shape of this alveolus may not accurately reflect the
condition that would be found in a fully mature individual, because
the unerupted P4 talonid forms the anterior border, and no bony
partition is preserved between the tooth and alveolus. The posterior

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

alveolus is roughly rectangular in outline; the anterioposterior length
is 1.2 mm and the mediolateral width is 1.3 mm. The bony parti-
tion between the forward and rear alveoli displays the paiied bony
ridges common in erinaceids.

M^. — In outline, the two alveoH for this tooth are crudely circu-
lar except in the area of the bony partition which separates them
from one another; here they are straight or actually concave in the
case of the anterior alveolus. Their diameters are approximately
1.0 mm. As on the Mi, the bony partition between the two alveoU
has bony ridges.

Ms. — The prevallid is expanded anteriorly, the length of the
tiigonid being about 10 percent greater than the width. No distinct
paraconid is developed at the anterior end of the paralophid; this
region is markedly lower than the posterior area. Along the rear
of the trigonid is a tall protolophid. At the lingual and buccal ends
of this crest there are slight swellings that represent the metaconid
and the slightly taller protoconid, respectively. No talonid is de-
veloped, but a narrow postcingulum is present that is continuous
with a buccal cingulum that extends to the anterior extremity of
the tooth.

Upper Dentition. — (Figs. 8 and 10) P. — The single root of this
mediolaterally flattened tooth passes posteriorly above the roots
of the succeeding incisors and terminates near the premaxilla-
maxilla suture. Although broken off at the base of the crown, this
tooth is clearly much larger than the posterior incisors.

P-C^. — Morphologically, these three teeth are quite similar, but
there is a marked progressive increase in size posteriorly. In outline,
each tooth is crudely elliptical with the major axis directed antero-
posteriorly. P and C^ are double — rooted but P has only one root;
however on the buccal side of the P root there is a clear division into
anterior and posterior lobes. On the crown of all three teeth there
is a single, tall cusp, the height of which is nearly equal to or
slightly greater than the length of the respective teeth. The anterior
edge of this cusp dips posteroventrally and the posterior edge is
nearly vertical. The tip of the cusp is ventral to the posterior root
on the I' and C^ and to the posterior lobe on the single root on
the P. Extending posteriorly from the base of the cusp along the
midline of each tooth is a short crest which slopes posterodorsally.

P^. — Two roots are present on this small tooth. As on P-C^,
the outline of the tooth is crudely elliptical and there is a single,
prominent cusp, but its height is only two-thirds the length of the
tooth. Moreover, the tip of the cusp is further forward than on
P'-C^ for it is ventral to the anterior root. The posterior edge of the
cusp dips anteroventrally. Extending anteriorly from the base of
the cusp is a short crest on a precingulum which projects forward
of the anterior root. Extending posteriorly from the base of the
cusp is a second, much longer crest that terminates at a small

NORTH AMERICAN ERINACEINE HEDGEHOGS 25

cuspule on a cingulum which projects posteriorly beyond the rear
root.

P^. — In occlusal view, the straight buccal border of the tooth
is directed anteromedially; the anterior and posterior borders are
concave, and the lingual border is convex. The prominent meta-
crista extends posterobuccally from the paracone along the midline
of the well developed metastylar spur to the posterior tip of that
structure. The protocone is somewhat less than half the height
of the paracone, and anteromedial to it. Behind the protocone is a
flat surface that dips posterodorsally. This surface lacks any trace
of a hypocone. A narrow cingulum extends along the anterior side
of the base of the paracone. Three roots are developed on this
tooth, one above the protocone, another over the paracone, and a
third over the metacrista.

PK — This tooth is several times larger than the preceding P^.
The height of the paracone, the tallest and most prominent cusp
on the tooth, is subequal to the tooth length. The metastylar spur
projects posteriorly and somewhat laterally away from the buccal
side of the main body of the tooth; thus, the main body of the tooth
is more anteropostcriorly elongated buccally than lingually. One —
third the height of the paracone, the protocone is located lingual
and slightly anterior to that cusp, as in modern hedgehogs. The
hypocone is equal in basal dimensions to the protocone but lower.
Linking these two cusps is a low crest. A similar crest extends
anterolaterally from the protocone to the base of the paracone. On
the anteromedial side of the hypocone base are two cuspules. Pos-
terior to the hypocone is a small planar surface that dips steeply
anteroventrally. Along the posterior and medial edges of this sur-
face is a weak cingulum. Although no distinct parastyle is present,
a prominent parast\'lar spin- is de\cloped on the anterior side of the
base of the paracone. Along the margin of the parastylar spur is a
distinct cingulum. Three roots are developed on this tooth, one
about the protocone-hypocone, a second above the metacrista, and
the third above the paracone-parastylar spur.

M'. — Although this tooth is slightly shorter anteroposterior^
than the P^, it is markedly broader. The paracone and metacone
are subequal in height, their heights being about one-half the
length of the tooth. The paracone is anterobuccal to the metacone.
Third tallest cusp on the tooth, the protocone is lingual and slightly
anterior to the paracone. Posterolingual to the protocone is the
lowest of the four principal cusps, the hypocone. Unlike modern
erinaceines, the protocone and paracone are no closer to one another
than the hypocone and metacone. Because no protoconule is de-
veloped, the preprotocrista and preprotoconule crista grade im-
perceptibly into one another. Linking the protocone and paracone,
this loph extends along the anterior border of the tooth and in
anterior view has a broad U-shaped profile. The postprotocrista

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

and postmetaconule crista grade imperceptibly into one another
because no metaconule is present. They form a loph which extends
posterobuccally from the protocone and gradually turns so that it is
convex posteromedially and directed mediolaterally at its termina-
tion near the base of the metacone. This loph lacks any connection
with the hypocone, for not even a weak crest is developed between
them. A weak paracrista linking the paracone and precingulum is
developed on the small parastylar spur, which, despite its name,
lacks any trace of a distinct parastyle. From the metacone, the
metacrista extends posterobuccally for a distance equal to two-fifths
the tooth length, and terminates in a weak metastyle. Except for
the posterior tip of the metastylar spur, the entire margin of the
tooth is bounded by a weak cingulum. In occlusal view, the
anterior and medial borders are nearly straight and the buccal and
posterior borders, concave. The metastylar spur forms a prominent
projection posterobuccally away from the body of the tooth while
the parastylar spur is a much weaker anterior projection. Three
roots are developed — one above the paracone-paracrista region, a
second above the metacone-metacrista region, and a third above the
protocone-hypocone region.

M^. — The circular alveolus above the paracone-parastylar region
is approximately 0.8 mm in diameter. Posteromedial to this alveolus
is the alveolus above the metacone-metastyle region and medially
is the one above the protocone-hypocone region. Only the anterior
part of the rims of these latter two alveoli are preserved. Judging
from the placement of the three alveoli, the angle formed by the
buccal edge of this tooth with that of the M^ appears to have been
about 135°, and the width of this tooth was slightly less than that
of the Ml.

Face. — As is typical of erinaceids, the nasals are long and nar-
row, their posterior ends contacting the frontals. Due to extensive
damage on the specimen available, it is not clear how far forward
the nasals extended relative to the premaxilla or if there was direct
contact between the frontal and premaxilla. The maxilla-premaxilla
suture begins ventrally between the C^ and I'' and forms an irregular
arc that curves upward and backward. The supraorbital crest and
the region posterior to it are not preserved.

Palate. — Most of the palate is formed by the maxilla, the pre-
maxilla contributing only in the area immediately adjacent to the
upper incisors. The anterior border of the maxilla extends as far
forward as the anterior side of the P. The palatine fissure is de-
veloped on the maxilla-premaxilla suture close to the midline of the
palate. The palate extends for a short distance behind the transverse
crest, as in the Erinaccinae. Becaucs the lateral margin of the
palate has been destroyed in the region of the transverse crest on all

NORTH AMERICAN ERINACEINE HEDGEHOGS

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OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

specimens available, the presence or nature of posterior palatine
foramina in that crest cannot be determined.

A slight groove is developed on either side of the midline at the
anterior end of the palatine; it extends posteriorly to a small fora-
men. Presumably this groove is homologous to the similar one
found in galericines, in which the palatine nerve and artery lie.
Midway between the aforementioned foramen and the transverse
crest is a smaller palatine foramen. The bones of the palate are as
thick and solid as those of the living galericines. No vacuities typical
of the modern erinaceines are present in the palate.

Although the snout is more elongate than in living erinaceines,
the molars are more posterior relative to the zygomatic arch. The
elongation is reflected in the high length-to-width ratio of the palate
(Table 5). On KU 18163 this ratio" is 2.41, on species of Amph-
echinus edicardsi it is from 2.30 to 2.40, and in a sample of recent
erinaceines it ranges from 1.84 to 1.99. The location of the base of
the zygomatic arch, mainly opposite the IVP, is indicative of the
degree of posterior displacement of the molars; however, a small
segment of the base lies opposite the M-. In most modern erina-
ceines the base of the arch is equally developed above both molars.

Mandible. — (Fig. 9) The mental foramen lies below the anterior
root of P4. A wide groove extends anterodorsally from the mental
foramen and terminates below the partition between the alveoli for
P2 and Ci. In the region beneath Pj-Ni., the mandible depth is
nearly uniform. Moving away from this region in either direction,
the depth gradually decreases.

Fig. 11. Method of measuring the elevation of the anterior
edge of the ascending ramus ( / a), elevation of the condyle
{ /_ h), and inchnation of the dorsal border of the midline
symphysis { /_ c) on the lower jaw.

NORTH AMERICAN ERINACEINE HEDGEHOGS 29

Extending forward from beneath Po, the unfused symphx^sis
occupies the ventral three-quarters of the mandibular surface. The
dorsal border of the symphysis is convex dorsally and roughly paral-
lel to the ventral border, dipping posteriorly at an angle of 25°
( Z cin Fig. 11).

The ascending ramus lies slightly labial to the midline of the
mandible. Due to damage sustained by the only mandible avail-
able, the anteroposterior length of the ascending ramus, precise
position of the condyle, and extent of the angle cannot be deter-
mined. The straight anterior border of the ascending ramus is
posteriorly inclined at an angle of 76° ( Z a in Fig. 11). This angle
is the same as Butler's (1948:469-470) "elevation of anterior edge
of coronoid process." Contact bet\veen the dorsal and anterior
edges of the ascending ramus is at a gently rounded corner. A strong
lateral flange on the leading edge of the ascending ramus forms the
anterior margin of the masseteric fossa. There is a ridge near the
base and on the medial side of the ascending ramus. It extends
posterordorsally from the anterior edge of the ascending ramus,
passes immediateK^ above the mandibular foramen (which is lex'el
with the tooth row), and continues a short distance further before
being abruptly cut off by the damage described above. Because
ridges similar to this are directed at the condyle on complete
erinaccine mandibles, it appears that the condyle was quite low,
with an angle of elevation ( Z b in Fig. 11) of about 15°. This
angle is the same as the "elexation of the condyle" as defined by
Butler (loc. cit.). On the medial side of the angle can be seen
the most anterior part of the inflected ridge which is directly below
the mandibular foramen.

Comparisons. — Each of the three specimens from Montana
identified as Amphechinus horncloudi are from different localities
and stratigraphic positions in the Cabbage Patch beds. All are of
the proper size to be parts of the unknown dentition, jaw, or skull
of Amphechinus horncJoudi originally described from South Dakota.
However, with the exception of KU 18162, there is no reason other
than geographic proximit\- to regard these specimens as conspecific
with A. horncloudi rather than the slightly larger Amphechinus
edwardsi of Europe or slighth- smaller Amphechinus rusing,ensis of
Africa. See Butler (1956: Tables 4 and 7) and Hiirzeler (1944:461)
for measurements of the dentitions of these two species.

KU 18097, a maxilla fragment with P'^-M^ is similar to the
species of Amphechinus in the proportions of the upper molars.
Molars of modern erinaceines, Gymnurechinus, Mioechinus, and
Postpolerinaceus are all more anteroposteriorly elongate. Paloeo-
scaptor and Sfenoechinus have much smaller molars.

KU 18162, a mandible with Ii, P4, and M3 is similar to other
species of Ampliechinus and unlike other erinaceines except PaJaeo-
scaptor and Gymnurechinus. The P4 is similar to that of the t}'pe of

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 12. Paivericius montamis. A. YPM 13956, type, right maxilla
fragment, occlusal view, from Arikareean or Hemingfordian, Deep
River beds, sec. 3, T 10 N, R 5 E, Meagher Co., Montana. x6. B.
UMMP V61033, right P\ occlusal view, from late Barstovian, Egelhoff
Quarry, from an unnamed lithic unit beneath the Valentine Formation
and overlying the Rosebud Formation, SW corner of NE %, SW %, sec.
29, T 33 N, R 23 W, Keya Paha Co., Nebraska. xl2. C. UMMP
V61026, right M\ occlusal view, same locality as B. Xl2.

Amphechinus horncloudi (SDSM 62113) and unlike any other
known species of the genus in that the maximum transverse diam-
eter is in the talonid region, not further forward across tlie trigonid.
KU 18163, a partial skull with P-P-, resembles Amphecliinus
edwardsi and Ainphechinus rusingeiisis in the presence of an en-

NORTH AMERICAN ERINACEINE HEDGEHOGS

Fig. 13. Parvericius montanus, UMMP V56569, left mandible.
A. Lateral view. B. Occlusal view. C. Lingual view. From the
late Barstovian, Egelhoif Quarry, from an unnamed lithic unit be-
neath the Valentine Formation and overlying the Rosebud Forma-
tion, SW comer of NE Ya, SW )i, sec. 29, T 33N, R 23 W, Keya
Paha Co., Nebraska. X4.5.

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

larged P. Among the other genera of Erinaceinae, only the much
smaller Palaeoscaptor is known to possess as enlarged an P (Tro-
fimov, 1960:38 and Fig. 1).

Parvericius Koerner 1940

Type Species. — Farvericius montanus Koerner 1940.

Known Distribiifion. — ?32 — ?31 my (medial? Oligocene) Asia;
medial Arikareean — late Barstovian (23 — 12.5 my, early to medial
Miocene), North America.

Diagnosis. — Distinguished from Palaeoscaptor and Stenoechimis
by the greater length of the M^ relative to its width; from Palaeo-
scaptor by absence of a metacone on M'^; from all other erinaceines
except Palaeoscaptor and Amphechimis by the presence of an en-
larged Ii; from Stenoechimis by a prominent paraconid taller than
the metaconid on Pj; from Amphechimis, Dimylechimis, Mioechinus,
Palaeoscaptor, and Postpalerinaceiis by the anteroposteriorly com-
pressed Ml trigonid; from Gymmirechimis, Palaeoscaptor, and
Stenoechimis by absence of a talonid or postcingulum on M3; and
from Dimylechimis by presence of M3 and M'^

Parvericius montanus Koerner 1940

Figures 7d, 12-15

Parvericius montanus Koerner 1940:841.

Palaeoerinaceus minimus Bohlin 1942:23.

Amphechinus (Palaeoerinaceus) cf. ?n fnfr/ius Sulimski 1970:64.

Holotype. — YPM 19356, right ma.xilla fragment with P^-M'^ com-
plete except for lingual halves of P''"^.

Type Locality and Stratigraphic Position. — Collected from the
Deep River beds in sec. 3, T 10 N, R 5 E, Meagher Co., Montana.

Diagnosis. — Only known species of genus.

Referred Material. — MPUM 1551, right mandible with highly
worn P4-M;5, tip of Ii root, fragment of Ci, P^ alveolus, horizontal
ramus complete from C, to base of leading edge of the ascending
ramus. Found in the middle Cabbage Patch beds, Bert Creek lo-
cality 2 (MV6504-2), NW }i, NE )i, SW }i, SE K, sec. 1, T 10 N,
R 12 W, Granite Co., Montana (Rasmussen, 1969:131-132).

UCM 29507, isolated left Mi. Found in the Split Rock Forma-
tion, SW M, NW }i, sec. 36, T 29 N, R 90 W, Fremont Co., Wvoming.

UCM 29853, isolated right M,. UCM 29955, left mandible with
Ml-:;, posterior half of rear alveolus of P^, horizontal ramus com-
plete from rear alveolus of P4 to base of leading edge of ascending
ramus. Found in the Split Rock Formation, NE K, NE K, NW }i,
sec. 36, T 29 N, R 90 W, Fremont Co., Wyoming.

F:AM 76704, complete left mandible, edentulous except for
P4. UMMP V56569, left mandible lacking only P. and the angle.
UMMP V57331, left mandible fragment lacking the region pos-

NORTH AMERICAN ERINACEINE HEDGEHOGS

terior to the alveolus for the forward root of Mo with Ii, Ci, P4 and
alveoH for lo, P3, M: and anterior alveolus for Mo. UMMP V61029,
isolated left Mj (this specimen appears to have been etched by
acid). UMMP V61031, isolated right M,. UMMP V61027, isolated
right Mo. UMMP V61033, isolated left P^. UMMP V61034, isolated
right M\ UMMP V61026, isolated left M^. Found at Egelhoff
Quarry in the SW corner of NE K, SW Ji, sec. 29, T 33 N, R 2,3 W,
Keya Paha Co., Nebraska, in an unnamed lithic unit that uncon-
formably overlies the Rosebud Formation and is unconformably
overlain by the Valentine Formation.

UMMP V61022, isolated left Mj. Found at Norden Bridge
Quarry in the SE %, SW 'A, sec. 33, T 33 N, R 23 W, Brown Co.,
Nebraska, in the same lithic unit as Egelhoff Quarry described in
the previous paragraph.

Upper Dentition. — (Fig. 12) P\ — Only the buccal half of this
tooth is preserved on the single specimen axailable. The paracone
is the largest and tallest cusp preser\cd on the tooth. The anterior
side of the paracone is straight and the posterior side, concave.
Wear has blunted the tip of the paracone, but, when unworn, the
apical height of this cusp was probably no more than two-thirds
the crown length. Directed posteriorly from the metacone, the
metacrista abruptly turns in a buccal direction at the posterior
border of the tooth, but no metastyle or other cuspulcs are devel-
oped there. On the anterior side of the paracone base there is a
narrow precingulum with a weak paracrista which extends ventrally

Fig. 14. Parvericius montanus, MPUM 1551, right mandible. A. Lateral
view. B. Occlusal view. From the medial Arikareean, middle Cabbage Patch
beds, Bert Creek locality 2 (MV6504-2), NW %, NE %, SW %, SE %, sec. 1,
T 10 N, R 12 W, Granite Co., Montana (Rasmussen, 1969:131-132). x6.

34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 15. Parvericiiis montaniis, UCM 29955, left mandible. A. Lateral view.
B. Occlusal view. From tlie Hemingfordian, Split Rock Formation, NW %,
sec. 36, T 29 N, R 90 W, Fremont Co., Wyoming. x6.

from the cingulum toward, but not reaching, the apex of the cusp.
Examination of the broken surface where the hngual half of the
tooth broke away suggests that it was markedly shorter anteropos-
teriorly than the buccal half; thus, the metastylar spur projected
away from the body of the tooth as on the P^. On the buccal side
of the tooth along the base of the paracone is a weakly developed
cingulum. Two prominent roots are preserved, one above the
parastylar region, the other over the metacrista. If a third root
were present above the missing lingual region of the tooth, all trace
of it has been lost.

P'. — This tooth is several times larger than the preceding P^.
The height of the paracone, tallest cusp on the tooth, is about five-
sixths of the tooth length. Two-fifths the height of the paracone,
the protocone is lingual and slightly anterior to the cusp, as in living
hedgehogs. Equally as large in basal dimensions, the hypocone is

Table 6. Measurements (mm) of the Upper Dentition of
Parvericius montanus

YPM 13956 UMMP UMMP UMMP

type V61033 V61026 V61034

P^ anteroposterior diameter — _ 1.1

P*, anteroposterior diameter ... 2.1 2.2 — —

P*, transverse diameter — 2.2

M\ anteroposterior diameter .... 2.1 .... 2.0 2.2

M\ transverse diameter 2.5 — 2.4 2.6

M", anteroposterior diameter ... 1.6

M", transverse diameter 1.8 .— . —

M'', anteroposterior diameter -... 0.4 — —

M^ transverse diameter 1.1

NORTH AMERICAN ERINACEINE HEDGEHOGS 35

lower than the protocone and posteroHngual to it. Although these
two lingual cusps are linked together by a low crest, neither cusp
is joined to the buccal structure of the tooth by a crest. Between
the hypocone and the base of the metastylar spur is a planar surface
that slopes posterodorsally. A small parastyle is present on the
well-developed precingulum anterior to the base of the paracone.
Although no paracrista is present, the prominent metacrista is di-
rected posterobuccally away from the paracone and terminates
posteriorly without a distinct metastyle. Likewise, the metastylar
spur projects posterobuccally away from the body of the tooth so
that its lingual half is narrower antcroposteriorly than the buccal
half. Continuous with the precingulum is a weaker cingulum along
the buccal side of the tooth. The three roots of the tooth lie above
the protocone and hypocone, paracone, and metacrista, respectively;
all are subequal in length and diameter.

M^ — Largest tooth in the skull, the M^ is somewhat broader
but subequal in length to the P^. The height of the paracone, third
tallest and third largest of the four principal cusps, is half the
length of the tooth. The paracone is anterior to the metacone and
their bases are confluent. Subequal in height and basal dimensions,
the metacone and protocone are the tallest and largest of the four
principal cusps. The anterior side of the metacone is steep; the
posterior side is more gently dipping. The protocone is lingual and
slightly anterior to the paracone. The hypocone is the lowest and
smallest of the four principal cusps. The hypocone and metacone
are as close to one another as the protocone and paracone. Linking
the anterobuccal corner of the protocone and the anterolingual
corner of the paracone are the well-developed preprotocrista and
preprotoconule crista which grade imperceptibly into each other
in the absence of a paraconule. In anterior \iew this loph has a
broad, inverted V-shaped profile. The postprotocrista is directed
posterobuccally from the protocone. This crista divides at a point
lingual to the anterior edge of the metacone; one branch extends
posterolingually to the hypocone whereas the other, the postmeta-
conule crista, extends buccally to the metacone. No metaconule is
developed on this tooth. A short paracrista crosses the precingulum
and terminates against the base of the paracone at the anterobuccal
corner of the tooth. No parastyle is present. The metacrista extends
posterobuccalK' from the metacone for a distance equal to one-half
the tooth length. A cingulum is developed along the entire margin
of the tooth except at the posterior tip of the metastv'lar spur. In
outline the tooth is approximately square, with prominent excava-
tions on the posterior and lingual side. A well-developed metast\dar
spur projects posterobuccally away from the body of the tooth,
and a much smaller parast\'lar spur projects anteriorly. Three roots
are present — one above the paracone-paracrista region, a second

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

above the metacone-metacrista region, and a third above the proto-
cone-hypocone region.

M~. — Although smaller, the M- is basically similar to the NP.
The protocone and paracone are subequal in height and basal di-
mensions, being the tallest and largest of the four principal cusps.
The lingual region of the M- is the same as the M'; however, the
buccal region, particularly the posterobuccal area, is different. The
metacone has shifted position so that it is posterolingual to the
paracone, not posterior. The metastylar spur projects posteriorly,
not posterobuccally, and is reduced. The buccal border of the
tooth is not subparallel to the midline of the skull, but is directed
posteromedially. Similarly, the parastylar spur has shifted so that
it projects anterobuccally rather than anteriorly. Narrow cingula
are developed on all sides except the lingual. Only the two buccal
roots of the tooth can be observed; relative to structures on the
crown, their positions are the same as on the M^

M^. — This tooth is basically a mediolaterally elongated blade
with a protocone at the lingual end and an e([ually large and tall
paracone at the buccal end; the two cusps are joined together by a
short, straight crest. A weak precingulum is developed along the
anterior margin of this tooth. Two roots are developed on this
tooth — one above the protocone and a second over the paracone.

Lower Dentition. — (Figs. 13-15) /,. — In morphology and size,
this tooth is (juite distinct from the incisor behind (L'). It is elon-
gated, extends forward, and cur\cs slightly upward from the an-
terior end of the mandible for a distance about equal to the Ii;-P?,
length, or 3.7 mm on the least worn specimen (UMMP V56569).
Posteriorly, the root extends beneath and medial to the roots of
L.-P.! to terminate just in front of the anterior root of P4. In cross-
section at the aheolar border, the tooth is a slightly distorted,
dorsoventrally elongated oval with the blunt end dorsal and the
medial side straight rather than curved. The tooth is mediolaterallv
compressed so that its dorsoventral diameter is about half again as
great as its mediolateral diameter. At its alveolar border, the tooth
is nearly as wide and deep as the mandible itself. Further posterior,
beneath the canine, the dorso\'entral diameter of the tooth is equal
to about half the height of the mandible.

I ;. — This tooth and the somewhat larger canine behind are more
similar in form to one another than either is to the I1. Presumably,
the same could be said for the P2, although the tooth is known only
by its alveolus in this species. The single root of the L- dips pos-
teriorly at a shallow angle. Overlapping the Ii ahead, the crown
of the I2 projects forward from the root. Along the midline is a
low crest with two prominent swellings. The anterior swelling lies
slightly forward of the midpoint of the tooth and is the protoconid;
the second swelling lies at the posterior margin of the short talonid.
The length of the tooth is about twice the width.

NORTH AMERICAN ERINACEINE HEDGEHOGS 39

C ,. — In addition to its somewhat larger size, the Ci differs from
the L. primarily in the relati\e sizes of protoconid and talonid cusp,
which are slightly taller relative to the length and width of the
tooth.

Pj. — This tooth is known in this species only by its single al-
veolus. The P2 was probably similar in size and morphology to the
I2, because the alveoli of the two teeth are of similar dimensions;
in other erinaceines, they are similar in morphology. The posterior
dip of the Pj aKeolus is steeper than in L. or Ci.

P;. — This tooth is markedly larger than the three immediately
anterior to it. Tlie protoconid is most prominent of the cusps; its
lieight is nearly as great as the length of the tooth. Anteromedial
to the protoconid is the lower paraconid, a well-developed cusp.
In lateral \iew the paralophid linking the two cusps has a deep,
acute V-shaped profile. The protolophid is directed medially from
the protoconid and has a slight swelling on the lingual end, but no
distinct metaconid. Along the buccal border of the tooth is a weak
cingulum. This cingulum passes into the narrow talonid which is
de\eloped for the full width of th(> trigonid.

On no specimen were the aheoli of this double-rooted tooth
visible.

M,. — The length of the trigonid is four-fifths its width and
approximately equal to half the length of the tooth. Lowest of the
trigonid cusps, the paraconid is anterior to the metaconid; the
bases of the two cusps are separated b>- a deep cleft. The proto-
conid is posterobuccal to the paraconid; the two cusps are linked
by a paralophid. In lateral view, the paralophid has a V-shaped
profile that is acute but nearly a right angle, whereas in occlusal
view, the paralophid is somewhat buccally convex. The protoconid
is the tallest cusp on the tooth and has the greatest basal dimensions.
On the least worn specimen ( UCM 29(S53), the height of th(> proto-
conid is equal to 85 percent of the tooth length. The protolophid
joins the metaconid and protoconid; when viewed posteriorly, it has
an acute, V-shaped profile. The metaconid is intermediate in height
between the protoconid and paraconid.

Ten percent wider than the trigonid, the talonid has onl\- t\\o
cusps, one in each posterior corner. The entoconid is the taller of
these two cusps and is intermediate in height bet^^•een the meta-
conid and paraconid. The entoconid is directly posterior to the
metaconid and the hypoconid is slightly more buccally placed than
the protoconid. In anterior view, the buccal margin of the proto-
conid is nearly straight and that of the hypoconid, convex. Be-
tween the entoconid and that part of the posterior wall of the tri-
gonid beneath the apex of the metaconid is a low entocristid which
encloses the talonid basin lingually. Directed anteriorly and slightly
lingually from th(> hypoconid, the cristid obliqua abuts against the
postvallid immediately behind the protoconid. A continuous, well-

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

developed, narrow cingulum extends along the entke buccal side
of the tooth. At the base of the hypoconid, the buccal cingulum
passes into the posterior cingulum which is directed dorsomedially
so that it terminates lingually midway between the hypoconid and
entoconid just below the crest of the postcristid. In the region of
transition between the buccal and posterior cingula, these narrow
cingula are even narrower.

The partition between the alveoli of this double-rooted tooth
has twinned vertical ridges characteristic of erinaceids. On the
posterior wall of the rear alveolus is a single vertical ridge. The
anterior alveolus is nearly circular with the posterior side slightly
flattened. Its diameter is approximately 0.7 mm. The larger, pos-
terior alveolus is roughly rectangular with an anteroposterior length
of 0.8 mm and width of 0.9 mm.

M2. — Although the Mo is only four-fifths as large as the Mi, in
other respects the teeth are quite similar. Therefore, only differ-
ences between these teeth will be noted in the following two para-
graphs.

The trigonid length is only two-thirds as great as its width. The
paraconid is not developed as a distinct cusp but rather is a small
swelling at the anterolingual end of the paralophid. The antero-
lingual quarter of the protolophid is at an abrupt angle with respect
to the remainder of that crest, being oriented mediolaterally rather
than anterolingually.

The entoconid is more buccally placed on the tooth than the
metaconid. When viewed anteriorly, the buccal margin of the
protoconid is buccally convex. No buccal cingulum is developed
on the most anterior quarter of the prevallid.

Ms. — This tooth differs from the two anterior molars in several
ways: markedly smaller size; complete absence of a talonid; more
anteroposteriorly expanded trigonid, its length being one-fourth
greater than its width; and a single anteroposteriorly elongated root
that may be weakly bifurcated. Only two distinct cusps are present
on the trigonid — the protoconid and metaconid. The protoconid is
taller and somewhat larger in basal dimensions than the metaconid;
the bases of the two cusps are so close to one another that only a
narrow notch separates them. No swelling is present at the lingual
end of the paralophid to mark the presence of a distinct paraconid.
Only the worn base of the paralophid remains on the best available
specimen of this tooth. A well-developed, continuous, narrow basal
cingulum is present along the buccal margin of the tooth, but no
cingulum is present along the posterior side of the trigonid.

Oval in outline, the single alveolus of this tooth departs slightly
from the vertical so as to dip posteriorly. The length of the oval is
0.9 mm and the width, 0.5 mm.

MandihJe. — (Figs. 13-15) Circular in outline, the mental fora-
men Hes below the middle of P^, midway between the dorsal and

NORTH AMERICAN ERINACEINE HEDGEHOGS

ventral borders of the ramus. Beneath P3-M2, the mandible is of a
uniform depth; it gradually decreases in depth posteriorly and
abruptly anteriorly. The angle of the jaw protrudes ventrally and
posteriorly away from the body of the mandible and is slightly in-
flected. The unfused midline symphysis of the mandible extends
posteriorly to a point beneath the C^-P.^ juncture. The dorsal
boundar)' of the symphysis dips posteriorly at an angle of 25° with
respect to the dorsal edge of the mandible ( Z c in Fig. 11). Be-
neath I], the symphysis covers the entire medial surface of the
mandible; posteriorly it covers a progressively decreasing amount.
Parallel and adjacent to the ventral margin of the midline symphysis
is a faint depression which opens posteriorly and may ha\'e been
the site for the insertion of the geniohyoideus muscle.

The ascending ramus lies slightly labial to the midline of the
mandible. Its anteroposterior width, measured from the condyle,
is twice the maximum depth of the jaw. The anterior border of the
ascending ramus departs slightly from the vertical and dips an-

FiG. 16. Method of measuring distances between occluding surfaces. Cor-
responding measurements on the upper and lower teeth are indicated by the
same letter of the alphabet, capital letters for the upper teeth and small letters
for similar measiu-ements on the lower teeth.

42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

teriorly; the angle of elevation ( Z a in Fig. 11) is 80°. Near its
dorsal end, the anterior edge curves slightly posteriorly and dorsally
and passes smoothly into the convex dorsal edge of the ascending
ramus. On the leading edge of the ascending ramus, there is a
sharp lateral flange that forms the anterior margin of the masseteric
fossa. The prominence of this flange decreases dorsally. There is a
ridge on the medial side of the ascending ramus. Beginning in the
anteroventral corner of the ascending ramus immediately behind
the tooth row, the ridge extends posterodorsally for a distance less
than one-fourth the \\'idth of the ascending ramus. To the rear,
the ridge terminates somewhat anterior to the mandibular foramen;
between that point and the condyle, there is no ridge developed on
the medial side of the mandible. The mandibular foramen is mid-
way between the anterior and posterior borders of the ascending
ramus and level with the tooth row. The condyle is well above the
level of the tooth row, the angle of elevation ( Z b in Fig. 11) being
about 35°. In posterior view, the margin of the mandible has a
knife-edge thinness except immediately below the condyle. The
posterior border of the ascending ramus curves anterodorsally from
the condyle and meets the dorsal edge of the ascending ramus at a
sharp angle.

Comvariso7is. — In a living erinaceine such as Erinacens etiro-
paeiis, the distances between adjacent occlusal surfaces of the
upDcr dentition arc found to be close to those of the corresponding
occlusal surfaces of the lower dentition. For example, the distance
from the prevallum of M^ to that of M- is nearly the same as that
from the postvallid of M, to that of M^. Because the prevallum of
M^ is adjacent to the postvallid of Mi when the two teeth are in
occlusion and likewise, the prevallum of M- is adjacent to the post-
vallid of M- in the same circumstances; this result is hardly sur-
prising. See figure 16 for a diagramatic exulanation and table 8
for a set of measurements in a sample of five recent specimens of
E. europaeus.

If the distances between the orevalla of the upper molars of
Parvericius montamis (YPM 13956, Fig. 12a) are compared to the

Table 8. Measurements (mm) of the distances between occlusal planes in
Erinaceus europaeus (See Fig. 16 and text for explanation)

AMNH(M)

AMNII(M)

34820

34821

149412

160470

164391

8.9

9.2

8.4

9.1

8.8

8.7

9.1

8.6

4.6

4.9

4.4

4.6

4.5

4.6

4.8

4.7

4.6

4.5

4.3

4.0

4.5

4.3

4.1

4.3

3.9

4.0

4.1

NORTH AMERICAN ERINACEINE HEDGEHOGS

o
c

Co"
3

^
<»

O
U

■S3

■ — I >-
ca o

^ bO

pa
<

l>D

09 CO

•<?*

►c

u
^

S ^

a „

■*- OJ

CD
lO

s
o

=o CO

S >— 1

2 >^

O
00

^ 2

c:5

C<l

C30

q
CO

CM — ; I-;
CM cm' ^

05 ^ U

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

distances behveen the postvallids of the mandible fragment pre-
sumed to be that of P. montamis from the SpHt Rock Formation
(UCM 29955, Fig. 5), the corresponding measurements are found
to be similar to the degree found between indi\iduals of Erinaceus
europaeus of slightly different size (compare Tables 8 and 9). In
addition, these upper and lower dentitions have been modified in
the characteristic crinaceine pattern; M''' reduced to a straight blade
between the protocone and paracone; Mo diminished to a trigonid
lacking a talonid. These specializations of the most posterior upper
and lower molars are functionally correlated with one another and
are not known to occur singly.

On the basis of the above evidence only, it could be postulated
that the mandibles of Sienoechimis tantalus were the previously
unknown lower jaws of Parvericius montanns. The distances be-
tween the corresponding occlusal surfaces are nearly the same
(Table 9); and, although the M- of S. tantalus does have a wide
postcingulum, it conceivably did occlude with an M*^ similar to that
on P. montamis. However, the M^ and M- referred to S. tantalus
(found at the same locality as the majority of known lower denti-
tions of that species) aopears to be c|uite unlike those of P. mon-
tanus; thus, the possibility of taxonomic synonymy seems improb-
able. The material from Egelhoff Quarry further supports the
assignment of the lower dentions, here inferred to belong to P.
monfanus, with the upper dentition on which the species is based.
At that locality only one form of small crinaceine tooth is known
for each position of the dental series represented, suggesting that
all such teeth belong to a single sriecies. The isolated P^ and tsvo
M^'s from Egelhoff Quarry are quite similar to those of the type of
P. montamis, whereas the lower teeth from there are 10 percent
larger than those of the Split Rock mandible.

Dentitionally, specimens of Parvericius montanus from Taben-
buluk (originally described as Palaeoerinaceus inini7nus Bohlin,
1942), and from Nareen Bulak (originally described by Sulimski,
1970, as Amphechinus {Palaeoerinaceus) cf. mini7nus), are similar
to specimens of P. montamis from North America. On the basis of
dimensions of the teeth, this Asian material is most similar to older,
smaller Arikareean and Hemingfordian North American specimens
of P. montamis. There is a marked difference in the depth of the
mandible; the North American specimens are about half again as
deep as the Asiatic ones (Bohlin, 1942: Fig. 3d, d', Plate 1, Fig. 4;
Sulimski, 1970: Fig. 2d-f; Plate 19, Figs. 5-7). Besides P. minimus,
Bohlin described another species in 1942 from Taben-buluk, Pa-
laeoerinaceus kansuensis Bohlin (1942). This species is based on
an incomplete edentulous mandible fragment. The mandibular
depth is closer to the North American P. montanus than P. minimus,
being only one-fifth larger. Furthermore, based on the alveoli, the

NORTH AMERICAN ERINACEINE HEDGEHOGS 45

molars appear to have been about as large as the later, larger Bar-
stovian North American specimens of P. montanus.

Comparisons with Stenoechinus tantalus were made on p. 18.

Direct comparison of the type and referred specimen of Amph-
echintis horncloudi (SDSM 62113, and 6278; J. R. Macdonald, 1970:
Fig. 5) together with the descriptions and figures of Amphechinus
edwardsi in Viret (1938: Figs. 4, 8, and Plate I, Fig. 2) and Hiirze-
ler (1944: Figs. 7, 10, 13) and of AmpJwcJiimis arvernensis in Viret
(1929: Plate 28, Figs. 1-2), reveals two significant differences be-
tween these three species and Parvericiiis montanus. The corre-
sponding dimensions of the teeth are only about one-half to three-
fifths as large, and the trigonid of Mi is anteroposteriorly compressed
on Parvericius montanus.

Erinaceinae genus and species indet.
Figure 17

Material— KU 18396, isolated right M-. Found in the middle
Cabbage Patch beds. Cabbage Patch localitv 4 (KU-i\It-12), SW }i,
SE }i, SW .'i, SE ii, sec. 1, T 10 N, R 12 W, Granite Co., Montana
(=MV6504-4, Rasmussen, 1969:131-132).

M^. — (Fig. 17) Length of the tooth is only two-thirds the width
(length = 1.5 mm, width r= 2.4 mm). Few signs of wear mark the
surface of the tooth. Tallest of the four principal cusps, the para-
cone is anterobuccal to the metacone and their bases are confluent.
Second tallest of the cusps, the metacone has more gently sloping
anterior and posterior sides than the paracone. Nearly equal in
height to the metacone, the protocone is approximately lingual to
the paracone. The base of the protocone projects medially beyond
the body of the tooth. Posterolingual to the protocone is the lowest
of the four principal cusps, the h\pocone. The base of the hypocone
projects posterolingually away from the body of the tooth. The
hypocone and metacone are nearly as far apart as the protocone

Fig. 17. Erinaceinae, genus and species indet., KU 18396, right M", occlusal
view. From tlie medial Arikareean, middle Cabbage Patch beds, Cabbage
Patch locality 4 (KU-Mt-12), SW }l SE 'i, SW 'i, SE K, sec. 1, T 10 N, R 12 W,
Granite Co., Montana (=MV6504-4, Rasmussen, 1969:131-132). Xl5.

46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

and paracone. Linking the anterobuccal corner of the protocone
and the paracrista are the well-developed preprotocrista and pre-
protoconule crista which grade imperceptibly into one another, in
the absence of a paraconule. In anterior view, this loph has a broad
inverted V-shaped profile. The postprotocrista is directed postero-
buccally from the protocone and terminates posteriorly lingual to
the metacone. A short crest is directed from the posterior tip of
the postprotocrista posterolingually toward the hypocone. How-
ever, the length of the crest is only about one-third of that necessary
to link the postprotocrista to the hypocone; therefore, the cusp is
isolated. A similar gap exists between the postprotocrista and the
metacone because the postmetaconule crista extends buccally from
the cusp but does not reach the crest. No metaconule is developed
on this tooth. Along the straight anterior margin is a well-developed
precingulum that buccally becomes progressively wider and curves
ventrally until it merges with the paracrista buccal to the paracone.
Together the precingulum and paracrista extend further buccally
to the margin of the prominent parastylar spur. No distinct para-
style is developed. From the metacone, the metacrista extends
posterobuccally for about two-thirds of its length and then turns
to an anterobuccal direction to terminate finally without develop-
ment of a distinct metastyle. A well-developed ectocingulum is
present between the paracrista and metacrista. Along the posterior
margin of the base of the metacone is a similarly well-developed
postcingulum.

Comparisons. — This specimen differs from the upper molars of
all other erinaceids possessing a distinct metacrista in that the
buccal third of that crest is directed anterobuccally rather than
posterobuccally.

In general form, this tooth is quite similar to the M- of Valueo-
scaptor acridens and Parvericius moniamis, although it is about 10
to 15 percent larger in every dimension and possesses a more
prominent parastylar spur. In contrast to P. acridens, the postproto-
crista and postmetaconule crista are not parts of one continuous
crest, but instead form two short crests quite distinct from one
another. Unfortunately, the M- of the only specimen of P. moniamis
is too worn to discern the condition of these crests in that species.

The M-'s of Amphechimis, Nenrog,ymnurus, Mioechimis, Post-
palerinaceus, and the modern Erinaceinae are markedly longer
relative to their respective widths and larger in size than this speci-
men. With the exception of Postpalerinaceiis and some specimens of
Ajnp] I echinus, this group also differs from this isolated M- in that
the base of the hypocone does not project away from the main body
of the tooth.

NORTH AMERICAN ERINACEINE HEDGEHOGS

133J Ni anvos

48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

STRATIGRAPHIC RELATIONSHIPS

Stratigraphic positions of the erinaceine insectivores discussed
in this paper from Granite and Powell Counties, Montana, are
illustrated in figure 18. Cabbage Patch beds shown in this figure
range in age from medial to late Arikareean (late Oligocene and
early Miocene), as based on correlations of the enclosed Cabbage
Patch faunas to Arikareean faunas in the John Day of Oregon and
Great Plains of South Dakota, Nebraska, and Wyoming. Cabbage
Patch localities 2 (KU-Mt-9) and 4 (KU-Mt-12) and Bert Creek
locality 2 (MV6504-2) occur within a single measured section.
Cabbage Patch locality 13 (KU-Mt-46) occurs not far from this
measured section and can be directly correlated into it. Because
of poor exposures, erosional gaps, and distances up to several
kilometers, correlation of Cabbage Patch locality 3 (KU-Mt-11) and
Tavenner Ranch locality 2 (KU-Mt-21) to the measured section dis-
cussed above is difficult, but each can be approximately correlated
using the stratigraphic position of the range zone of the gopher
Pleurolicus. Thus, each locality is correlated by recording whether
it occurs above, within, or below the FleuroUcus range zone. Like-
wise, an approximate correlation of this biostratigraphic zone can
be made to the Pleurolicus range zone in the John Day Formation
of Oregon (Rensberger, 1971:141, Fig. 69).

Ten of the eleven specimens of Stenoechinus tantalus occur at
a single horizon and locality (Tavenner Ranch locality 2) in the
upper Cabbage Patch beds above the Pleurolicus range zone. Al-
though several rich fossil localities above and below this strati-
graphic position of Tavenner Ranch locality 2 have been extensively
searched and large bulk samples have been washed from some of
them, only one other specimen of Stenoechimis has been found (at
Cabbage Patch locality 13 which lies within the Pleurolicus range
zone in the middle Cabbage Patch beds). Also occurring at Tav-
enner Ranch 2 is a partial maxillary (KU 18097) and partial isolated
tooth (KU 18405) of Amphechinus horncloudi. Other mammals
found at this locality include Nanodelphys, Peratherium, Domnina,
Proscalops, Meniscomys, Niglarodon, Paciculus, Leidymys, Plesios-
minthus, Gregorymys, Mookomys, Paleocastor, Archaeolagus, Notho-
cyon, Leptomeryx, and oreodonts, and will be reported on in a
later paper by Rasmussen.

Two additional specimens of Amphechinus horncloudi have
been found within the Pleurolicus range zone in the middle Cabbage
Patch beds (Fig. 18)— KU 18162 from Cabbage Patch locality 3,
and KU 18163 from Cabbage Patch locality 2. Each was found
associated with a fauna that is less varied but similar to that ac-
companying A. horncloudi at Tavenner Ranch locality 2, except
that Pleurolicus is present and Gregorymys is absent.

An upper second molar of a small erinaceine (KU 18346) was

NORTH AMERICAN ERINACEINE HEDGEHOGS 49

found at Cabbage Patch locality 4 within the Pleuroliciis range zone
in the middle Cabbage Patch beds. Although more than one ton
of matrix has been washed from this locality and several hundred
specimens have been recovered, this is the sole record of a hedge-
hog in the fauna other than the genus Ocajilo. Mammals from the
locality include Peratlierhnu, a mole, Ocajila, Domnina, Allomys,
Nigkirodon, PleiiwUciis, Mookomys, a beaver, Eutypomys, Archae-
olagiis, an ochotonid, and Diceratherium.

A lower jaw of a small hedgehog (MPUM 1551) referred to
Parvericius montanus was found in the middle Cabbage Patch beds
in association with Perather'mm^ a bca\er, Pleurolicus, and Archaeo-
lagm at Bert Creek locality 2. The Pletiwliais specimens collected
at this locality are among the most primitive in the Cabbage Patch
beds, indicating this locality is near the base of the Pleurolicus
range zone.

Several specimens of StenoecJiinus have been found at Tavenner
Ranch locality 2 in the Cabbage Patch beds and were not found
at other localities stratigraphically above and below (except the
single specimen from Cabbage Patch locality 13). Even at localities
which were intensively sampled, the absence of Stenoechinus may
be due to paleoecological differences among the \arious localities.
The type locality of Stenoechinus (Tavenner Ranch locality 2) is
unif|uely rich in terrestrial gastropods, whereas all other localities
where hedgehogs were found (including Cabbage Patch locality 1.3)
have very numerous freshwater gastropods. The scarcity of fresh-
water gastropods and other aquatic organisms and the abundance
of terrestrial animals at Tavenner Ranch locality 2 suggests that the
area was relati\ely remote from aquatic en\ironments during most
of the episode of deposition of this particular b(>d. The sediment
at Tavenner Ranch locality 2 is composed almost entirely of silt-
sized glass shards with little clay and sand, and may have been
partially wind-blown into the site of deposition. Root burrows are
extremely common, indicating abundant vegetation. A grassy or
wooded habitat in an area free from frequent flooding or standing
water is indicated. Thus Stenoechinus may ha\'e li\'ed in habitats
which were drier than those indicated at the other localities where
other types of hedgehogs have been found.

Two isolated lower molars and a mandibular fragment of
Parvericius nwntonus were found in Fremont Co., east central
Wyoming, in the upper porous sandstone sequence of the Split
Rock Formation in association with mammals of the Split Rock local
fauna considered to be of Hemingfordian age, medial Miocene
(Love, 1961:24; Munthe and Lander, 1973:497). All these speci-
mens were found at or near locality' 11-V of Love (1970).

Two mandibles, three isolated lower teeth, and three isolated
upper teeth of Parvericius montanus were found at Egelhoff Quarry
in Keya Paha Co., north-central Nebraska, in an unnamed lithic

50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

unit that unconformably overlies the Rosebud Formation and in
turn is unconformably overlain by the Valentine Formation (R. H.
Tedford, pers. com., 1972). In the same lithic unit and about 2 km
southeast of Egelhoff Quarry, is the Norden Bridge Quarry in
Brown Co., Nebraska, where an isolated Mi of P. monfomis was
found. Both local faunas are regarded as Barstovian in age (C. W.
Hibbard, pers. com., 1972).

The type of Parvericius monfanus Koerner (1940), a maxilla
fragment, was found in Meagher Co., central Montana, in the Deep
River beds. Unfortunately, Koerner never published a complete
faunal list for the Deep River beds. However, the oreodont material
he collected and described (Koerner, 1940), together with his ro-
dent and lagomorph specimens described by Black ( 1961 ) , indicate
that some parts of the unit (as Koerner conceived it) are of Arik-
areean age and other parts Hemingfordian. In other areas of North
America, Promerycochoenis and Pacictihis are known from Arik-
areean deposits while Hypolaa^us and MonosanJax are known from
Hemingfordian or later deposits (Schultz and Falkenbach, 1949;
Black, 1961; Dawson, 1958; R. W. Wilson, 1960).

CONCLUSIONS

In the Arikareean (latest Oligocene and earliest Miocene), a
general faunal interchange of mammals, particularly small forms,
took place between the Old World and North America after an
extended interval of almost no interchange during the medial and
late Oligocene (R. W. Wilson, 196S). It is at that time, and pre-
sumably as part of that episode of widespread interchange, that
the North American history of the Erinaceinae appears to have
begun with an invasion from Asia by at least two species. These
are documented by the presence of two erinaceine species in Arik-
areean deposits of North America — Amphechinus horncloudi and
Parvericius montanus. Each species appears to be more closely
related to known Oligocene species of Europe or Asia than to any
other North American erinaceines. The Arikareean appearance of
Stenoechiniis tantalus n. gen. and n. sp. in North America may be
the result of yet another invasion immediately prior to the Arik-
areean. Or it is equally as likely that S. tantalus could represent
the first record of a lineage endemic to North America through the
Oligocene. Because no earlier species in either the Western or
Eastern Hemisphere closely resembles S. tantalus, it is not possible
at the present time to strongly support one alternative over the
other. A poorly known species described here as "Erinaceinae,
genus and species indet." may be congeneric with one of the three
other species.

Largest of the North American erinaceine species, Amphechinus
horncloudi can be readily allied with the early Oligocene to early

NORTH AMERICAN ERINACEINE HEDGEHOGS 51

Miocene European group Amphechinus cayluxi, Amphechinus ar-
vernet%sis, and Amphechinus edwardsi. The three European species
and A. horncloudi are all quite similar to one another in size and
morphology; apparently little change took place in this group
during the time of its existence.

Except for a somewhat deeper mandible, Parvericiiis montanus
from North America is nearly identical to medial or late Oligocene
east Asian specimens originally described by Bohlin (1942) as
Falaeoerinaceus minimus. Parvericius would seem to be most simi-
lar and closely allied to Amphechinus among the known erinaceine
genera but more primitixe in the shorter length of the NP and M-
relativ e to their respective widths and in the more anteroposteriorly
compressed Mi trigonid.

The M- described as "Erinaceinae, genus and species indet."
appears to be yet another distinct species, because it is too large
to be the M- of either Parvericius montonus or Sienoechinus tanta-
his and much too small to be the M- of A)npJ)ec]}inus horncloudi.
In addition, it is distinguished from the M- of PoJeoscaptor acridens
in having a more prominent parastylar spur, and the postprotocrista
and postmetaconule crista developed as two distinct ridges rather
than as one continuous ridge.

One of the three erinaceine genera described here has a post-
Arikareean record in North America and a second one may. Par-
vericius surxived until the late Barsto\ian; this genus has been
found in the samples of that age collected at Egelhoff and Norden
Bridge Quarries in north-central Nebraska. Rich will describe these
specimens in a future paper. John Storer is describing three molar
talonids of a large erinaceid of the proper size to be Amvhechinus,
collected at the medial Barstoxian Kleinfeld(T Farm localit\' in
southern Saskatchewan. Although none of the genera discussed
here endured in North America after the Barstovian, the Erinaceinae
were represented on this continent until at least Clarendonian time
as attested by the presence of a single, isolated lower molar belong-
ing to a member of this subfamilv in the WaKeeney local fauna of
Kansas (see R. L. Wilson, 1968: Fig. 12a-b).

SUMMARY

The history of the Erinaceinae in North America begins in the
Arikareean (latest Oligocene and earliest Miocene) with the ap-
pearance of four species: Amphechinus horncloudi, Parvericius
montanus, Stenoechinus tantahts n. gen. and n. sp., and Erinaceinae,
genus and species indet. The first two species listed are more closely
related to various species known from Oligocene deposits of Europe
and Asia than to any North American ones; hence, at least t\vo
separate erinaceine groups appear to ha\e inxaded North America.
Presumably these invasions took place immediately prior to the

52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

first record of these species in North America and were part of an
episode of renewed, widespread interchange of mammals between
North America and the Old World that occurred at that time ( R. W.
Wilson, 1968). StenoecJiintis- tantalus is not closely related to any
known earlier forms; therefore, the Arikareean record of this species
may be either the first discovery of a lineage whose unknown Oligo-
cene history was in North America, or the descendant of yet another
group that invaded North America from Asia in the Arikareean. The
fourth, poorly known species may have been derived from one of
the stocks that gave rise to one of the first three mentioned sDecies,
or it may represent yet another lineage with an independent history
extending well back into the Oligocene.

LITERATURE CITED

Aymard, Auguste

1850. (No title.) Ann. SocAgric. Pay, 14:104-114.
Berggren, W. a.

1971. Neogene chronostratigraphy, planktonic foraminiferal zonation and
the radiometric time scale. Hungar. Geol. Soc. Bull., 101:162-169,
.3 tables.
Black, C. C.

1961. Rodents and lagomoiphs fr(jm the Miocene Fort Logan and Deep
River Formations of Montana. Postilla, (48): 1-20, 6 figs.
DE Blaixville, H. M. D.

18.38. Recherches sur I'anciennete des mammiferes insectivores a la surface
de la terre; precedees de I'historie de la science a ce sujet, des
l^rincipes de leur classification et de leur distribution geographique
actuelle. C. R. Acad. Sci. Paris, 6:738-744.

BOHLIN, B.

1942. The fossil mammals from the Tertiary deposit of Taben-buluk,
western Kansu. Part I: Insectivora and Lagomorpha. In Hedin,
Sven, Reports from tlie scientific expedition to tlie northwestern
provinces of China. Stockholm. Vertebrate Paleontology. 6 (3):
1-113, 32 figs., 1 pi.
Butler, P. M.

1948. On the evolution of the skull and teeth in the Erinaceidae, with
special reference to fossil material in the British Museum. Proc.
Zool. Soc. London, 118:446-500, 28 figs.
1956. Erinaceidae from the Miocene of East Africa. Brit. Mus. Nat.

Hist., Fossil Mamm. Afr., (11): 1-75, 18 figs., 4 pis., 16 tables.
1969. Insectivores and bats from the Miocene of East Africa: New ma-
terial. In Leakey, L. S. B., ed.. Fossil Vertebrates of Africa. Aca-
demic Press, New York and London. 1:1-37, 13 figs.
Dawson, M. R.

1958. Later Tertiary Leporidae of North America. Uni\'. Kansas Paleont.
Contrib., Vertebrata, (art. 6): 1-75, 39 figs., 2 pis., 14 tables.
FiLHOL, Henri

1879. fitude des mammiferes fossiles de Saint Gerand le Puy (Allier).
Ann. Sci. geol. (Paris), 10:1-253, 30 pis.
Friant, M.

1961. Les insectivores de la famille des Erinaceidae. L'evolution de leurs
molaries au cours des temps geologiques. Ann. Soc. Geol. Nord.
81:17-30, 6 figs., 2 pis.

NORTH AMERICAN ERINACEINE HEDGEHOGS 53

HiJRZELER, J.

1944. tjber einem dimyloiden Erinaceiden { Dimijlechinus nov. gen.) aus
dem Aquitanien der Limagne. Eclog. Geol. Helvetiae. 37 (2):460-
467, 13 figs.

KOERNER, H. E.

1940. The geology and xertebiate paleontology of the Fort Logan and
Deep River formations of Montana. Part I: New vertebrates.
Amer. Jour. Sci. 238 ( 12) : 837-862, 7 pis.
Love, J. D.

1961. Split Rock Fonnation (Miocene) and Moonstone Fonnation (Plio-
cene) in central Wyoming. U. S. Geol. Surv. Bull. ( 1121-1): 1-39,
6 figs., 3 tables.
1970. Cenozoic geology of the Granite Mountains area, central Wyoming.
U. S. Geol. Surv. Prof. Paper (495-C): viii+154 pp., 61 figs., 13
tables.
Macdoxald, J. R.

1970. Review of the Miocene Wounded Knee faunas of southwestern
South Dakota. Bull. Los Angeles Co. Mus. Nat. Hist. (8):l-82,
32 figs., 53 tables, 2 maps.

Macdonald, L. J.

1972. Monroe Creek (early Miocene) microfossils from the Wounded
Knee area, South Dakota. South Dakota Geol. Surv. Rept. Invests.
105: iii+43 pp., 14 figs.

Matthew, W. D., and Walter Granger

1924. New insecti\ores and nmiinants from the Tertiary of Mongolia,

with remarks on tire correlation. Amer. Mus. Novitates (105): 1-7,

3 figs.
Mellet, J. S.

1968. The Oligocene Hsanda Gol Formation, Mongolia: a revised faunal
list. Amer. Mus. Novitates (2318): 1-16, 4 figs., 2 tables.

MuNTHE, Jexs, Jr. and E. Bruce Lander

1973. A reevaluation of the age of the Split Rock vertebrate fauna,
Wyoming. Geol. Soc. Amer. Abs. witlr Programs 5(6):497.

Rasmussen, D. L.

1969. Late Cenozoic geology of the Cabbage Patch area, Granite and
Powell Counties, Montana. Univ. Montana unpubl. M.A. thesis:
1-188, 15 figs., 10 plates, 3 tables.

Rensberger, J. M.

1971. Entoptychine pocket gophers (MammaHa, Geomyidae) of the Early
Miocene John Dav Fonnation, Oregon. Uni\-. Calif. Publ. Geol.
Sci. 90:1-209, 76 figs., 22 pis., 15 tables.

Rich, T. H. V. and P. V. Rich

1971. Brachyerix, a Miocene hedgehog from western North .\merica with
a description of the t\inpanic regions of Faraechinus and Podo-
gymnura. Amer. Mus. Novitates (2477): 1-58, 22 figs., 4 tables.
ScHULTZ, C. B. and C. H. F.\lkenbach

1949. Promerycochoerinae, a new subfamily of oreodonts. Bull. Amer.
Mus. Nat. Hist. 93 (art. 3):69-198, 26 figs., 8 tables, 6 charts.
Stole, N. R., et al., eds.

1961. International Code of Zoological Nomenclature. International Tiust
for Zoological Nomenclature. London: x\iii+176 pp.

SULIMSKI, A.

1970. On some Oligocene insectix'ore remains from Mongolia. Palaeont.
Polonica. ( 21 ) :53-70, 2 figs., 2 pis., 6 tables.

Trofimov, B.

1960. [The insecti\ore genus Palaeoscaptor from the Oligocene of Asia.]
Tmdy Pal. Inst. Akad. Nauk SSSR 77:35-40, 3 figs., (Russian).

54 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Van Couvering, J. A.

1972. Radiometric calibration of the European Neogene. In W. W. Bishop
and M. A. Miller, eds., Calibration of Hominoid Evolution. Edin-
burgh, Scottish Academic Press: 247-271, 2 figs., 2 tables.

Van Valen, L.

1967. New Paleocene insectivores and insectivore classification. Bull.
Amer. Mus. Nat. Hist. 135 (art. 5):217-284, 7 figs., 2 pis., 7 tables.

ViRET, J.

1929. Les faunes de mammiferes de I'Oligocene superieur de la Limagne

Bourbonnaise. Ann. Univ. Lyon, (n. s. 1), (fasc. 47): viii+328 pp.,

32 figs., 32 pis.
1938. fitude sur quelques Erinaceides fossiles specialement sur le genre

Palaerinaceus. Trav. Lab. Geol. Univ. Lyon, (fasc. 34, Mem. 28):

1-32, 12 figs., 1 pi.
Wilson, R. L.

1968. Systematics and faunal analysis of Lower Pliocene vertebrate as-
semblage from Trego County, Kansas. Contrib. Mus. Paleo., Univ.
Mich. 22(7):75-126, 17 figs., 20 tables.

Wilson, R. W.

1960. Early Miocene rodents and insectivores from Northeastern Colo-
rado. Univ. Kansas Paleont. Contrib., Vertebrata, (art. 7):l-92,
131 figs.
1968. Insectivores, rodents, and intercontinental correlation of tlie Mio-
cene. XXIII International Geological Congress 10:19-25, 1 table.
Wood, A. E. and R. L. Koxizeskt

1965. A new eutypomyid rodent from the Arikareean (Miocene) of Mon-
tana. Jour. Paleontology. 39(3):492-496, 2 figs., 2 tables.

University of Kansas Publications
MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
History, beginning with volume 1 in 1946, was discontinued
with volume 20 in 1971. Shorter research papers formerly pub-
lished in the above series are now published as Occasional
Papers, Museum of Natural History. The Miscellaneous Publica-
tions, Museum of Natural History, began with number 1 in 1946.
Longer research papers are published in that series. Monographs
of the Museum of Natural History were initiated in 1970. All
manuscripts are subject to critical review by intra- and extra-
mural specialists, final acceptance is at the discretion of the
publications committee.

Institutional libraries interested in exchanging publications
may obtain the Occasional Papers and Miscellaneous Publica-
tions by addressing the Exchange Librarian, University of Kan-
sas Library, Lawrence, Kansas 66045. Individuals may pur-
chase separate numbers of all series. Prices may be obtained
upon request addressed to Publications Secretary, Museum of
Natural History, University of Kansas, Lawrence, Kansas 66045.

Editor: Linda Trxieb
Managing Editor: William E. Duellmax

PRINTED BY

UNIVERSITY OF KANSAS PRINTING SERVICE

LAWRENCE, KANSAS

657

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