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(Chiroptera: Natalidae) from the Early Miocene of Florida, with Comments on Natalid Phylogeny

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(Chiroptera: Natalidae) from the Early Miocene of Florida, with Comments on Natalid Phylogeny Journal of Mammalogy, 84(2):729±752, 2003 A NEW BAT (CHIROPTERA: NATALIDAE) FROM THE EARLY MIOCENE OF FLORIDA, WITH COMMENTS ON NATALID PHYLOGENY GARY S. MORGAN* AND NICHOLAS J. CZAPLEWSKI New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, NM 87104, USA (GSM) Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK 73072, USA (NJC) We describe a new extinct genus and species of bat belonging to the endemic Neotropical family Natalidae (Chiroptera) from the Thomas Farm Local Fauna in northern peninsular Florida of early Miocene age (18±19 million years old). The natalid sample from Thomas Farm consists of 32 fossils, including a maxillary fragment, periotics, partial dentaries, isolated teeth, humeri, and radii. A proximal radius of an indeterminate natalid is reported from the I-75 Local Fauna of early Oligocene age (about 30 million years old), also from northern Florida. These fossils from paleokarst deposits in Florida represent the 1st Tertiary records of the Natalidae. Other extinct Tertiary genera previously referred to the Natalidae, including Ageina, Chadronycteris, Chamtwaria, Honrovits, and Stehlinia, may belong to the superfamily Nataloidea but do not ®t within our restricted de®nition of this family. Eight derived characters of the Natalidae sensu stricto are discussed, 5 of which are present in the new Miocene genus. Intrafamilial phylogenetic analysis by parsimony of the Natal- idae suggests that the 3 living subgenera, Natalus (including N. major, N. stramineus, and N. tumidirostris), Chilonatalus (including C. micropus and C. tumidifrons), and Nyctiellus (including N. lepidus), deserve full generic rank. The Natalidae apparently evolved in North America before the late Oligocene, went extinct in what is now the Nearctic region (i.e., Florida) after the early Miocene, and survived in tropical Middle America during the re- mainder of the Tertiary. The presence of 2 endemic genera and 4 endemic species suggests that natalids reached the West Indies by overwater dispersal early in their history (Oligocene or Miocene). The lack of a Tertiary fossil record, marginal distribution, and limited species richness and endemism of natalids in South America are suggestive of a comparatively late arrival on that continent, possibly in the late Pliocene after the beginning of the Great American Faunal Interchange. Key words: bat, biogeography, Chiroptera, Florida, fossil, Miocene, Natalidae, Nataloidea, paleo- karst, phylogeny We describe a new extinct genus and spe- West Indies. The species described in this cies of bat in the endemic Neotropical fam- study is the 1st known extinct member of ily Natalidae from the early Miocene (early the Natalidae, excluding 5 other extinct Ter- Hemingfordian land mammal age) Thomas tiary genera (Ageina, Chadronycteris, Farm Local Fauna in northern peninsular Chamtwaria, Honrovits, and Stehlinia) that Florida. The Natalidae are a small family have been considered natalids based on the composed of 6 living species restricted to concept of the family proposed by Van Val- Middle America, South America, and the en (1979). These 5 genera are referred to the superfamily Nataloidea following Sim- * Correspondent: [email protected] 729 730 JOURNAL OF MAMMALOGY Vol. 84, No. 2 mons and Geisler (1998) but are not con- ric tons of sediment screenwashed between 1981 sidered members of the Natalidae sensu and 1985 by Ann Pratt and Arthur Poyer as part stricto. of Pratt's dissertation on the taphonomy and pa- We also report a proximal radius of a na- leoecology of the Thomas Farm vertebrate fauna talid from the Oligocene I-75 Local Fauna, (Pratt 1989, 1990; Pratt and Morgan 1989). The use of ®ne-mesh screens (1-mm opening) re- located near Gainesville, Alachua County, sulted in the recovery of large numbers of iso- Florida, about 80 km southeast of Thomas lated bat teeth that passed through the standard Farm. The I-75 site is the oldest land-ver- window screening used by earlier workers. Bat tebrate locality in Florida (Patton 1969), fossils were recovered throughout the 3-m sec- dating to the Whitneyan land mammal age tion of sediments excavated at Thomas Farm in Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 (early Oligocene, about 30 3 106 years the early 1980s, but the greatest number of bats ago). The radius from the I-75 site cannot occurred in a 1-m-thick unit of lime sand near be identi®ed below the family level but is the top of the section (unit 15; Pratt 1989, important because it represents the oldest 1990). fossil record of the Natalidae. All fossils cited are from the vertebrate pale- The discovery of a new genus and spe- ontology collection, FLMNH, University of cies of Natalidae from the early Miocene of Florida, Gainesville (UF). We follow the chirop- teran dental terminology of Legendre (1984) and Florida provides new information on the Menu and Sige (1971). We use standard abbre- historical biogeography and phylogeny of viations for tooth positions in mammals, with this small and enigmatic group of New uppercase letters for upper teeth and lowercase World bats. The identi®cation of a natalid letters for lower teeth: I and i (upper and lower nearly 20 million years older than any other incisors), C and c (upper and lower canines), P known member of this family prompted us and p (upper and lower premolars), and M and to examine the phylogenetic relationships m (upper and lower molars). The terminology of all species within the Natalidae. Al- for chiropteran postcranial elements follows though our primary aim was to determine Smith (1972) and Vaughan (1959). All measure- the systematic position of the new Florida ments of fossils are in millimeters. Miocene natalid, we also examined the re- We compared the Thomas Farm natalid fossils lationships among the 6 extant species of with at least 2 skulls and 2 complete skeletons natalids because no formal hypothesis of re- of each of the 6 extant species of Natalidae and the potentially related families Furipteridae, lationships has been proposed for the fam- Thyropteridae, and Vespertilionidae (see Appen- ily. dix I for comparative material examined). Methods of phylogenetic analysis.ÐThere is MATERIALS AND METHODS no previous intrafamilial phylogenetic analysis In the late 1950s and early 1960s, Clayton for the Natalidae. We developed a hypothesis of Ray of the Florida State Museum (now the Flor- relationships based on parsimony analysis (using ida Museum of Natural History [FLMNH]) and PAUP 4.0b10 softwareÐSwofford 2000) of the Pierce Brodkorb of the University of Florida De- Floridian fossils and all 5 extant species gener- partment of Zoology began screenwashing sed- ally recognized as members of the Natalidae iments from Thomas Farm for small vertebrates. (Koopman 1993, 1994; Nowak 1994; Appendix These early screenwashing efforts led to system- I) plus Natalus major, which we recognize as a atic studies of frogs (Holman 1965, 1967), sal- valid species following Morgan (1989b). Mor- amanders and lizards (Estes 1963), snakes (Auf- phological comparisons disclosed 50 characters fenberg 1963), birds (Brodkorb 1956), and ro- that may be phylogenetically informative (Ap- dents (Black 1963). Bat fossils also were recov- pendix II). We scored 24 cranial, dental, and hu- ered during this early screenwashing program, meral characters that are present in the fragmen- but early ®nds did not include specimens of the tary fossils as well as in extant natalids (Appen- new natalid. The majority of bat fossils from dix II, characters 1±24). The remaining 26 cra- Thomas Farm, including all natalid specimens nial and postcranial characters were available described here, were collected from about 2 met- only in the extant natalids (with missing values May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 731 coded for the fossil taxon). Characters and char- acter states are based on Dalquest (1950), Good- win (1959), Koopman (1994), Miller (1907), Ot- tenwalder and Genoways (1982), Silva Taboada (1979), Simmons (1998), Simmons and Geisler (1998), Van Valen (1979), and our personal ob- servations. There has been controversy surrounding the higher-level relationships of the Natalidae. In 1 study by Simmons and Geisler (1998), the Na- talidae were part of a clade (Nataloidea) that Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 also included the Furipteridae, Thyropteridae, and Myzopodidae. In their analysis (Simmons and Geisler 1998), this clade was sister to the Vespertilionoidea (Vespertilionidae and Molos- sidae) and farther removed from the Noctilion- oidea. In a contrasting analysis based on mito- chondrial DNA, Van Den Bussche and Hoofer (2001) found the nataloids (except Myzopodi- FIG. 1.ÐLocation of the 2 sites in Florida that dae) strongly associated with the Noctilionoidea have produced Tertiary fossils of Natalidae, I-75 (Noctilionidae, Mystacinidae, Mormoopidae, (Oligocene) and Thomas Farm (early Miocene). and Phyllostomidae) and far removed from the Vespertilionidae and Molossidae. More recently, cated 12 km north of Bell in Gilchrist County in using additional evidence from nuclear genes, northern peninsular Florida (Fig. 1), has pro- Hoofer et al. (in press) reexamined yangochirop- duced the best known early Miocene (early teran bats, including all the aforementioned fam- Hemingfordian land mammal age) vertebrate ilies. In their analysis Nataloidea was found to fauna in eastern North America. There are de- be polyphyletic; Natalidae clustered with Ves- tailed accounts of the discovery, excavation, ge- pertilionidae and Molossidae, whereas Furipter- ology, and stratigraphy of Thomas Farm (Auf- idae and Thyropteridae clustered with the Noc- fenberg 1963; Pratt 1989, 1990; Simpson 1932; tilionoidea. Based in part on the analysis of White 1942), as well as faunal lists and bibli- Hoofer et al. (in press) and in part on the tra- ographies (Olsen 1962; Ray 1957; Webb 1981).
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