AMERICAN MUSEUM

NovlttatesPUBLISHED BYTHE AMERICAN MUSEUMOF NATURAL HISTORYCENTRAL PARK WEST AT 79TH STREETNEW YORK, N.Y. 10024 U.S.A.

NUMBER 2610 DECEMBER 2, 1976

RICHARD H. TEDFORD AND DAVID FRAILEY

Review of Some Carnivora (Mammalia)From the Thomas Farm Local Fauna(Hemingfordian: Gilchrist County, Florida)

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AMERICAN MUSEUMNov itatesPUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORYCENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024Number 2610, pp. 1-9, figs. 1, 2 tables 1, 2 December 2, 1976

Review of Some Carnivora (Mammalia)From the Thomas Farm Local Fauna

(Hemingfordian: Gilchrist County, Florida)

RICHARD H. TEDFORD1 AND DAVID FRAILEY2

ABSTRACT

Species of the carnivore genera Cynelos(Amphicyonidae), Hemicyon (Ursidae), andEuoplocyon (Canidae) are recognized in themedial Hemingfordian Thomas Farm LocalFauna. In addition early Mustelinae are indicatedby the ramus referred to Miomustela(?) by Olsen.The record of the latter and that of Hemicyonand Euoplocyon represent the earliest occur-

rences of each taxon in North America; all fourrecords constitute the first recognized occurrenceof each in eastern North America. The ThomasFarm Cynelos, Hemicyon and musteline are veryclosely related to European representatives ofthese taxa from the early Burdigalian, confirmingprevious correlations of the Burdigalian Age withat least part of the Hemingfordian "Age."

INTRODUCTION

Since its discovery in 1931, the Thomas FarmQuarry has remained the most productive Terti-ary locality in eastern North America and hasyielded the largest sample of Hemingfordianvertebrates from this part of the continent.Various taxa of the local fauna have been de-scribed and complete faunal lists have been pub-lished (Ray, 1957; Olsen, 1962; Patton, 1967)that have established the Thomas Farm LocalFauna as a reference point for Miocene faunalcomparisons between eastern and western NorthAmerica. The recognition of Cynelos (Amphicyo-nidae), Hemicyon (Ursidae), and Euoplocyon(Canidae) in the Thomas Farm Local Fauna con-stitutes a refinement in our knowledge of thiscritical local fauna.

ACKNOWLEDGMENTSWe are grateful to Mr. Henry Galiano, De-

partment of Vertebrate Paleontology, the Ameri-can Museum of Natural History, for discussionsand suggestions during the preparation of thepresent paper. We further express our gratitudeto Drs. Robert M. Hunt, Jr., Department of Geol-ogy, University of Nebraska and Thomas H. Pat-ton, Department of Geology, University ofFlorida for their critical comments on the manu-script of this work. Dr. Farish Jenkins lent usmaterial from the Thomas Farm Collection ofthe Museum of Comparative Zoology, HarvardUniversity. Mr. Raymond Gooris skillfully ren-dered the figures. The manuscript was typed byMs. Janice Ebenstein.

'Chairman, Department of Vertebrate Paleontology, the American Museum of Natural History.2Curatorial Assistant, Department of Vertebrate Paleontology, the American Museum of Natural History.

Copyright © The American Museum of Natural History 1976 ISSN 0003-0082 / Price 90 cents

AMERICAN MUSEUM NOVITATES

INSTITUTIONAL ABBREVIATIONSAMNH, Department of Vertebrate Paleontology,

the American Museum of Natural HistoryF:AM, Frick Collection, the American Museum

of Natural HistoryMCZ, Museum of Comparative Zoology, Harvard

UniversityMNHN, Museum National d'Histoire Naturelle

SYSTEMATICS

FAMILY AMPHICYONIDAE TROUESSART, 1885

SUBFAMILY AMPHICYONINAE TROUESSART, 1885

GENUS CYNELOS JOURDAN, 1862

Cynelos caroniavorus (White, 1942)

Daphaenus caroniavorus White, 1942Parictis bathygenus White, 1947Absonodaphoenus bathygenus (White, 1942),

Olsen, 1958

RevisedDiagnosis. About the size of C. schlos-seri (Dehm, 1950). Ml triangular, crown con-

stricted lingual to paracone and metacone; M2rectangular approximately 3/4 the width of Ml;M3 oval, approximately half the width of M2,with no distinct cusps. P2-M1 separated by dia-stemata approximately equal to half the lengthof P3 (P4-M, diastema may be absent). Talonid ofM2 very reduced with an indistinct hypoconidand no entoconid.

Discussion. The species of D. caroniavorus andP. bathygenus were defined on disparate ele-ments (upper molars and a lower jaw fragmentrespectively), but Olsen (1958) recognized themas belonging to a single taxon for which he in-advertently used the wrong specific name. BothWhite and Olsen indicated that this taxon was

distinct generically and Olsen formally recog-nized this in proposing the generic name Absono-daphoenus for the Thomas Farm material. In hisdiscussion of the relationships of this form, hemade what appears to us to be a propheticremark in a comparison with the type specimen:"The upper molars of A. bathygenus compare

well with the general form and structure of thosefound in Amphicyon eseri from the Miocene ofEurope, but are not closely allied to any knowngenera from North America." Although the name"Amphicyon eseri" has been used for species of

at least three genera of amphicyonids (see Kuss,1965), the type may belong to a species of Cyne-los Jourdan, 1862, the genus to which we believethe Thomas Farm species should be assigned.There seems little doubt that Absonodaphoenusis an amphicyonine rather than an amphicyno-idontine (as Olsen had concluded) because of itssimple, transversely elongate, tribosphenic uppermolars, presence of an M3, strong hypoconid onM1, and large protoconid on M2. Its small sizeinvites comparison with Cynelos and PseudarctosSchlosser, 1899. It lacks the diagnostic wide,simple premolars, short trigonid, and broadtalonid of M1 and M2 of Pseudarctos and agreeswith the more primitive Cynelos in having a long,deep jaw with diastemata shorter than half thelength of the premolars; elongate slender pre-molars with variable development of a posteriorcusp on P4; talonids narrower than trigonids onM1-M2; M2 with short trigonid; M'-M2 relativelyshort and wide, often with concave anterior andposterior borders, in which the protocone formspart of a U-shaped crest with the variable presentconules.

In his study of the genus, Kuss (1965) re-stricted Cynelos to two species groups that rangefrom Aquitanian through Burdigalian time. LaterGinsburg (1966) relegated two Oligocene speciesto the genus, each of which appears to representa member of the two groups present in the earlyMiocene. Cynelos caroniavorus closely resemblesthe C. rugosidens group and particularly the formC. schlosseri (Dehm, 1950), known from Bur-digalian deposits in the Loire Basin in France(Ginsburg, 1974) and limestone fissure fillings atWintershof-West, Bavaria (Dehm, 1950). TheAmerican and European forms resemble oneanother closely in size, in the degree of reductionof M3, the small size of Ml to Mt, the small sizeof P4 in relationship to M1, and the lack of de-fined conules on the upper molars (see data inKuss, 1965). The Thomas Farm species differsvery little from its European relative; in size itfalls within the lower range of values for dimen-sions of the cheek teeth of C. schlosseri, but itslower premolars are relatively smaller and morewidely spaced. Further material is needed beforegeography can be overcome as one of the primeconsiderations in separation of the Americanform.

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TEDFORD AND FRAILEY: CARNIVORA

Hunt (1972) has previously recognized thepresence of Cynelos in North America, specifi-cally the Cynelos lemanensis group, which rangesfrom late Arikareean through the medialHemingfordian. It now appears that speciesclosely allied to the C. rugosidens group werealso present during part of medial Hemingfordiantime in North America.

FAMILY URSIDAE GRAY, 1825

SUBFAMILY HEMICYONINAE FRICK, 1926

GENUS HEMICYON LARTET, 1851

Subgenus Phoberocyon (Ginsburg, 1955)Hemicyon (Phoberocyon) johnhenryi

(White, 1947)Aelurodon johnhenryi White, 1947

Revised Diagnosis. Relatively unreduced, nar-row premolars with well-developed posterioraccessory cusps on P2-P4; maximum width of M1across metaconid equal to or slightly exceededby width of trigonid of M2; narrow talonids onM1 and M2 with large hypoconids and shallowbasins; small paraconid present on M2; proto-conid of M2 higher than metaconid; mandibleslender.

Description. Hemicyon johnhenryi fromThomas Farm was described as a new species ofAelurodon (White, 1947) using a ramus withlower dentition as the type specimen (MCZ4059). No upper teeth have been referred to thisspecies. Although the mandible and dentition ofH. johnhenryi bear a superficial resemblance tothat of Aelurodon, numerous features differ-entiate Hemicyon and H. johnhenryi from Aelu-rodon. These are as follows: (1) The mandible ofHemicyon has a premasseteric fossa not found inAelurodon (or any other nonursid carnivore); (2)the lower premolars are much smaller in relationto M1 than is seen in most Aelurodon; P4,especially, is large and robust in Aelurodon incontrast to Hemicyon; (3) M2 is larger, relative toM1, than in Aelurodon (in Aelurodon the M2 issmaller than P4).A second specimen of Hemicyon johnhenryi,

a left ramus with P1-M2, is housed in the FrickCollection, the American Museum of Natural His-

tory (F:AM 98608, fig. 1). Although unques-tionably of the same species, minor differencesdo exist between this referred specimen and theholotype of H. johnhenryi (MCZ 4059), whichprovide an indication of individual variationwithin this species. Also, P1 is missing in the typespecimen but present in the referred specimenand can be described for the first time, as can therelative placement and sizes of the incisor alveoli.

The lower incisor alveoli are crowded betweenthe large root of the canine and the very rugosesymphyseal suture. They form a triangular con-figuration with the 12 root falling behind those ofI1 and I3. The root of I1 indicates that it wassmaller than the subequal I2TI3. The P1 of H.johnhenryi is relatively large and bladelike inappearance. The highest point of the crown is inthe anterior part of the enamel crest although nocusp is readily apparent. All the teeth of thereferred ramus are smaller and more slender thanthose of the holotype (table 1). The alveolus forM3 is also smaller as was presumably M3. Thetalonid of M1 of the holotype has a small ento-conid, whereas that of the referred specimen hastwo equal-sized but small cusps in the position ofthe entoconid. The protoconid and metaconid ofM2 are lower on the referred specimen than thesecusps on the holotype, although the protoconidis the larger cusp as in the latter. The referredramus appears to be shallower anteriorly than theholotype but this may be due only to breakage inthe holotype. In both specimens the anterioredge of the masseteric fossa is bounded by a lowridge that passes posteroventrally toward theedge of the large muscle scar for the massetermedialis on the angular process. The premasse-teric fossa is shallow and extends anteriorly tobeneath the anterior part of M2, and then for-ward as a shallow groove to the position of themental foramina. The referred ramus of H. john-henryi has two large mental foramina, one belowP2 and one below P4. On the holotype ramusthere are three mental foramina; two smallerforamina beneath P4 in place of the large singleforamen seen on the referred ramus. The condyleof the holotype is much heavier than that of thereferred ramus and the height of the ascendingramus is less. The overall impression of the holo-type is of a relatively heavier, more massive jaw.

Discussion. Hemicyon johnhenryi is the

1976 3

AMERICAN MUSEUM NOVITATES

A

B/I

D

FIG. 1. Hemicyon (Phoberocyon) johnhenryi (White, 1947). Holotype, MCZ 4059, right (reversed,drawn from cast), C, P1 alveolus, P2-P4, M1-M2, M3 alveolus. A. Occlusal view. B. Lateral view. Referredspecimen, F:AM 98608, left ramus, C alveolus, P1-P4, M1-M2, M3 alveolus. C. Occlusal view. D. Lateralview. All X1/2.

earliest occurring species of Hemicyon in NorthAmerica, and, as might be expected, exhibitsnumerous features that may be considered primi-tive for the genus. These serve well to differ-entiate this species from the other North Ameri-can species ofHemicyon whose features are morecharacteristic of the genus, i.e., demonstrate agreater emphasis on a massive, bearlike mandibleand large, crushing M1-M2. The mandible ofHemicyon johnhenryi is relatively slender; themasseteric and premasseteric fossae are theshallowest among those known for species ofHemicyon. The teeth of H. johnhenryi are amongthe largest known for the species of Hemicyonbut the mandible is much less deep. The pre-molar series is not so drastically reduced as it isin the other species nor are diastemata asapparent between them. The talonids of M1 andM2 of H. johnhenryi are much less basined thanin most of the other species as the large hypo-conid occupies most of the talonid. The ento-conid is small and variable in expression. A smallparaconid is retained on M2.

In Europe where the record of Hemicyonextends from the Burdigalian to the Tortonian,species with the suite of morphological char-acters that typify H. johnhenryi are restricted tothe Burdigalian. These species are the sole repre-sentatives of the genus Phoberocyon Ginsburg,1955 [P. aurelianensis (Frick, 1926), and P.huerzeleri Ginsburg, 1955] distinguished by itsauthor from Hemicyon Lartet, 1851, by thefollowing features: P3-P4 with posterior accessorycusps; M1 long and slender, talonid low andinclined forward; M2 of the same width as MI,massive, low, with strong trigonid cusps, includ-ing a strong paraconid, protoconid more elevatedthan metaconid, talonid as in M1. As Ginsburg(1955, p. 89) pointed out, the species ofPhoberocyon show a curious mixture of archaicand specialized features, however they are mosteasily distinguished from the species of Hemi-cyon (and Plithocyon Ginsburg, 1955) by primi-tive features. Consequently, we have chosen toregard Phoberocyon as a plesiomorphous sistergroup to Hemicyon and have therefore reduced

4 NO. 2610

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TEDFORD AND FRAILEY: CARNIVORA

this taxon to the rank of a subgenus to show thisrelationship. We are aware of the fact that a

group so distinguished may be paraphyletic, buta far richer record of these rare ursids is neededfor useful phylogenetic analysis.

Hemicyon (P.) johnhenryi is closest in size toH. (P.) aurelianensis, but differs in that the para-

conid on M2 is smaller and there is no sharpgroove separating it from the metaconid. Un-fortunately, nothing is known of the premolarsof H. (P.) aurelianensis, but the smaller H. (P.)huerzeleri has posterior cusps on P3 and P4 (butnot P2). Hemicyon (P.) huerzeleri has an M2trigonid more nearly resembling H. (P.) john-henryi. Considering the great range in dentalmeasurements shown by individuals of thesexually dimorphic species of ursids, it is possiblethat the two European Phoberocyon species lienear the extremes of variation for a single species(compare the ranges with comparable dimensionsfor single taxa of better represented hemi-cyonines in the data given by Hurzeler, 1944, andGinsburg, 1955). If this is the case then H. (P.)johnhenryi would be weakly distinguished fromthe European forms only by having posterioraccessory cusps on P2 as well as P3-P4 and perhapsshorter diastemata between the premolars.

Hemicyon (Phoberocyon) johnhenryi is pres-ently known only from the Thomas Farmdeposit, but associated metacarpals IV-V (F:AM

68254) from the Hemingfordian Blick LocalFauna (Gawne, 1975, p. 2) of the Zia Sandrecords the presence of a long-footed Hemicyonin New Mexico that is comparable in size withthe limbs expected of H. (P.) johnhenryi.

FAMILY CANIDAE GRAY, 1821

SUBFAMILY BOROPHAGINAE SIMPSON, 1945

GENUS EUOPLOCYON MATTHEW, 1924

Euoplocyon spissidens (White, 1947)

Aelurodon spissidens White, 1947Enhydrocyon spissidens (White), Olsen, 1958

Revised Diagnosis. P4 more slender than thatof E. praedator Matthew, 1924. Talonids of Mland M2 are longer and wider than in E. praedator.The protoconid and metaconid of M2 are sepa-rate, distinct cusps. The lower tooth row of E.spissidens is less curved than that of E. praedator.

Discussion. Euoplocyon spissidens is knownonly from lower dentitions. Previous authorshave primarily used the size and placement of thepremolars, the trenchant talonid, and absence ornear absence of a metaconid on M1 in deter-mining the generic affinities of this species. The

TABLE 1Comparative Measurements of Hemicyon johnhenryi and Hemicyon aurelianensis

(Measurements in Millimeters; parentheses indicate an approximate measurement.)

Hemicyon Hemicyonjohnhenryi aurelianensis

F:AM 98607 F:AM 98608 MNHN 228(Cast of MCZ 4059, holotype) (Holotype)a

Length from canine to condyle 201.0 (198.0)Depth of ramus between P2 and P3 36.6 31.8Depth of ramus between M2 and M3 (42.0) 42.5Depth of condyle at center 17.0 11.1P, (length X width) - 8.1 X 4.2P2 (length X width) 12.4 x 6.5 10.9 X 5.4P3 (length X width) 14.0 x 6.9 12.9 x 6.0P4 (length X width) 18.2 X 9.7 16.8 X 8.0Ml (length X width) 33.6 X 14.3 33.2 X 13.1 35.0 X 13.5M2 (length X width) 22.0 x 15.0 19.7 X 13.0 (21.0 X 13.5)M3 (alv.) (length X width) 15.0 x 6.9 (11.0 X 6.0)

aMeasurements of M, from Hurzeler, 1944, p. 154; M2 from Ginsburg, 1955, fig. 2.

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AMERICAN MUSEUM NOVITATES

problems inherent in working with limited mate-rial are apparent in the taxonomic history of E.spissidens. White (1947) diagnosed the speciesand referred it to Aelurocyon. Olsen (1958),using Enhydrocyon crassidens Matthew, 1907,for comparison, noted the less crowded toothrow and the less robust teeth of Enhydrocyon,also seen in E. spissidens, and the absence of thelingual flexure in the trigonid of M1 of Aeluro-cyon, in contrast to E. spissidens, and transferred"Aelurocyon" spissidens to Enhydrocyon Cope,1879. Olsen (1958) also noted that the smallridge which occupies the position of the meta-conid on Ml was present in Enhydrocyon crassi-dens, E. spissidens, and a referred specimen ofEuoplocyon praedator (MCZ 17301) but feltthat E. spissidens did not otherwise closelyresemble Euoplocyon praedator.' For the follow-ing reasons, we believe that Euoplocyon is themore correct generic assignment for E. spissi-dens: (1) The teeth of Euoplocyon are lessrobust than those of Enhydrocyon; (2) P4 has an

anterior accessory cusp which is lacking inEnhydrocyon; (3) the talonids of Ml and M2 are

larger than those of Enhydrocyon; (4) the hypo-conid of Ml in Euoplocyon is more labiallyplaced and less bladelike, whereas that ofEnhydrocyon occupies most of the talonid; (5)Euoplocyon has a Pl, whereas Enhydrocyonoften lacks this tooth; and (6) the mandible ofEuoplocyon increases in depth between P3 andMI; the opposite being true in Enhydrocyon.

Euoplocyon spissidens differs from E.praedator, the only other described species ofEuoplocyon, in the reasons stated in the diag-nosis. The referred ramus MCZ 7310 is comparedin figure 2 with the type of E. praedator (herefigured for the first time). The most obviousdifferences are the distinct cusps on the trigonidof M2 in E. spissidens. In E. praedator thetrigonid of M2 is composed of a single crest inwhich the protoconid and metaconid cannot bedifferentiated.

The trenchant talonid and the absence of a

metaconid on Ml led Matthew (1924) to relate

'The presence of a ridge in the position of the meta-conid appears to be variable in Euoplocyon. The ridge istotally absent in the type specimen of E. praedator(AMNH 18261) but present as a different ridge in otherspecimens (F:AM 27314, 27315) referred to thatspecies.

Euoplocyon to the Simocyoninae Zittel, 1893.These features, however, are widely encounteredamong carnivores and for this and other reasonsthe Simocyoninae can be shown to be a highlypolyphyletic group whose members can beallocated to the Amphicyonidae, Ursidae, andCanidae. The unusual presence of anterior andposterior cingular cusps and a posterior accessorycusp on all four premolars, used by Matthew(1924) in his diagnosis of E. praedator, are foundonly in the borophagine canids Tomarctus Cope,1873, Prohyaena Schlosser, 1887, and AelurodonLeidy, 1858. As these three genera have a num-ber of features in common (such as a pre-maxillary-frontal bone contact, an enlarged I3with accessory cusps, a parastyle on P4) andprobably form a natural group, Euoplocyoncould perhaps be better placed with these generain the Borophaginae Simpson, 1945.

Euoplocyon is known from two species:Euoplocyon spissidens from the Thomas FarmLocal Fauna (medial Hemingfordian) and E.praedator from the Lower Snake Creek Fauna(early Barstovian). Gazin (1932) referred a singleMl from the Skull Spring Fauna (early Bar-stovian) of Oregon to ?Euoplocyon. This latteridentification remains uncertain, but the genus isnonetheless widespread over North America.

FAMILY MUSTELIDAE SWAINSON, 1835

SUBFAMILY MUSTELINAE SWAINSON, 1835

Miomustela(?) sp. Olsen, 1956

The small mustelid referred to Miomus-tela(?)sp. by Olsen (1956) appears to represent athird carnivore from Thomas Farm belonging toa genus also present in the Old World Miocene.Olsen's queried reference to Miomustela Hall,1930, must be rejected on the grounds of thedistinctly different morphology of the carnassialand the large size of the crowded premolars asindicated by their alveoli. The carnassial has avery high compressed trigonid with a small lowmetaconid and a narrow trenchant talonid. Thepresence of an Ml metaconid differentiates theThomas Farm species from contemporary Pale-ogale Meyer, 1846. Comparable forms are knownfrom the Runningwater Formation of Nebraskaand among the various species assigned to thegenus Plesictis Pomel, 1846, in Aquitanian andBurdigalian faunas of Europe. The latter differ so

NO. 26106

TEDFORD AND FRAILEY: CARNIVORA

A gC Q

B

C CP

D

FIG. 2. Euoplocyon praedator Matthew, 1924. Holotype, AMNH 18261, left ramus, C broken, P1,P2 broken, P3-P4, M1-M3. A. Occlusal view. B. Lateral view. E. spissidens (White, 1947), referred, MCZ7310, left ramus, P2-P3 alveoli, P4, M1-M2. C. Occlusal view. D. Lateral view. All natural size.

TABLE 2Comparative Measurements of Euoplocyon spissidens and Euoplocyon praedator

(Measurements are in Millimeters; parentheses indicate an approximate measurement.)

Euoplocyon Euoplocyonspissidens praedator

MCZ 4246a MCZ 7310 AMNH 18261Type Type

Depth of ramus at P3 19.0 19.4 21.7Depth of ramus at M2 21.0 22.5 22.9P, (length X width) - - 5.9 X 3.7P2 (length X width) length alveolus (6)P3 (length X width) - Length alveolus (8) 8.4 X 4.8P4 (length X width) 10.0 X 5.0 10.0 X 5.2 10.0 X 5.7M1 (length X width) 17.0 X 7.0 17.0 X 7.0 18.2 X 7.7M2 (length X width) 7.8 x 5.1 6.8 x 5.3M3 (length X width) 3.4 X 3.2

aMeasurements from White, 1947, p. 498.

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AMERICAN MUSEUM NOVITATES

radically from the holotype of the genotypic spe-cies (P. genettoides Pomel, 1846) that referenceto that genus seems as unlikely as reference toMiomustela. Hough (1948) and de Beaumont(1968) have shown that P. genettoides, and theclosely related P. lemanensis Pomel, 1853, areprocyonids although musteloid in dental adapta-tion. The Thomas Farm form is closest to "P."julieni Viret, 1929, and "P." sicaulensis Viret,1929, in size (see Dehm, 1950, fig. 78) andmorphology. "Plesictis" julieni is known from askull (the type, Viret, 1929) which shows theinflated bullae with strongly forward directedmeati, and widely separated paroccipital andmastoid processes typical of mustelids and con-trasting with contemporary Plesictis. Thesefeatures and the parasagittal crests are matchedin specimens from the Runningwater Formation.Mustelids such as these have many of the featuresof the later Cenozoic and living Mustelinae, andpossibly represent the earliest occurrences of thatsubfamily in the New World.

SUMMARY

The presence of the Holarctic carnivore generaCynelos and Hemicyon in the Thomas FarmLocal Fauna and their very close morphologicalcorrespondence with European counterpartsyields further data on the intercontinental corre-lation of these early Miocene mammal faunas. Asthe comparisons detailed above reveal, Cyneloscaroniavorus and Hemicyon (Phoberocyon)john-henryi closely resemble the exclusively Burdi-galian C. schlosseri and H. (P.) aurelianensisrespectively. These European species occurtogether in the Sables de l'Orleanais of the LoireBasin of France. Cynelos schlosseri (type) and H.(P.) huerzeleri [closely related to H. (P.) aureli-anensis] occur in the fissure fills at Wintershof-West in Bavaria. Both assemblages are regardedas early Burdigalian Age following the conceptof that Age held by European paleomammalogists.

Euoplocyon is recognized in the ThomasFarm Local Fauna extending both the temporaland geographic ranges of this genus from earlyBarstovian faunas in Oregon and Nebraska intothe Hemingfordian of Florida.

The identification of Miomustela(?)sp. byOlsen (1956) is rejected. The mustelid in ques-tion is apparently a true musteline related to spe-

cies that are now incorrectly included within theprocyonid genus Plesictis. Such musteline speciesare known from Aquitanian and Burdigalianfaunas of Europe and Hemingfordian faunas ofNorth America.

LITERATURE CITED

de Beaumont, G.1968. Note sur la region auditive de quelques

carnivores. Arch. Sci. Geneve, vol. 21,no. 2, pp. 213-224.

Dehm, R,1950.

Frick, C.

Die Raubtiere aus dem Mittel-Miocan(Burdigalium) von Wintershof-West beiEichstatt in Bayem. Abh. BayerischenAkad. Wiss. Math.-Nat. KI., vol. 58, pp.1-141.

1926. The Hemicyoninae and an AmericanTertiary bear. Bull. Amer. Mus. Nat.Hist., vol. 56, art. 1, pp. 1-119.

Gawne, C. E.1975. Rodents from the Zia Sand Miocene of

New Mexico. Amer. Mus. Novitates, no.2586, pp. 1-25.

Gazin, C. L.1932. A Miocene mammalian fauna from

southeastern Oregon. Publ. CarnegieInst. Washington, no. 418, part III, pp.37-86.

Ginsburg, L.1955. De la subdivision du genre Hemicyon

Lartet (carnassier du Miocene). Bull.Soc. Geol. France, 6th series, vol. 5, pp.85-99.

1966. Les amphicyons des Phosphorites duQuercy. Ann. Paleont., vol. 52, pp.23-63.

1974. Les faunes de mammiferes burdigalienset Vindoboniens des bassins de la Loireet de la Garonne. Mem. Bur. du Rech.Geol. et Min., vol. 1, no. 78, pp.153-167.

Hough, J. R.1948. The auditory region in some members

of the Procyonidae, Canidae and Ursi-dae. Bull. Amer. Mus. Nat. Hist., vol.92, art. 2, pp. 67-118.

Hunt, R. M., Jr.1972. Miocene amphicyonids (Mammalia,

Carnivora) from the Agate SpringsQuarries, Sioux County, Nebraska.Amer. Mus. Novitates, no. 2506, pp.1-39.

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TEDFORD AND FRAILEY: CARNIVORA

Hurzeler, J.1944. Zur Revision der europfiischen Hemi-

cyoniden. Verhandl. Naturf. Gesell.Basel, vol. 55, pp. 131-157.

Kuss, S. E.1965. Revision der europaischen Amphi-

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5-168.Lartet, E.1851. Notice sur la Colline de Sansan, J.-A.

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