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Early Eocene Bats (Mammalia, Chiroptera) and Other Vertebrates in Freshwater Limestones of the Willwood Formation, Clark's Fork Basin, Wyoming

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Early Eocene Bats (Mammalia, Chiroptera) and Other Vertebrates in Freshwater Limestones of the Willwood Formation, Clark's Fork Basin, Wyoming CONTRIBLTIO\S FRO\l THE \IL SEL \I OF PALEO1TOLOGY THE UNIVERSITY OF MICHIGAN VOL 27. No I I. p 275-320 October 23. 1987 EARLY EOCENE BATS (MAMMALIA, CHIROPTERA) AND OTHER VERTEBRATES IN FRESHWATER LIMESTONES OF THE WILLWOOD FORMATION, CLARK'S FORK BASIN, WYOMING BY PHILIP D. GINGERICH MUSEUM OF PALEONTOLOGY THE UNIVERSITY OF MICHIGAN ANN ARBOR CONTRIBUTIONS FROM THE MUSEUM OF PALEONTOLOGY Charles B. Beck. Director Jennifer A. Kitchell, Editor This series of contributions from the Museum of Paleontology is a medium for publication of papers based chiefly on collections in the Museum. When the number of pages issued is sufficient to make a volume, a title page and a table of contents will be sent to libraries on the mailing list, and to individuals upon request. A list of the separate issues may also be obtained by request. Correspond- ence should be directed to the Museum of Paleontology, The University of Michigan, Ann Arbor, Michigan 48 109. VOLS. 11-XXVII. Parts of volumes may be obtained if available. Price lists are available upon inquiry. EARLY EOCENE BATS (hIAMMALI.4, CHIROPTERA) AND OTHER VERTEBRATES IN FRESHWATER LIhlESTONES OF THE WILLWOOD FORMATION, CLARK'S FORK BASIN, WYOMING PHILIP D. GINGERICH Abstract.-Fossil vertebrates are preserved in a variety of freshwater limestones in the Willwood Formation of the Clark's Fork Basin. Wyoming. Six limestones are described in detail. Three are interpreted to represent still water environments including the centers of hollow trees, flooded remnants of tree stumps or burrows, and shallow vegetated pans. The remaining three limestones represent a spectrum of moving water deposits ranging from a sizable perennial stream to an ephemeral stream or storm wash. Fifteen limestone lenses are weathering from a single bed or narrow stratigraphic interval at the site where vertebrate fossils were first discovered in limestones. These lenses resemble trees in size and spacing. and concentration of fractured bone, isolated teeth, and fragmentary jaws of terrestrial vertebrates within one small lens suggests Joggins-like preservation in hollow trees. The vertebrate-bearing lens at the discovery site is late Clarkforkian in age, and it includes eight species of land snails (one prosobranch and seven pulmonates), one or more salamanders, a frog, three anguimorph lizards, the oldest known heron (new genus and species), and thirteen species of mammals (including a new genus and two new species of Nyctitheriidae, a new genus and species of Palaeochiropterygidae, and a new species of Viverravidae). The palaeochiropterygiid is the oldest bat known to date. INTRODUCTION The Willwood Formation was named by F. B. Van Houten for "red-banded strata in the Big Horn Basin that in their normal position overlie the Polecat Bench, or 'Fort Union,' formation and underlie the Tatman" (Van Houten, 1944, p. 176). The "Big Horn Basin" of Van Houten included both the Clark's Fork and Bighorn drainage basins in northwestern Wyoming. These drainage basins appear to have had different histories of subsidence preceding Willwood deposition (Gingerich, 1983), and consequently the two are regarded as contiguous but distinct structural basins. The name of the formation is taken from the Willwood agricultural settlement at the base of McCullough Peaks in the northern Bighorn Basin. Van Houten (1 944, p. 178) wrote that the Willwood Formation generally consists of about 2500 feet (ca. 760 meters) of variegated shale. numerous beds of unsorted. occasionally arkosic, white and yellow sandstone. and locally abundant conglomerate lenses. He noted that small, irregular, rust-colored calcareous concretions (nodules) abound in shale layers. Development of red banding and abundant ~alcareousnodules are both attributed to soil-forming processes (Neasham and Vondra, 1972: Bown. 1979: Bown and Kraus. 198 1 a). Paleosol horizons developed in Willwood mudstones yield abundant remains of fossil vertebrates spanning the early Clarkforkian (latest Paleocene) and middle Clarkforkian (earliest Eocene) through early Bridgcrian (middle Eocene) land-mammal ages (Bown, 1979; Gingerich et al.. 1980: Winkler. 1980. 1983; Bown and Kraus, 1981b; Rose, 1981; Gingerich. 1982. 1983: Torres and Gingerich. 1983). Paleocene Fort Union limestones often occur as more or less continuous beds 1&50 cm in thickness intercalated within mudstones (Gingerich. 1969: Hickey. 1980: see also Yuretich et al., 1984). Originally gray, these weather to brown. orange. and purple as they break up when exposed at the surface. In addition, extensive but discontinuous exposures of lenticular orange-weathering calcium carbonate occur in the Fort Union Formation: these are interpreted as mineralized remains of fossil trees (Kraus. 1985). Calcium carbonate soil nodules are rare in the Fort Union Formation. In contrast, Willwood limestones rarely occur as continuous beds. More often they appear as discontinuous lenses or pods. some. again, plausibly representing mineralized remains of fossil trees. Calcium carbonate soil nodules are abundant in the Willwood Formation. The Clark's Fork Basin is unusual in that aquatic mollusks (both bivalves and gastropods) and land snails are exceptionally common and well preserved, as are calcareous hackberry seeds (Celtis sp.). Hackberry seeds calcify during growth, and their abundance in latest Clarkforkian and Wasatchian sediments of the Clark's Fork Basin may reflect growth environments rich in calcium as well as depositional environments favoring preservation of the seeds. Evidence provided by soil nodules, mollusks, and hackberry seeds indicates that calcium carbonate was present in the early Eocene environment in unusual concentrations. Carbonate nodules, mollusks, and hackberry seeds are present but notably less common in adjacent parts of the northern Bighorn Basin. Soil nodules have long been known to form around isolated bones and teeth of fossil vertebrates, but associated diverse vertebrate remains were first found in Willwood limestones of the Clark's Fork Basin in 1975. The initial discovery included a partial dentary of the plesiadapiforrn primate Cnrpolestes, a dentary of the small insectivore Leptncodon, a broken shearing premolar of the multituberculate Echpodus, and fragmentary bird remains, all preserved in a small concentration of orange-weathering limestone rubble at University of Michigan locality SC-29 (late Clarkforkian). This site has now been studied in detail, and its lithology and fauna are described below. Ongoing work in the Clark's Fork Basin indicates that early Eocene vertebrates are present in a wide variety of freshwater limestones of both Clarkforkian and Wasatchian age. The purpose of this report is, first, to describe unusual modes of preservation of fossil vertebrates in Willwood carbonates that may prove productive when investigated in detail here and elsewhere, and second, to describe the invertebrate and vertebrate fossils, including bats, found at the 1975 discovery site. All specimen numbers prefaced by the acronym UM are in the fossil vertebrate collections of the University of Michigan Museum of Paleontology. UM(1) refers to molluscan specimens numbered separately in the invertebrate collection. Fossil localities prefaced by SC are in the Sand Coulee area of the Clark's Fork Basin in northwestern Wyoming. Specimens prefaced AMNH are in the American Museum of Natural History, New York; specimens prefaced UW are in the University of Wyoming Geological Museum, Laramie. VERTEBRATE-BEARING LIMESTONES OF THE WILLWOOD FORMATION Six different occurrences of freshwater limestones yielding remains of fossil mammals and other vertebrates are described here: (1) SC-29 Discovery Site. (2) SC-29 Curpolestes Skull Site, (3)SC-29 Ectocion Site, (4) SC-289 Unionid Coquina Site. (5) SC-133 Bird Quarry Site. and (6) SC-133 Calcirudite Site. The first four of these sites are late Clarkforkian in age (Phenacodzls-EctocionZone FRESHWATER LIlIESTOYES OF WILLh'OOD FORMATION 277 of Rose, 1980. 198 1: zone Cf-3 of Gingerich. 1983). while the latter two are early Wasatchian in age (late Sandcouleean or zone Wa-2 of Gingerich. 1983). The geographic placement of University of Michigan localities prefaced SC is shown on the Clark's Fork Basin map published by Gingerich and Klitz (1985). The six sites described here represent only a small sample of the richness and variety of vertebrate-bearing early Eocene carbonates in the Willwood Formation. Middle Clarkforkian vertebrate-bearing limestones are also under study. One of these yielded specimens of Ceurholestes dolosus described by Rose and Gingerich (1987). and another limestone discovered in 1986 yielded a skull and partial skeleton of Plesiadapis cookei with associated microfauna. Other vertebrate-bearing limestones will be described as they are studied in coming years. SC-29 Discovery Site The first discovery of fossil vertebrates in Willwood Formation limestones of the Clark's Fork Basin was made by the author in 1975 when a partial dentary of Carpolestes was found in a piece of orange-weathering limestone at University of Michigan locality SC-29 (late Clarkforkian). This locality is at the head of Big Sand Coulee, and the discovery site itself (Fig. 1) is near the center of the NW 114 of Section 36, T56N, R1O1 W, Park County, Wyoming. Fragments of limestone concentrated at this site were collected and washed in an attempt to find more of the Carpolestes dentary. NO additional remains of Curpolestes were found, but close examination yielded a dentary of Leptacodon
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