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) By 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 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 10 c a <3 cc O >:» c OS CO S a c 3 s M o a s 3 O) cq s o ^ 00 s q Oh < CD O -^ 00 1—1 o 00 1— ( lO CO 00 rH D CO 00 1—1 i2 -t .c o a o ■^ 00 CO o o 00 1—1 o o 00 o o 00 1—1 q t^ CC lO o oi ,-< .-i ■-; o -< -r n lo Tt; q t-; C. — 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 17 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 19 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 27 a o •rH C O ISO H O O 0) J* '-5 la eq — "S cu p o O ^ -^JS U5 CD -M ■y. o o o c .2 - Oj o 05 C8 O Ol 1 — 1 Q S ■" 3 3 c ■^ (Z5 c4 o CO O CT) CD CT> 05 ^ ■* ^_ CO 02 05 00 00 00 c-i i-i >-( r-i CO 00 00 in t- p -f p 05 CO 1> CO lO CO CO C2 oi O O TT !>- p cq CO OO' CC 05 CO ^' 00 (M' ^ ^ fN (M (D 3 PQ CD in CD in > PL. :^ in in 05 OS (M u CO in 00 05 o in in in ino-iincocqoqppinTfpoq 1— ii— !(>ii— (T— ii— ic— io Ol (M 1H in I- C5 fM_ Ol J> CO oi 1— ! I— I 1— i 1— i o >— ! >— 5 o o cM_ o] cq t-; C>1 T— I ■— ( I— I ^ c^i CO in Oq T— I 1— ^ I— I M 05 C5 1— i o >— i CO CO in O) c CD i—Ji— ii— iO'— ir- ioO s OS .ii c8 C Sjr 2 _^ .X M S -^ -S 2 o a o „ > 4-1 S •5 o "5 c .ii (b - c c *- M M O C tftfs^s;^^;^;^;^;^ NORTH AMERICAN ERINACEINE HEDGEHOGS 37 n3 a •r-t •W a o o 1> >-l to < o I— < > Oh O CO < 1> cq o 1—1 > Oi o 1—1 CO > p-j CO o 1— ( CD > CM CO CO > :^ in CO iq CD O-l 1— i 1— J 1— i lO o q q lO in lo t; q oi 1— < — < 1— I 00 o 1< fM u ■M s .2 4-1 •t-t a c _3} 'C s O o 4-1 rt ■*-> •M 03 4-> ■J-' c o to " - ^ x; *^ =« -^ P^PMi^SSSi^^ 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 41 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) AMNH(M) AMNH(M) AMNH(M) AMNII(M) 34820 34821 149412 160470 164391 A 8.9 9.2 8.4 9.1 8.8 a 8.7 9.1 8.6 8.6 8.6 B 4.6 4.9 4.4 4.6 4.5 b 4.6 4.8 4.7 4.6 4.5 C 4.3 4.3 4.0 4.5 4.3 c 4.1 4.3 3.9 4.0 4.1 NORTH AMERICAN ERINACEINE HEDGEHOGS 43 C o c c CS Co" 3 ^ <» O U CO O ^? ■S3 ■ — I >- ca o CS ^ bO pa < l>D o ^ o c 09 CO • Pi a s o in CM CO in =o CO S >— 1 ^^ 2 >^ O 00 ^ 2 CO c:5 in q O) Cd. 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. 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