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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3092, 34 pp., 9 figures, 1 table February 17, 1994 Basicranial Anatomy and Phylogeny of Prinmitive Canids and Closely Related Miacids (Carnivora: Mammalia) XIAOMING WANG' AND RICHARD H. TEDFORD2 ABSTRACT Selected fossil carnivorans are studied in an at- internal carotid artery in an extrabullar position tempt to bridge the gap between caniform miacids between the entotympanic and the petrosal. Early and early canids, and to identify character trans- canids are further distinguished by their posses- formations within the clade leading to canids. Sev- sion of a shallow suprameatal fossa. Canids suc- eral basicranial and dental features are important cessively acquired the following dental characters: in characterizing the cladogenetic events that oc- presence ofposterior accessory cusps on upper and curred and culminated in Hesperocyon. lower third premolars, loss of M3, reduction of Basicranial transformations from miacids to the the parastyle on M1, and enlargement ofthe meta- first recognizable canids involved the formation conid on m2 so that it equals the protoconid in ofa rigid middle ear chamber, i.e., the ossification size. of an entotympanic bulla and development of a Although the middle ear structure of canids is low septum from the in-bent edge of caudal en- relatively stable when compared with that ofother totympanic. The middle ear region -s further families of caniform carnivorans, it does change strengthened by medial expansion of the petrosal in at least three aspects. Firstly, the low entotym- into contact with the basioccipital and basisphe- panic septum varies in size and extent, and tends noid, the ossification of the tegmen tympani, and to be confined to the anteromedial corner of the the closure of piriform fenestra. Another impor- bulla in derived forms. Secondly, the anterior loop tant character complex consists ofthe medial mi- of the internal carotid artery changes from intra- gration of the internal carotid utery, She loss of cranial to extracranial in position. Evidence for the stapedial artery, and the entrapment of the this latter feature is the loss of a bony signature of ' Frick Postdoctoral Fellow, Department of Vertebrate Paleontology, American Museum of Natural History. 2 Curator, Department of Vertebrate Paleontology, American Museum of Natural History. Copyright © American Museum of Natural History 1994 ISSN 0003-0082 / Price $3.60 2 AMERICAN MUSEUM NOVITATES NO. 3092 this loop in the basioccipital-an embayment on bony tube for the external auditory meatus de- the lateral wall of the basisphenoid anteroventral velops in later canids. to the middle lacerate foramen. Finally, a short INTRODUCTION Among caniform carnivorans, the basicra- does not prove that the reverse process may nial anatomy ofcanids has long been consid- not exceptionally take place, whether by re- ered relatively stable, and canid character- version or otherwise." Otocyon was therefore istics in the middle ear region are identifiable removed from the basal stock in Scott's phy- in the earliest canid Hesperocyon (Hough, logeny which still recognized Miacis as an- 1948; Tedford, 1976). This apparent stability cestral to canids. of the middle ear of canids, together with The miacid ancestry of canids has since their rather uniform dental patterns, makes been accepted by most students of carnivor- canids a morphologically conservative group. ans (e.g., Matthew, 1930; Clark, 1939; Ted- Basicranially, canids can be distinguished ford, 1978; Flynn and Galiano, 1982; Gus- from other families of carnivorans by pos- tafson, 1986). Various authors, however, session of an inflated bulla, formed mostly adopted different ways of defining the Cani- from an inflated caudal entotympanic; a low dae-some favored the inclusion of certain septum across the inner surface of the bulla; miacid genera in the Canidae consistent with a fully functional internal carotid artery in an their philosophy of a vertical classification extrabullar position; an external auditory (see Flynn and Galiano, 1982, for a recent meatus lacking a prominent bony tube (ex- review). As may be expected from such a cept derived forms); and a small mastoid pro- basal group, miacoids could be claimed as cess. ancestral to every modern family of carni- Cope (1877, 1883) first proposed that the vorans. Indeed, Flynn and Galiano (1982) ancestry ofcanids was to be found among the associated certain miacoids with the phylog- forms placed in his Creodonta. More specif- eny ofprimitive feliforms, and the rest ofthe ically, the Miacidae was postulated as the miacoids were set aside in an unresolved probable ancestral stock (Cope, 1880: 81). Caniformia. In a more recent analysis (Wyss The central theme of Cope's phylogeny re- and Flynn, 1993), however, the Miacidae was volved around a linear sequence of dental removed from direct relationship to the liv- reduction from four upper and lower molars ing families of Carnivora, and placed one in the African bat-eared fox (Otocyon mega- branch below the common ancestor of Viv- lotis), to a more "normal" dental formula in erravidae and Carnivora. Amphicyon (three upper and three lower mo- Among the various North American Eo- lars), and to variously reduced dentitions (e.g., cene miacids, Miacis, with its tendency to loss of pl, M3, and sometimes M2) in later elongate the carnassial shear and reduce the true canids. Even though the number of up- M3 (a "typical cynoid" character according per molars in Miacis was not known to Cope to Matthew, 1909: 362), appears morpholog- (1883), he correctly predicted that it would ically closest to the canids. Clark (1939), and be the same as in Amphicyon. Cope thus pre- later Gustafson (1986), went further and sug- dicted discovery of Otocyon-like fossils pre- gested that "Miacis" gracilis was closest to ceding other extinct dogs. Hesperocyon. The proposed affinity between Cope's theme of progressive dental reduc- "M. "gracilis and Hesperocyon was primarily tion was soon challenged by Scott (1895: 74) based on postcranial evidence, i.e., the ad- who realized that the supernumerary teeth in vanced canidlike limb bones of "M. "gracilis; Otocyon could be a result of character rever- dentally, however, "M. "gracilis was little dif- sal: "I am by no means convinced of the ferent from other species of Miacis. impossibility of the addition of new teeth to Apart from the abovementioned attempts the molar series. That modification in the to link a particular taxon to the ancestry of mammalian lines is very generally by way of the Canidae, the relationships between var- reduction in the number ofteeth, is true, but ious species of miacids on the one hand, and 1 994 WANG AND TEDFORD: CANIDS AND MIACIDS 3 most primitive canids on the other, remains of upper fourth premolar and lower first mo- unresolved. Tedford (1976: 364) remarked: lar. Miacoids are otherwise more or less de- "The fact that very few miacoids have been fined by the paucity of derived characters identified as phyletically related to members shared with advanced (mostly extant) fami- of the modern superfamilies only increases lies of carnivorans. The taxon Miacoidea is the isolation of the archaic and modern car- therefore one of convenience and has long nivore families. In other words, there are few been a taxonomic wastebasket waiting to be miacoids that possess derived characters emptied. Recent phylogenetic treatments uniquely shared with representatives of the range from dividing miacoids into feliform modern families. This represents one of the and caniform components (Flynn and Galia- largest gaps in our knowledge of the phylog- no, 1982), to placing them within a multi- eny of the Carnivora." chotomy of Feliformia + Viverravidae + Recent discoveries offorms from the Vieja Miacidae + Nimravidae etc. (Flynn et al., Group of Texas (Chadronian) intermediate 1988), and to postulating a sister-group re- between miacids and canids has significantly lationship between Miacidae and a Viverrav- bridged this gap in our knowledge of canid idae + Carnivora clade (Wyss and Flynn, ancestry (Gustafson, 1986). The Texas ma- 1993). In this discussion, the Miacidae in- terial includes the last known representatives cludes such archaic caniforms as Miacis, Uin- of Miacis in North America, "M." cognitus tacyon, Vulpavus, etc. (roughly equivalent to and M. australis. Their dental morphology is Miacinae of Simpson, 1945: 108), whereas also the most derived among all Miacis. An- the Miacoidea includes the caniform Miaci- other important taxon, also from the Vieja dae and the feliform Viverravidae. The pres- Group, is "Hesperocyon" wilsoni Gustafson ent study considers only a few putative mia- (1986), which reduces, in yet another way, cids, which share several derived characters, the morphological gap between the well- such as reduced or absent P4 parastyle, re- known White River H. gregarius and various duced P4 protocone, and internal cingulum Miacis. "H. " wilsoni has a fully ossified bulla of upper molars surrounding the protocone, with a high degree ofinflation resembling that in addition to various postcranial characters of H. gregarius. However, the dental mor- (Flynn and Galiano, 1982). Preservation of phology of "H." wilsoni is still primitively the basicranium is also important to the scope similar to that of miacid carnivorans. It re- of this study. tains a primitively short shearing blade on As discussed in the introduction, of the the ml with a relatively high trigonid and known miacid genera Miacis seems in general narrow talonid, and has a relatively large closest to canids because of its tendency to parastyle on the Ml. The Texas material dis- elongate the carnassials and to reduce the M3. plays an interesting combination of charac- The genus was established by Cope (1872) on ters, which not only offers an example ofwhat the type species M. parvivorus from the Bridg- a transitional canid may have looked like, er Formation (Bridgerian), Wyoming. Nearly but also raises new questions regarding the two dozen nominal species of Miacis have origin of canids.
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