Taxonomy and biostratigraphy of the early Tertiary Taeniodonta (Mammalia: Eutheria): Summary
ROBERT M. SCHOCH Department of Geology and Geophysics and Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06511
INTRODUCTION (northeastern Utah), Huerfano basin (south-central Colorado), San Juan basin (northwestern New Mexico and southwestern Colo- The Taeniodonta is an order of archaic mammals known rado), and Tornillo Flat area (western Texas). Here, I place special exclusively from the early Tertiary of western North America emphasis on the San Juan basin (Fig. 3), from which the early (Schoch and Lucas, 1981a). Their remains are found in Puercan Puercan to Wasatchian taeniodonts are best known. The history of (lower Paleocene) to Uintan (upper Eocene) strata of the Rocky study and nomenclature of the Tertiary strata of the San Juan basin Mountain intermontane sedimentary basins (Figs. 1, 2; Table 1). In (Fig. 4) has been discussed and reviewed in numerous papers, the San Juan basin of New Mexico and Colorado, the Puercan to Wasatchian (lower Eocene) sedimentary sequence includes the type localities for the Puercan, Torrejonian (middle Paleocene), and Tif- Figure 1. Localities at fanian (late Paleocene) land mammal "ages" (Wood and others, which taeniodonts have been 1941), as well as a classic Wasatchian fauna (Lucas and others, found. Numbers correspond 1981). Thus, the San Juan basin mammalian faunas provide a to localities listed in Table sequence of faunas which can provide the basis for interbasinal 1. For localities'in the San correlation. The Taeniodonta are an important element of these Juan basin (SJB), see Fig- faunas; many of the type specimens and important referred speci- ure 3. mens of these animals come from the San Juan basin. However, except for Patterson's (1949) brief review, a comprehensive study of this order has not been published for more than 80 yr (Wortman, 1897). Before a group of animals can be used for biostratigraphic correlation, its species- and genus-level taxonomy and its known stratigraphic occurrences must be well established. In Part //of this paper, I revise the species-level taxonomy of the Taeniodonta and document their geographic and temporal (stratigraphic) distribu- tion. Here, I discuss their usefulness and implications for biostrati- graphic correlation in western North America.
THE GEOGRAPHIC AND BIOSTRATIGRAPHIC DISTRIBUTION OF THE TAENIODONTA
Introduction
All unequivocally known taeniodonts come from the Rocky Mountain early Tertiary intermontane sedimentary basins of west- ern North America (Figs. 1, 2; Table 1), which were formed or rejuvenated during the Laramide orogeny. The major areas where taeniodonts occur are, from north to south: the Crazy Mountain Field (south-central Montana), Bighorn basin (north-central Wyoming), Togwotee Pass area (northwestern Wyoming), Wind River basin (central Wyoming), Green River-Bridger basin (southwestern Wyoming), Washakie basin (south-central Wyoming), Uinta basin
The complete article, of which this is a summary, appears in Part II of the Bulletin, v. 92, p. 1982-2267.
Geological Society of America Bulletin, Part I, v. 92, p. 933-941, 5 figs, 1 table, December 1981.
933
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including Baltz and others (1966), Gardner (1910), Granger (1914), "zones." These "/.ones" have been considered to represent super- Kues and others (1977), Lucas (1981), Matthew (1937), Reeside posed biostratigraphic units well separated temporally. Alterna- (1924), Simpson (1948, 1959, 1981), Sinclair and Granger (1914), tively, they have been thought to represent different facies or to Tsentas (1981), and Wood and others (1941). reflect collecting biases (Matthew, 1937). Previously, Onychodectes tisonensis has been thought to occur throughout the Puercan in both Puercan-Torrejonian "zones," whereas O. "rarus" and Wortmania otariidens were re- stricted to the Taeniolabis "zone" (Russell, 1967). Here, O. "rarus" Taeniodonts of Puercan and Torrejonian age occur in the is considered a junior subjective synonym of O. tisonensis at the Paleocene Nacimiento Formation at several localities in the specific level and W. otariidens is now known from the Ectoconus southwestern and south-central San Juan basin (Fig. 3; Table 1). "zone" in Betonnie Tsosie and Kimbeto washes. However, as far as The Nacimiento Formation is composed of red and green, buff and is known, O. t. tisonensis does occur in both zones, whereas O. t. gray clay shales and siltstones, black clay shales, and lenticular rarus is known only from the Taeniolabis "zone." Thus, better arkosic and quartzose sandstones (Baltz and others, 1966; Tsentas knowledge of the Puercan taeniodonts reduces the distinctiveness of and Lucas, 1980, Tsentas and others, 1981). In the upper part of the these "zones" and does not strongly support the idea that they are Nacimiento Formation, a northern facies of relatively high-energy separated by a significant span of time. fluvial deposits (with a greater over-all percentage of sandstone) Both O. tisonensis and W. otariidens are restricted to the and a southern facies of lower-energy fluvial and swamp deposits Puercan. The occurrence of O. tisonensis in the Wagonroad local are recognizable (Tsentas and others, 1981). This distribution of fauna of the upper part of the North Horn Formation of east- facies suggests a northern source area for much of the upper part of central Utah supports the correlation of this locality with the the Nacimiento Formation (Baltz, 1967; Tsentas and others, 1981). Puercan-aged strata of the Nacimiento Formation (Robison and The Puercan strata of the Nacimiento Formation have been Lucas, 1980). subdivided into two "zones," a lower Ectoconus "zone" (also known The Torrejonian strata of the Nacimiento Formation also have as the Hemithlaeus "zone"; Van Valen, 1978) and an overlying been divided into two "zones" on the basis of mammalian faunas Taeniolabis "zone" (formerly known as the "Polymastodon," (Lindsay and others, 1978; Matthew, 1937; Osborn, 1929). The pre- (-Taeniolabis) zone; Lindsay and others, 1978; Matthew, 1937; sumably lower Deltatherium "zone" occurs in Kutz Canyon, Tor- Osborn, 1929; Sinclair and Granger, 1914). Localities in Betonnie reon wash, and Kimbeto wash, whereas the overlying Pantolambda Tsosie wash and Kimbeto wash are in the Ectoconus "zone," where- "zone" is well known from Torreon wash and University of Kansas as localities in De-na-zin wash and Alamo wash include both New Mexico locality 15, as well as the areas south-east of Kimbeto
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UINTAN "5>
Figure 2. Biostratigraphic distribution of the Taeniodonta.
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and just south of Cedar Hill (Tsentas, 1981; Wilson, 1956). These 1937; Tsentas, 1981; Wilson, 1956). Recently, a specimen of Panto- "zones" also have been thought to represent superposed biostrati- lambda was found in a Deltatherium "zone" horizon in Kutz graphic units separated by a significant length of time (Lindsay and Canyon, supporting the idea of collecting biases (Lucas and O'Neill, others, 1978; Taylor and Butler, 1980). Alternatively, it has been 1981). Conoryctes comma is known with certainty only from the suggested that the differences betweeen the faunas of the two Pantolambda "zone." The specimens reported by Taylor (1981) as "zones" reflect facies differences and/or collecting biases (Matthew, Conoryctes comma from a Deltatherium "zone" horizon in Kutz
TABLE 1. SUMMARY OF THE STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION OF THE TAENIODONTA»
Locality Formation Age Taxa Reference
1. Betonnie Tsosie wash Nacimiento Puercan Onychodectes tisonensis tisonensis Simpson, 1959; Sinclair and Granger Wortmania otariidens 1914 2. Kimbeto wash Nacimiento Puercan O. t. tisonensis Simpson, 1959; Sinclair and Granger, W. otariidens 1914 3. De-na-zin wash Nacimiento Puercan O. t. tisonensis Sinclair and Granger, 1914 O. t. rarus W. otariidens 4. Alamo wash Nacimiento Puercan O. l. tisonensis Sinclair and Granger, 1914 O. 1. rarus W. otariidens 5. Wagonroad Ridge North Horn Puercan O. l. tisonensis Robison and Lucas, 1980 stylinodontine indet. 6. Dragon Canyon North Horn Torrejonian Conoryctella dragonensis Gazin, 1941; Schoch and Lucas, 1981c Conoryctella pattersoni tPsitlacotherium sp. or ? Wortmania sp 7. Kutz Canyon Nacimiento Torrejonian C. pattersoni Granger, 1917; Schoch and Lucas, Psittacotherium multifragum 1981c; R. W. Wilson, 1956 8. Kimbeto wash Nacimiento Torrejonian Huerfanodon torrejonius Schoch and Lucas, 1981b P. multifragum 9. Torreon wash Nacimiento Torrejonian Conoryctes comma Matthew, 1897; Tsentas, 1981 P. multifragum 10. UK NM locality 15 Nacimiento Torrejonian C. comma R. W. Wilson, 1956 P. multifragum 11. Gallegos Canyon Nacimiento Torrejonian P. multifragum Sinclair and Granger, 1914 12. Escavada wash Nacimiento Torrejonian conoryctine indet. Kues and others, 1977 P. multifragum 13. Simpson's locality 226 Nacimiento Torrejonian P. multifragum Simpson, 1959 14. Gidley Quarry Lebo Torrejonian conoryctine indet. Simpson, 1937 15. Silberling Quarry Lebo Torrejonian ? Huerfanodon sp. Simpson, 1937 P. multifragum 16. Rock Bench Quarry Polecat Bench Torrejonian Huerfanodon polecatensis Gingerich and others, 1980 17. Swain Quarry Fort Union Torrejonian P. multifragum Rigby, 1980 18. Douglass Quarry Melville Tiffanian P. multifragum Douglass, 1908; Simpson, 1937 19. Tornillo Flat Black Peaks Tiffanian P. multifragum Schiebout, 1974; J. A. Wilson, 1967 20. Plateau Valley Wasatch Tiffanian- Ectoganus gliriformis lobdelli Patterson, 1936, 1949a; Sloan and ("DeBeque") Clarkforkian others, 1980 21. Polecat Bench Polecat Bench Clarkforkian E. g. lobdelli Gingerich and others, 1980 22. Bear Creek Fort Union Clarkforkian E. g. lobdelli Simpson, 1929a, 1929b 23. Togwotee Pass area "Lower varie- Clarkforkian? E. g. gliriformis McKenna, 1972, 1980a gated beds" E. cf. E. copei bighornensis 24. Bighorn basin Willwood Wasatchian E. g. gliriformis Bown, 1980; Schankler, 1980 E. copei copei E. c. bighornensis 25. Almagre Arroyo San Jose Wasatchian E. g. gliriformis Granger, 1914; Simpson, 1948 26. Gobernador San Jose Wasatchian E. g. gliriformis Kues and others, 1977 27. Cerrillos Galisteo Wasatchian Ectoganus sp. Lucas and Kues, 1979; Lucas, 1982 28. Wind River basin Wind River Wasatchian E. g. gliriformis Granger, 1910; Guthrie, 1967, 1971 Stylinodon mirus 29. Huerfano basin Huerfano Wasatchian- S. mirus Robinson, 1966 Bridgerian 30. Green River-Bridger basin Bridger Bridgerian S. mirus West, 1972 31. Washakie basin Washakie Bridgerian S. mirus Turnbull, 1972, 1978 Stylinodon inexplicatus 32. Uinta basin Uinta Uintan S. mirus Black and Dawson, 1966 33. Brewster County, Texas Pruett Bridgerian- Stylinodon sp. J. A. Wilson, 1972, 1974 Uintan
•Descriptions, illustrations, and synonyms of taxa are given in Part //, as are references cited in this table but not elsewhere in Part I.
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Lucas, 1981b). Huerfanodon polecatensis is known from the Rock Bench Quarry of the Polecat Bench Formation, Bighorn basin (Schoch and Lucas, 1981b), and ?Huerfanodon sp. is known from the Silberling Quarry, Libo Formation, Montana. Both Rock Bench and Silberling Quarries are considered to be in the upper Torrejonian (Pantolambda "zone") (Gingerich and others, 1980). Although Huerfanodon polecatensis may appear to be slightly more "advanced" (and perhaps younger?) than Huerfanodon torre- jonius, it is unwise to try to hypothesize that one species is the ancestor of another and then base the relative correlation and dat- ing of the strata in different basins on such hypothetical lineages (icontra Gingerich, 1976). Rather, the occurrence of Huerfanodon in New Mexico, Wyoming, and Montana suggests that these strata all are approximately the same age. Tomida (1981) recently has described a typical Torrejonian mammalian faunal assemblage from the San Juan basin in sedi- ments that can be magrietostratigraphically correlated with the Dragon local fauna of the North Horn Formation, Utah (Tomida and Butler, 1980), the type locality of the "Dragonian" land mam- mal "age" (Wood and others, 1941). These sediments also are strati- graphically below a Deltatherium "zone" horizon and suggest the presence of a third, earliest Torrejonian "zone." Conoryctella pat- tersoni occurs in Tomida's (1981) "Dragonian" zone in the San Juan basin, in the Kutz Canyon Deltatherium "zone" (Wilson, Figure 3. Geologic map of lower Tertiary strata of the San 1956), and in the Dragon local fauna of Utah (Schoch and Lucas, Juan basin, New Mexico, showing major Paleocene localities which 1981c). Thus, these occurrences of Conoryctella suggest that all have produced taeniodont fossils. For lower Eocene localities in the three localities do not significantly differ temporally. The Dragon San Juan basin, see Figures 2 and 3 in Lucas and others (1981) (see local fauna thus can be correlated with the "lower" Torrejonian also Table 1; geology modified after Fassett and Hinds, 1971, PI. 1). strata of the Nacimiento Formation. Canyon here are assigned to Conoryctella (Schoch and Lucas, Psittacotherium is a far-ranging genus, both geographically 1981c). However, because it is relatively rare, I do not stress the and temporally. It occurs throughout the Torrejonian strata (all possible biostratigraphic significance of the absence of C. comma in three "zones") of the Nacimiento Formation. It may also occur in the Deltatherium "zone." the Dragon local fauna, and, if this is substantiated, it would Huerfanodon torrejonius is now known in the San Juan basin further support a Torrejonian age for this fauna. Psittacotherium is only from the Deltatherium "zone" in Kimbeto wash (Schoch and present in the Torrejonian Swain Quarry, Fort Union Formation,
SW UTHOLOGY [schematici NE STRAT. UNITS & LOCAL FAUNAS N. frMERPBOV. "AGES' EUROP. STAGES LEGEND
Q ALLUVIUM
[H CONGLOMERATE
SANDSTONE
G] ANDESITE
HH MUDSTONE
• COAL
UNCONFORMITY
["""] UNFOSSILIFEROUS
ALMAGRE LOCAL A FAUNA
MAASTRKXT1AN LATE CRETACEOUS R. SCHOCH. 1961
Figure 4. The stratigraphy of the lower Tertiary strata of the San Juan basin. Lithologies are schematic. Stratigraphic units after Baltz (1967). Local faunas after Sinclair and Granger (1914), Matthew (1937), Simpson (1935a, 1935b, 1935c), and Lucas and others (1981). North American land mammal "ages" after Wood and others (1941). Correlation of North American "ages" with European stages after Berggren and others (1978). Abbreviations for faunas are: A = Almagre local fauna, AW = Alamo Wash local fauna, L = Largo local fauna, P = Puerco fauna, T = Torrejonian fauna, Ti = Tiffany fauna.
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Wyoming, and in the upper Torrejonian-lower Tiffanian strata of from the "west branch of Almagre Arroyo, upper beds." This local- Montana (Gidley, Lebo, and Douglass Quarries) and Texas (Tor- ity corresponds to the lower part of the Largo of Granger (1914) nillo Flat, Black Peaks Formation). and is in the upper part of the Regina Member of the San Jose Formation. Tiffanian-Uintan The Wasatchian land mammal "age" is usually divided into three "subages," the Graybullian, Lysitean, and Lostcabinian, from In the northeastern San Juan basin, the Nacimiento Formation oldest to youngest (Granger, 1914; Wood and others, 1941). Ecto- grades laterally into the upper part of the Animas Formation, and ganus gliriformis gliriformis (= Ectoganus "simplex"-, Schankler, the Ojo Alamo Sandstone grades into the lower Animas (McDer- 1980, p. 104) occurs in Graybullian and Lysitean strata of the Will- mott Member; Baltz, 1967; Barnes and others, 1954; Reeside, 1924). wood Formation, Bighorn basin, Wyoming, but not in the Lostcab- The type locality of the Tiffanian land mammal "age" (Wood and inian strata of the Willwood. Likewise, in the Wind River others, 1941), Mason Pocket, is in the Animas Formation in south- Formation of the Wind River basin, Wyoming, the type area of the ern Colorado, but has not produced any taeniodonts (Barnes and Lysitean and Lostcabinian, E. g. gliriformis occurs only in the Lysi- others, 1954; Simpson, 1935a, 1935b, 1935c). Psittacotherium mul- tean strata and is superseded by the taeniodont Stylinodon mirus in tifragum does occur in lower Tiffanian strata of Montana (Melville the Lostcabinian (Guthrie, 1967, 1971). Thus, Ectoganus is not Formation; Simpson, 1937) and Texas (Black Peaks Formation; known from strata younger than Lysitean. The presence of E. g. Schiebout, 1974). gliriformis in the Almagre fauna thus suggests a Graybullian or "Lampadophorus lobdelli" and "L. expectalus" are junior sub- Lysitean rather than a Lostcabinian age for that fauna (contra jective synonyms of Ectoganus gliriformis. Ectoganus gliriformis Lucas, 1977), and the occurrence of E. g. gliriformis in the lower lobdelli occurs in upper Tiffanian-Clarkforkian strata of Colorado part of the Largo would also suggest that that horizon of the Largo and in the Clarkforkian of the Bighorn basin, Wyoming. E. gliri- (Granger, 1914) in the Regina Member of the San Jose Formation formis gliriformis occurs in the possible Clarkforkian strata of the is of Graybullian or Lysitean age. Nevertheless, the majority of the Togwotee Pass area, but otherwise is restricted to the lower-middle Largo fauna, which occurs stratigraphically higher in the Tapicitos Wasatchian of Wyoming and New Mexico. Except for a single Member, may be of Lostcabinian age (Lucas, 1977). specimen of Ectoganus cf. E. copei bighornensis from the possible Southeast of the San Juan basin in north-central New Mexico Clarkforkian strata of the Togwotee Pass area, both E. copei copei in the Wasatchian Cerrillos local fauna of the Galisteo Formation and E. copei bighornensis are restricted to the lower Wasatchian of (Lucas and Kues, 1979), a humerus of Ectoganus sp. recently has the Bighorn basin, Wyoming. Rose (1977) has used "Lampado- been found (Lucas, 1982), supporting the assignment of a pre- phorus"\n part to support the validity of the Clarkforkian land Lostcabinian Wasatchian age to the Cerrillos local fauna. mammal "age"; the revised taxonomy of Ectoganus eliminates The earliest occurrence of Stylinodon mirus is in the upper "Lampadophorus" as a distinctive Clarkforkian, or even latest Wasatchian (Lostcabinian) strata of the Wind River basin, Wyo- Paleocene, genus. ming. It also occurs in the upper Wasatchian-lower Bridgerian of The San Jose Formation ("Wasatch" of early workers), a series Colorado, in the Bridgerian of the Green River-Bridger and Washa- of variegated continental mudstones and sandstones (Simpson, kie basins of Wyoming, and in the middle Uintan ("Horizon B") of 1948), unconformably overlies the Nacimiento Formation and is in the Uinta basin, Utah. Stylinodon inexplicatus is known from only the structural center of the San Juan basin. Granger (1914) distin- one specimen of Bridgerian age in the Washakie basin. Two speci- guished two Wasatchian facies in the San Jose: the lower variegated mens of Stylinodon sp. are known from upper Bridgerian or lower "Almagre beds" exposed in Almagre and Blanco arroyos near the Uintan-aged strata of the Pruett Formation, Trans-Pecos, west present town of Regina, and the upper red "Largo beds" exposed Texas (Schoch and Lucas, 1981d). The presence of Stylinodon, near Lindrith and Gavilan (see Simpson, 1948). Cope's specimens therefore, indicates a Lostcabinian-Uintan age. of Ectoganus and "Calamodon" as well as Marsh's "Dryptodon" almost surely came from the Almagre fauna, as have most taenio- Conclusions donts since then (Lucas, 1977; Lucas and others, 1981). Recently (1977), a specimen of Ectoganus was collected from rocks strati- In conclusion, the early to middle Paleocene conoryctines graphically equivalent to the "Almagre beds" near Gobernador. (Onychodectes, Conoryctella, Conoryctes, Huerfanodori) appear to Baltz (1967) defined and mapped four formal members of have the most potential for dating and biostratigraphic correlation. Simpson's San Jose Formation. The Cuba Mesa Member is the The species and subspecies of stylinodontines(,H/o/7maw'a, Psitta- lowest and is essentially unfossiliferous (Lucas, 1977); the overlying cotherium, Ectoganus, Stylinodon) are either known only from a Regina Member includes the Almagre fauna and the lower part of few specimens or are so broadly distributed geographically and of the Largo fauna; the Llaves Member is essentially unfossiliferous such long durations temporally that at present they are not useful (Lucas, 1977) and either overlies the Cuba Mesa Member or the except for very general correlations no more refined than the level Regina member, or grades into and intertongues with the Regina of land mammal "ages." Member. The Tapicitos Member is the youngest member and either overlies or laterally grades into the Llaves Member; it includes the APPENDIX 1. SYSTEMATIC PALEONTOLOGY majority of the Largo fauna (Lucas, 1977). The only taeniodont taxon known from the San Jose Forma- I recognize 8 genera and 15 species and subspecies of taeniodonts, tion is Ectoganus gliriformis gliriformis. Although the vast majority limited in distribution to western North America (Figs. 1, 2; Table 1). The of specimens of Ectoganus has come from the Almagre local fauna, taxonomies of Onychodectes and Ectoganus are revised here in order to fulfill the rules of the International Code of Zoological Nomenclature (Stoll E. gliriformis gliriformis is also known from the Largo local fauna. and others, 1964). They are more fully described, discussed, and illustrated American Museum of Natural History (New York) 16425, a lower- in Part II of this paper, along with the other six genera of taeniodonts. jaw fragment of E. g. gliriformis, was collected by Granger in 1912 References cited only in this appendix to Part I are given in Part II.
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Order Taeniodonta Cope, 1876 lingyally placed and narrower than the trigonids; M1-3 transversely bilo- Onychodectes Cope, 1888d phodont with subequal trigonids and talonids. Onychodectes Cope, 1888d, p. 317 Type Species. Onychodectes tisonensis Cope, 1888d (= Onychodectes Ectoganus gliriformis Cope, 1874 rarus Osborn and Earle, 1895). Ectoganus gliriformis Cope, 1874, p. 592. Included Species. Only the type species. (see synonymies under the subspecies) Distribution. Puercan of New Mexico and Utah. Type Subspecies. Ectoganus gliriformis gliriformis Cope, 1874. Revised Diagnosis. Small taeniodonts; teeth moderately hypsodont Included Subspecies. The type subspecies and Ectoganus gliriformis (relatively less hypsodont than Conoryctella); P4 nonmolariform with a lobdelli (Simpson, 1929b). well-developed protocone, paracone and incipient metacone, parastyle, Revised Diagnosis. Largest species of Ectoganus (see Table 7 of Part II). stylocone, and metastyle; P" metastylocone small to moderately well devel- oped; lower molar trigonids bear large, subequal, and sharply punctate Ectoganus gliriformis gliriformis Cope, 1874, new rank protoconids and metaconids with only slightly smaller, lingually placed Ectoganus gliriformis Cope, 1874, p. 592. paraconids; lower molar talonids bear high punctate hypoconids, slightly Calamodon simplex Cope, 1874, p. 593. smaller and punctate entoconids, and small hypoconulids. Calamodon arcamaenus Cope, 1874, p. 593. Calamodon novomehicanus Cope, 1874, p. 594. Onychodectes tisonensis Cope, 1888d Dryptodon crassus: Marsh, 1876b, p. 403. Onychodectes tisonensis Cope, 1888d, p. 318 Ectoganus novomehicanus: Cope, 1877, p. 159. (see synonymies under the subspecies) Ectoganus gliriformis: Cope, 1877, p. 160. Type Subspecies. Onychodectes tisonensis tisonensis Cope, 1888d. Calamodon arcamaenus: C)ope, 1877, p. 163. Included Subspecies. The type subspecies and Onychodectes tisonensis Calamodon simplex: Cope, 1877, p. 166. rarus Osborn and Earle, 1895. Calamodon simplex: Cope, 1884c, p. 189. Diagnosis. Same as that for the genus. Calamodon simplex: Wortman, 1897b, p. 88. Calamodon arcamnaeus (lapsus calami)-. Wortman, 1897b, p. 89. Onychodectes tisonensis tisonensis Cope, 1888d, new rank Ectoganus gliriformis: Gazin, 1936, p. 610 (in part). Onychodectes tisonensis Cope, 1888d, p. 318. Ectoganus cf. simplex: Guthrie, 1967, p. 23. Onychodectes tissonensis (lapsus calami): Osborn and Earle, 1895, p. 40. Ectoganus simplex: Schankler, 1980, p. 104. Onychodectes tissonensis (lapsus calami): Wortman, 1897b, p. 97. Type Specimen. USNM (= National Museum of Natural History, S. tissonensis (lapsus calami): Wortman, 1897b, p. 97. Washington, D.C.) 1137, right and left I3 and C1 fragments, (?)right dP3"4, Onychodectes tisonensis: Matthew, 1937, p. 239. upper (?)deciduous incisor fragment, right P2, left dP», partial lower molar Onychodectes n. sp.?: Robison and Lucas, 1980, p. 302. trigonid, partial lower molar talonid, fragmentary upper molar and asso- Type Specimen. AMNH (= American Museum of Natural History, New ciated bone and tooth fragments. York, New York) 3405, right and left maxillae with P4-M3, left dentary with Horizon and Locality of the Type. Collected by E. D. Cope in 1874 M2, and associated right astragalus. from Wasatchian strata of the San Jose Formation, probably in Almagre Horizon and Locality of the Type. Collected by David Baldwin in 1885 Arroyo, San Juan Basin, New Mexico. from presumably Puercan strata of the Nacimiento Formation, San Juan Revised Diagnosis. Large Ectoganus with all teeth extremely hypso- basin, New Mexico. dont; incisors and P|l2 approach the totally rootless condition of the Revised Diagnosis. Subspecies of Onychodectes tisonensis with rela- canines. tively simpler crowned premolars and molars than in O. t. rarus; anterior internal accessory cusp absent on P4; external accessory cusp between tri- Ectoganus gliriformis lobdelli (Simpson, 1929b), new combination gonid and talonid lobes of M1-2 absent. and new rank Psittacotherium sp. indet.: Simpson, 1929a, p. 121. Onychodectes tisonensis rarus Osborn and Earle, 1895, new rank ! Psittacotherium lobdelli Simpson, 1929b, p. 11. Onychodectes rarus Osborn and Earle, 1895, p. 42. ?Psittacotherium sp.: Patterson, 1936, p. 397. Onychodectes rarus: Wortman, 1897b, p. 97. Lampadophorus expectatus Patterson, 1949a, p. 42. Onychodectes rarus: Matthew, 1937, p. 249. Lampadophorus lobdelli: Patterson, 1949a, p. 42. Type Specimen. AMNH 824, left dentary fragment with M1-2. Type Specimen. AMNH 22234, right M3. Horizon and Locality of the Type. Puercan strata of the Nacimiento Horizon and Locality of the Type. Clarkforkian strata of the Fort Formation, De-na-zin wash, San Juan basin, New Mexico. Union Formation, Eagle Coal Mine, Bear Creek Montana. Revised Diagnosis. Subspecies of Onychodectes tisonensis with the fol- Revised Diagnosis. Large Ectoganus with P5 only slightly more hypso- lowing characters moderately to well developed (as compared to O. t. dont than in Psittacotherium; posterior cheek teeth (P3-M3) moderately rarus): P4-M3 slightly broader and with better developed ectocingula; lower hypsodont with relatively low, rounded, bulbous crowns, and relatively shal- premolars slightly more molariform with better developed talonids on P3-4 low, compressed roots. and an anterior internal accessory cusp on P4; M1-2 with external acces- sory cusp or cusps between the trigonid and talonid lobes. Ectoganus copei Schoch, new species Ectoganus gliriformis: Gazin, 1936, p. 597 (in part). Ectoganus Cope, 1874 Type Subspecies. Ectoganus copei copei Schoch, new subspecies. Ectoganus Cope, 1874, p. 592. Included Subspecies. The type subspecies and Ectoganus copei bighor- Calamodon Cope, 1874, p. 593. nensis Schoch, new subspecies. Dryptodon Marsh, 1876b, p. 401. Etymology. Named for Edward Drinker Cope, the great Nineteenth- Conicodon Cope, 1894, p. 594. Century American paleontologist. non Calamodon Amaral, 1935, p. 203. Diagnosis. Smallest species of Ectoganus (see Table 8 of Part II). Lampadophorus Patterson, 1949a, p. 41. Type Species. Ectoganus gliriformis Cope, 1874 (- Calamodon simplex Cope, 1874 = Calamodon arcamaenus Cope, 1874 'Calamodon novomehi- Ectoganus copei copei Schoch, new subspecies (Appendix Fig. 1, a-d) canus Cope, 1874 = Dryptodon crassus Marsh, 1876b = 1 Psittacotherium Ectoganus gliriformis: Gazin, 1936, p. 597 (in part). lobdelli Simpson, 1929b = Lampadophorus expectatus Patterson, 1949a). Type Specimen. USNM 12714, skull and mandible with right and left Included Species. The type species and Ectoganus copei Schoch, new P-M1 (P4's unerupted), left M2, right and left dP4, alveoli for right M2 and species. left M3, roots or right and left Ci, right and left P2, left P4 (unerupted), left Distribution. Tiffanian-Wasatchian of Colorado, Clarkforkian of dP4, right and left M|, left M2, right and left M3, roots of right P4, alveoli Montana, Clarkforkian-Wasatchian of Wyoming, and Wasatchian of New for right MI_2. Mexico. Horizon and Locality of the Type. Wasatchian strata of the Willwood Revised Diagnosis. Medium- to large-sized taeniodonts; canines Formation, 13 km northwest of Worland, Bighorn basin, Wyoming. enlarged, rootless, and compressed posteriorly with enamel limited to the Diagnosis. Small Ectoganus with all teeth extremely hypsodont; inci- antero-labial aspect; cheek teeth moderately to extremely hypsodont; M1 3 sors and P\~_2 approach the totally rootless condition of the canines; cusps on transversely bilophodont; P3_4 submolariform to molariform with talonids upper premolars larger and better developed than in E. c. bighornensis.
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Appendix Figure 1. The type specimens of Ectoganus copei copei Schoch, new species and subspecies (a-d) and Ectoganus copei bighornensis Schoch, new subspecies (e-s). (a) USNM 12714, palate with right and left I3-M' (P4's unerupted), right and left dP4, left M2, and alveoli for right M2 and left M3: occlusal view, (b) USNM
12714, left P2: posterior view, (c) USNM 12714, left dP4, partial M,, M2, and M3 talonid: occlusal view, (d) USNM 12714, right M,: occlusal view, (e-g) PU 14678,, canine fragments, (h-i) PU 14678, right P3: (h) occlusal view; (i) posterior view, (j-k) PU 14678, left P3: (j) occlusal view; (k) anterior view. (1-m) PU 14678, right M1: (1) occlusal view; (m) posterior view, (n-o) PU 14678, right M2: (n) occlusal view; (o) posterior view, (p-q) PU 14678, left M3: (p) occlusal view; (q) posterior view, (r-s) PU 14678, right M,: (r) occlusal view; (s) lingual view. The bar below a is 1 cm long and is for a-d, h, j, I, n, p, r. The bar above o and q is 1 cm long and is for e-g, i, k, m, o, q, s.
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Ectoganus copei bighornensis Schoch, new subspecies Lucas, S. G., and O'Neill, F. M., 1981, Occurrence of Pantolambda (Mam- (Appendix Fig. 1, e-s). malia; Pantodonta) in the Torrejonian Deltatherium "zone," San Juan Type Specimen. PU (= Princeton University, Princeton, New Jersey) basin, New Mexico: American Journal of Science, v. 281, p. 187-191. 14678, right and left P3, right M1"2, left M3, right M| and canine fragments. Lucas, S. G., Schoch, R. M„ Manning, E., and Tsentas, C„ 1981, The Horizon and Locality of the Type. Lower Wasatchian strata of the Eocene biostratigraphy of New Mexico: Geological Society of America lower part of the Willwood Formation, southern tip of Polecat Bench, T. 55 Bulletin, v. 92 (this issue). N., R. 100 W., Bighorn basin, Wyoming. Matthew, W. D., 1937, F'aleocene faunas of the San Juan basin, New Etymology. Named after the Bighorn basin, Wyoming. Mexico: American Philosophical Society Transactions, n.s., v. 30, Diagnosis. 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Stoll, N. R., Dollfus, R. P., Forest, J., Riley, N. D., Sabrosky, C. W„ Juan basin paleontology: Albuquerque, New Mexico, University of Wright, C. W., and Melville, R. V., 1964, International Code of Zoo- New Mexico Press, p. 264-292. logical Nomenclature: London, International Trust for Zoological Tsentas, C., and Lucas, S. G., 1980, Position of the Paleocene-Eocene Nomenclature, 176 p. boundary in the south-central San Juan basin, New Mexico: Geological Taylor, L. H., 1981, The Kutz Canyon local fauna, Torrejonian (middle Society of America Abstracts with Programs, v. 12, p. 538. Paleocene) of the San Juan basin, New Mexico, in Lucas, S. G., Rigby, Tsentas, C., Lucas, S. G., and Schoch, R. M., 1981, Lithofacies of the upper J. K., Jr., and Kues, B. S., eds., Advances in San Juan Basin paleon- part of the Nacimiento Formation, San Juan basin, New Mexico: tology: Albuquerque, New Mexico, University of New Mexico Press, Geological Society of America Abstracts with Programs, v. 13, p. 229. p. 242-263. Van Valen, L., 1978, The beginning of the age of mammals: Evolutionary Taylor, L. H., and Butler, R. F., 1980, Magneticrpolarity stratigraphy of Theory, v. 4, p. 45-80. Torrejonian sediments, Nacimiento Formation, San Juan basin, New Wilson, R. W., 1956, A new multituberculate from the Paleocene Torrejon Mexico: American Journal of Science, v. 280, p. 97-115. fauna of New Mexico: Kansas Academy of Science Transactions, v. 59, Tomida, Y., 1981, "Dragonian" fossils from the San Juan basin and status p. 76-84. of the "Dragonian" land mammal "age," in Lucas, S. G., Rigby, Wood, H. E., Chaney, R. W., Clark, J., Colbert, E. H„ Jepsen, G. L., J. K., Jr., and Kues, B. S., eds., Advances in San Juan basin paleon- Reeside, J. B., Jr., and Stock, C., 1941, Nomenclature and correlation tology: Albuquerque, New Mexico, University of New Mexico Press, of the North American continental Tertiary: Geological Society of p. 222-241. America Bulletin, v. 52, p. 1-48. Tomida, Y., and Butler, R. F., 1980, Dragonian mammals and Paleogene Wortman, J. L., 1897, The Ganodonta and their relationship to the Eden- magnetic polarity stratigraphy, North Horn Formation, central Utah: tata: Bulletin of the American Museum of Natural History, v. 9, American Journal of Science, v. 280, p. 787-811. p. 59-110 Tsentas, C., 1981, Mammalian biostratigraphy of the middle Paleocene (Torrejonian) strata of the San Juan basin: Notes on Torreon wash and the status of the Pantolambda and Deltalherium faunal "zones," in MANUSCRIPT RECEIVED BY THE SOCIETY OCTOBER 8, 1981 Lucas, S. G., Rigby, J. K., Jr., and Kues, B. S., eds., Advances in San MANUSCRIPT ACCEPTED OCTOBER 8, 1981
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