Morphology of the Auditory Region in Paramys Copei and Other Eocene Rodents from North America

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Morphology of the Auditory Region in Paramys Copei and Other Eocene Rodents from North America PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3307, 16 pp., 4 ®gures, 1 table 13 December, 2000 Morphology of the Auditory Region in Paramys copei and Other Eocene Rodents from North America JOHN H. WAHLERT1 ABSTRACT The morphology of the external surface of the petrosal and its junction with the basioccipital in Paramys copei is described. Comparison with an outgroup of the Cretaceous Vincelestes and Recent Didelphis and Solenodon reveals that the auditory region retains many primitive features. The Eocene paramyines from North America show only slight differences. Sciuravus is set apart by the lack of a ventral petrosal sinus canal between the petrosal and basioccipital and by the facial nerve and stapedial artery sharing a common foramen in the petrosal. Eu- ropean theridomyids, too, are not as primitive as Paramys but share with it and Sciuravus a ridge on the promontorium that separates the transpromontorial continuation of the internal carotid artery from the origin of the tensor tympani muscle. Twelve characters of the auditory region are analyzed and summarized in a data matrix for use in future studies. Relative prim- itiveness of the auditory region in all of these Eocene rodents suggests that derived characters in later rodent taxa may represent intraordinal relationships but only homoplasy with nonro- dents. INTRODUCTION in publications on cranial anatomy. Thus, a Rodent auditory regions are suf®ciently wealth of additional characters exists that can known for some characters to be used in phy- be described and evaluated for polarity (prim- logenetic analyses. However, many visible itive or derived). Paramys (family Ischyro- structural details are not routinely described myidae) is a key taxon in this category. It is College, City University of New York; member of doctoral faculty, CUNY Ph.D. program in biology: ecology, evolutionary biology, and behavior subprogram. 2 AMERICAN MUSEUM NOVITATES NO. 3307 one of the oldest fossil rodents in which suf- 1962, ®gs. 13 and 14; Wahlert, 1974, ®g. 2. ®cient detail has been preserved, and, in Similarly, independent versions of the audi- North American paleontology, the genus has tory region have been published: Wood, been placed near the base of the rodent family 1962, ®g. 14a; Wahlert, 1974, ®g. 4; Parent, tree as a kind of structural ancestor or ex- 1980, pl. 2, ®gs. 2, 4; Lavocat and Parent, ample of primitiveness (Gregory, 1951: ®g. 1985, ®g. 1c. However, I found that there 19.40; Wood, 1962: ®g. 90). One expects it was still matrix obscuring morphology in the to be almost wholly primitive for rodents. auditory region and even a matrix-covered Other contenders for structural primitiveness petrosal displaced posterolaterally on AMNH among rodents are the North American genus 4755. To sit at a microscope and carefully Sciuravus, which McKenna and Bell (1997: use the sharpened tip of a thin beading nee- 186) put in a new and different suborder from dle to push away the ®ne covering matrix the Ischyromyidae, and the European theri- gives one an intimate feel for the shapes and domyids (Lavocat and Parent, 1985: 338). Li edges as they are revealed. Identi®cation of et al. (1989) described the auditory region of these features and comparisons with other Cocomys, a rodent from the early Eocene of living and extinct mammals were made from China; future restudy of this specimen should the literature mentioned below and from the yield important comparative details. ontogenetic studies by MacPhee (1981) and My primary purpose in this paper is to de- Wible (1984). scribe and illustrate the auditory region in Among the ®rst major comparative studies Paramys copei, AMNH 4755 (holotype) and on auditory regions in mammals was that of 4756. These two skulls preserve the ®nest de- van Kampen (1905), who built primarily on tail that I have seen in any fossil rodent, and the ®ne German work in anatomy and de- their early Eocene age, approximately 51 mya velopment. Van der Klaauw (1931) made (million years ago), makes them important by fossil mammals a part of his monograph; he antiquity. The rodent branch of mammalian was able to take advantage of the descrip- phylogeny may have been separate already in tions and collections of fossils from the Asia in the early Paleocene (Heomys Li, American West that had accumulated since 1977); undisputed rodents are known from the the late 1800s. Recent publications provide late Paleocene: Alagomyidae in Asia and character analyses and detailed information North America and Ischyromyidae in North about early mammals (Wible, 1990; Rougier America. In North America, the appearance et al., 1992; Wible and Hopson, 1993; Wible of ischyromyid rodents de®nes the beginning et al., 1995; Rougier et al., 1996; McKenna of the Clarkforkian Mammal Age (Woodbur- et al., 2000) and insectivores (McDowell, ne, 1987: 61) about 56 mya (McKenna and 1958; Novacek, 1986). The pioneering com- Bell, 1997: ®g. 1). The early Eocene saw si- parative studies on rodent auditory regions multaneous diversi®cation of rodents in Asia, are relatively recent (Oaks, 1968; Parent, North America and Europe. My second pur- 1980; Lavocat and Parent, 1985). Many stud- pose is to compare the auditory region of Par- ies of living rodents have focused on taxa amys copei with an outgroup of thoroughly with in¯ated bullae (Webster, 1961, 1962, described and illustrated mammals including 1975; Pye, 1965; Webster and Webster, 1971, the early mammal Vincelestes, the marsupial 1975, 1977, 1984; Lay, 1972, 1993; Wahlert, Didelphis, and the insectivore Solenodon, and et al., 1993). Segall (1971) illustrated gliding with other Eocene rodents. My goals are to and non-gliding sciurids. Information is also describe character polarities as a starting point available about speci®c extinct taxa (Lavo- for comparison of rodent auditory regions and cat, 1967; Meng, 1990; Wahlert, 1974, 1977, to assess the relative primitiveness of Para- 1978, 1983; Carrasco and Wahlert, 1999). mys auditory morphology. The two specimens of Paramys copei were SPECIMENS EXAMINED collected by Jacob L. Wortman in 1880 for Edward D. Cope. Images of the skulls have Specimens are listed according to the been independently illustrated and published: classi®cation of McKenna and Bell (1997); Cope, 1884, pl. 24a, ®gs. 1, 1a, 2, 2a; Wood, generic assignments of paramyine species are 2000 WAHLERT: AUDITORY REGION IN PARAMYS COPEI 3 according to Korth (1984, 1985, and 1994). Suborder Sciuravida, Family Sciuravidae Abbreviations: AMNH, American Museum Sciuravus nitidus, AMNH 12531 and 12551, of Natural History; USNM, United States USNM 17683 and 22477, Blacks Fork Mem- National Museum of Natural History. ber, middle Eocene, Bridger Formation, Class Mammalia, Legion Cladotheria, Suble- Bridger Basin, Wyoming gion Zatheria, Family Vincelestidae Vincelestes neuquenianus La Amarga Forma- AUDITORY REGION OF tion, early Cretaceous, Southern NeuqueÂn PARAMYS COPEI Province, Argentina (information from Rougier et al., 1992) The late early Eocene species Paramys copei (AMNH 4755, 4756, and 103390) pre- Sublegion Zatheria, Supercohort Theria, Cohort serves the most ancient, complete example Marsupialia, Magnorder Ameridelphia, Order of auditory region morphology that is avail- Didelphimorphia, Family Didelphidae, Subfam- able for rodents. I describe it below as a stan- ily Didelphinae dard, and then note the differences of other Didelphis virginiana, AMNH 2070, Recent, no Eocene rodent taxa. The region is illustrated data. in slightly oblique ventral view in ®gures 1, Sublegion Zatheria, Supercohort Theria, Cohort 2, and 3. Placentalia, Magnorder Epitheria, Superorder In ventral view the junction of the petrosal Preptotheria, Grandorder Lipotyphla, Order with the basioccipital is oblique, and the ba- Soricomorpha sioccipital is widest posteriorly. The anterior Solenodon paradoxus, AMNH 28270, Recent, part of the basioccipital and adjacent basi- Haiti sphenoid appears swollen and may be pneu- Sublegion Zatheria, Supercohort Theria, Grandor- matized in the anterior portion. The posterior der Anagalida, Mirorder Simplicidentata, Order part is a ¯ange that ¯oors the ventral petrosal Rodentia. All specimens are of Eocene age. sinus. The posterior lacerate foramen is a len- ticular gap medial to the posterior part of the Suborder Sciuromorpha, Family Ischyromyidae, auditory chamber and situated between the Subfamily Paramyinae: petrosal and the occipital. A broad ascending Tribe Paramyini keel of the petrosal partly divides the anterior Paramys copei, AMNH 4755 (holotype) and part of the gap, which transmitted the sig- 4756; Lost Cabin Member, late early Eocene, moid sinus, from the posterior, for nerves IX, Wind River Formation, Wind River Basin, X, and XI. A prominent dimple in the petro- Wyoming; AMNH 103390, late early Eo- sal just anterior to the keel is the opening of cene, San Jose Formation, San Juan Basin, the cochlear canaliculus. A broad, covered New Mexico passage between the basioccipital and the pe- Paramys delicatus, AMNH 12506 and 13090, trosal may mark the course of the ventral (in- Blacks Fork Member, middle Eocene, Bridg- ferior) petrosal sinus. It opens just anterior to er Formation, Bridger Basin, Wyoming Notoparamys costilloi (type species) (5 Lep- the posterior lacerate foramen. The sigmoid totomus costilloi), AMNH 55110 and 55111 sinus and ventral petrosal sinus joined here (holotype), Lost Cabin Member equivalent to form the internal jugular vein. A deep, (Wind
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