VOL. 60, NO. 3, PP. 133-234 BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY August 23, 2023 UNIVERSITY OF FLORIDA GAINESVILLE The FLORIDA MUSEUM OF NATURAL HISTORY is Florida’s state museum of natural history, dedicated to understanding, preserving, and interpreting biological diversity and cultural heritage. The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is an on-line, open-ac- cess, peer-reviewed journal that publishes results of original research in zoology, botany, paleontology, archaeology, and museum science. Multi-author issues of related papers have been published together, and inquiries about putting together such issues are welcomed. Address all inquiries to the Editor of the Bulletin. The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online regis- tration system for the ICZN. The ZooBank Publication number for this issue is D72FD99C-656E-4FF5- 9813-2F1C71A53B9. Jon Bloch, Editor for this issue Bulletin Committee Michal Kowalewski Michelle J. LeFebvre Jacqueline Miller Roger W. Portell Jonathan I. Bloch, Ex officio Member ISSN: 2373-9991 Copyright © 2023 by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, and personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum. Publication Date: August 23, 2023 This and other issues of the Bulletin can be freely downloaded at: https://www.floridamuseum.ufl.edu/bulletin/publications/ Send communications about this publication to: Editor of the Bulletin; Florida Museum of Natural History; University of Florida; P.O. Box 117800; Gainesville, FL 32611-7800 USA FAX: 352-846-0287; Email: bulletin@flmnh.ufl.edu URL: https://www.floridamuseum.ufl.edu/bulletin/home/ Cover image: Top row, left, upper molars of Oligopteryx floridanus; center, Brooksville 2 Quarry, late Oli- gocene, Hernando County, Florida; right, lower molars of Floridopteryx poyeri. Bottom row, left, Thomas Farm site, early Miocene, Gilchrist County, Florida, left to right, Nicholas Czaplewski, Arthur Poyer, and Gary Morgan; right, Rhynchonycteris naso (proboscis bats) hanging under the eaves of a building in Costa Rica. mailto:bulletin@flmnh.ufl.edu https://www.floridamuseum.ufl.edu/bulletin/home/ NEW BATS IN THE TROPICAL FAMILY EMBALLONURIDAE (MAMMALIA: CHIROPTERA) FROM THE OLIGOCENE AND EARLY MIOCENE OF FLORIDA Gary S. Morgan1 and Nicholas J. Czaplewski2 ABSTRACT Three new genera and four new species in the tropical bat family Emballonuridae (Mammalia: Chirop- tera) are described from the Oligocene and early Miocene of peninsular Florida. These are: a new genus Oligopteryx and two new species, O. floridanus and O. hamaxitos, from the late Oligocene (early Arika- reean) Brooksville 2 Local Fauna in Hernando County, with referred specimens of each species from the early Oligocene (Whitneyan) I-75 Local Fauna in Alachua County; a new genus and species, Karstop- teryx gunnelli, from the latest Oligocene (late Arikareean) Buda Local Fauna in Alachua County; and a new genus and species, Floridopteryx poyeri, from the early Miocene (early Hemingfordian) Thomas Farm Local Fauna in Gilchrist County. Characters that distinguish Oligopteryx from other emballonurid genera are: M1 with a much reduced parastylar region and a large talon with a triangular posterolingual extension; M1 and M2 with a prominent hypocone separated from the protocone by a deep, V-shaped notch in the postprotocrista, and deeply concave talon basin; small, single-rooted p3; m1 and m2 with the trigonid narrower and shorter than the talonid, the paraconid and metaconid located close together along the lingual margin, and the entocristid blade-like, V-shaped, and oriented labially. O. hamaxitos is dis- tinguished from O. floridanus by its smaller size and well-developed paraloph and metaloph on M1 and M2. Karstopteryx gunnelli is separated from the two species of Oligopteryx by the prominent parastyle, anteriorly oriented preparacrista, and rounded talon. Floridopteryx poyeri is characterized by: M1 with a highly reduced parastylar region but with a well-developed parastyle, short preparacrista, small hypocone not separated from the protocone by a notch in the postprotocrista, and a squarish talon; m1/m2 with the talonid much broader than the trigonid, trigonid compressed, shorter than talonid, and with the paraconid and metaconid close together along the lingual margin, large hypoconulid, and anterior root flattened and compressed anteroposteriorly (m1 only). Oligopteryx and Karstopteryx are considered members of the subfamily Emballonurinae based on the reduced parastylar region of the M1 but are left unassigned at the tribal level. Oligopteryx differs from all living emballonurids in the presence of a p3. Floridopteryx has a reduced parastylar region on M1 and is also considered an emballonurine, but can be separated from Oligopteryx by the presence of a prominent parastyle and lack of a p3. Floridopteryx is placed in the monophyletic New World emballonurine tribe Diclidurini. Taphonomy indicates Oligopteryx was a colonial cave dweller, whereas Floridopteryx more likely roosted in trees, as do most living Neotropical emballonurids. A change from a tropical or subtropical climate in Florida in the Oligocene and early Mio- cene to a warm temperate climate from the middle Miocene to the present is reflected in the disappearance of emballonurids from Florida after the early Miocene. Molecular studies indicate the Emballonuridae dispersed from Africa to South America in the Oligocene, despite the oldest Western Hemisphere record of this family from the early Oligocene (~30 Ma) of Florida (Oligopteryx). We hypothesize emballonurids may have reached North America from Eurasia in the Eocene using an overland route and then dispersed overwater to the then-island continent of South America, with the oldest South American record of that family from the late Oligocene (~25 Ma) of Peru. 1New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, NM 87104, USA 2Oklahoma Museum of Natural History, University of Oklahoma, 2401 Chautauqua Avenue, Norman, OK 73072-7029, USA RESUMEN Se describen tres nuevos géneros y cuatro nuevas especies de la familia de murciélagos tropicales Embal- lonuridae del Oligoceno y el Mioceno temprano de la peninsula del estado de Florida. Estos son: un nuevo género Oligopteryx y dos nuevas especies, O. floridanus y O. hamaxitos, del Oligoceno tardío (edad Arikareense temprano) fauna local Brooksville 2 en el condado de Hernando, con especímenes referidos de cada especie de la fauna local I-75 del Oligoceno temprano (edad Whitneyense) en el condado de Ala- chua, y un nuevo género y especie, Karstopteryx gunnelli, del Oligoceno ultimo (edad Arikareense tardío) de la fauna local Buda en el condado de Alachua; y un nuevo género y especie, Floridopteryx poyeri, de la fauna local Thomas Farm del Mioceno temprano (edad Hemingfordiense temprano) en el condado de Gilchrist. Los caracteres que distinguen a Oligopteryx de otros géneros embalonúridos son: M1 con una region parastylar muy reducida y un talón de forma triangular con una extension posterolingual; M1 y M2 con un hipocono prominente separado del protocono por una muesca profunda en forma de V y una cuenca en talón profundamente cóncava; p3 pequeño, de raíz única; m1 y m2 con el trigónido más corto que el talónido, el paracónido y el metacónido ubicados muy juntos a lo largo del margen lingual, y el entocrístido en forma de cuchilla, enforma de V y orientado labialmente. Oligopteryx floridanus se distingue de O. hamaxitos principalmente por su mayor tamaño. Karstopteryx gunnelli está separado de las dos especies de Oligopteryx por el parastilo prominente, el preparastilo orientado anteriormente, y la talón redondeada. Floridopteryx se caracteriza por: M1 con una region parastilar muy reducida pero con un parastilo bien desarollado, preparacrista corto, pequeña hipocono no separado del protocono por una muesca, y un talón cuadrada; m1 / m2 con el talónido mucho más ancho que el trigónido, el trigónido comprimido, más corto que el talónido y con el paracónido y el metacónido muy juntos a lo largo del margen lingual, hipoconulido grande y raíz anterior aplanada y comprimida anterioposteriormente (solo m1). Oligopteryx y Karstopteryx se consideran miembros de la subfamilia Emballonurinae basada en la region parastilar reducida de M1, pero no se asignan a nivel tribal. Oligopteryx difiere de todos los em- balonúridos vivos en presencia de un p3. Floridopteryx también tiene una region parastilar reducida en M1 y también se considera una Emballonurinae, pero puede separarse de Oligopteryx por la presencia de un parastilo prominente y la falta de una p3. Floridopteryx se coloca en la tribu Diclidurini monofilética del hemisferio occidental. La tafonomía indica que Oligopteryx era un habitante colonial de las cavernas, mientras que Floridopteryx probablemente descansaban en los árboles, al igual que la mayoría de los em- balonúridos neotropicales vivos. Un cambio de un clima tropical / subtropical en Florida en el Oligoceno y Mioceno temprano a un clima templado cálido desde el Mioceno medio hasta el presente se refleja en la desaparición de los embalonúridos de Florida después del Mioceno temprano. Los estudios moleculares indican que los Emballonuridae se dispersaron desde África hasta América del Sur en el Oligoceno, a pesar del registro más antiguo del hemisferio occidental de esta familia desde el Oligoceno temprano (~30 Ma) de Florida (Oligopteryx). Hipotetizamos que los embalónuridos pueden haber llegado a América del Norte desde Eurasia en el Eoceno utilizando una ruta terrestre y luego disperarse sobre el agua hasta el continente de América del Sur, entonces insular, con el registro más antiguo de América del Sur de esa familia desde finales del Oligoceno (~25 Ma) Perú. Key words: Chiroptera, Emballonuridae, Florida, fossil, Oligocene, Miocene, paleoecology, biogeography, Neotropical TABLE OF CONTENTS Introduction ...........................................................................................................136 Methods and Materials ..........................................................................................136 Geologic Setting and Chronology .........................................................................137 Florida Fossil Sites Containing Emballonuridae ..................................................139 I-75 ...............................................................................................................139 Brooksville 2 ................................................................................................141 Buda .............................................................................................................144 Thomas Farm ...............................................................................................144 Systematic Paleontology .......................................................................................146 Oligopteryx floridanus new genus and species ..................................................147 Oligopteryx hamaxitos new species .................................................................178 Karstopteryx gunnelli new genus and species ...................................................186 Floridopteryx poyeri new genus and species ....................................................189 Discussion .............................................................................................................197 Systematic Relationships ................................................................................197 Taphonomy and Paleoecology .........................................................................200 Biogeography and Evolutionary History of the Emballonuridae ........................205 Brief Evolutionary History of New World Noctilionoidea, Molossidae, and Vespertilionidae ..............................................................................................210 Tertiary Fossil Record of the Neotropical Chiroptera: An Update ...............................212 Acknowledgments ...................................................................................................222 Literature Cited .......................................................................................................223 Appendix 1. Modern Comparative Specimens Examined Appendix 2. Characters of the dentition and dentary of Emballonuridae Morgan, G., S. and N. J. Czaplewski, 2023. New bats in the tropical family Emballonuridae (Mammalia: Chiroptera) from the Oligocene and early Miocene of Florida. Bulletin of the Florida Museum of Natural History 60(3):133–234. https://doi.org/10.58782/flmnh.wefq4531 https://doi.org/10.58782/flmnh.wefq4531 136 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) INTRODUCTION Much of what we know about the evolutionary his- tory of bats in eastern North America during the middle of the Cenozoic (between about 30 and 18 Ma) is derived from three paleokarst deposits in northern peninsular Florida: the early Oligocene (Whitneyan North American Land Mammal Age– NALMA, ~30 Ma) I-75 Local Fauna (LF), the late Oligocene (early Arikareean NALMA, ~26– 28 Ma) Brooksville 2 LF, and the early Miocene (early Hemingfordian NALMA, ~18 Ma) Thomas Farm LF. These three sites document a significant Paleogene-Neogene transition in Florida chiropter- an assemblages (Morgan and Czaplewski, 2012), from an Oligocene fauna characterized by families now restricted to the Neotropics (Emballonuridae and Mormoopidae) to an early Miocene fauna that primarily consists of Vespertilionidae but also in- cludes taxa with Neotropical affinities (Emballon- uridae, Natalidae, Molossidae). With a few excep- tions, most middle Miocene and younger fossil bats from Florida and elsewhere in North America belong to the Vespertilionidae (Czaplewski et al., 2008; Morgan and Czaplewski, 2012), the predom- inant family in the modern temperate North Ameri- can chiropteran fauna. The Emballonuridae, the sheath-tailed or sac-winged bats, occur worldwide in tropical re- gions, including Mexico, Central America, and South America, as well as Africa, Southeast Asia, Australia, and many Pacific islands (Simmons, 2005). Considering the modern pantropical distri- bution of emballonurids, it is rather surprising that the oldest fossil record of this group in the New World is from the early Oligocene of peninsular Florida, a region with a warm temperate climate at present that no longer supports members of this family. New World Tertiary records of emballon- urids are limited to eight sites, four from Florida, the Oligocene I-75, Brooksville 2, and Buda LFs, and the early Miocene Thomas Farm LF (Morgan and Czaplewski, 2012; this paper), and four locali- ties from South America, the middle Miocene La Venta Fauna from Colombia (Czaplewski, 1997; Czaplewski et al., 2003b) and three sites from Contamana, Peru, a late Oligocene fauna from the Chambira Formation, and early Miocene and late Miocene faunas from the Pebas Formation (An- toine et al., 2016). Three new genera and four new species of Oligocene and early Miocene emballonurids from Florida are described here and compared to fossil and living emballonurids from the Neotropical Re- gion and the Old World. The paleoecology of the Florida fossil emballonurids indicates a tropical or subtropical climate in the Florida peninsula during the Oligocene, transitioning into a warm temperate climate in the early Miocene. Taphonomy suggests the Florida Oligocene emballonurids were cave- dwelling bats which, together with a previously described extinct genus and species in the family Mormoopidae from the same fossil sites (Morgan et al., 2019), represent the earliest cavernicolous members of the Chiroptera documented in the Western Hemisphere. The single species of Flor- ida Miocene emballonurid was more likely a tree dweller like most modern Neotropical members of the family. Finally, we present a brief review of the pre-Pleistocene fossil record of the Neotropical Chiroptera from South America, as well as fossils from Oligocene and Miocene sites in Florida with Neotropical affinities. METHODS AND MATERIALS Chiropteran dental terminology follows Czaplews- ki et al. (2008) and Ravel et al. (2016). The ab- breviations for teeth in mammals are standard, with upper case letters for upper teeth and lower case letters for lower teeth: I/i (upper/lower inci- sors), C/c (upper/lower canines), P/p (upper/lower premolars), and M/m (upper/lower molars). Tooth positions are identified by numbers. For example, P4 is an upper fourth premolar and m3 is a lower third molar. The identity of the anteriormost lower premolar in bats that possess three lower premo- lars, such as Oligopteryx, is controversial. Most authors have followed Miller (1907) in considering p1 as the missing lower premolar in bats, with the remaining premolars being p2, p3, and p4. Howev- er, Thomas (1908), Simmons and Conway (2001), MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 137 and Giannini and Simmons (2007) regarded the first lower premolar as the p1 and considered the p2 to be the missing lower premolar in the Chirop- tera. We follow Miller (1907), Hand et al. (2015), and Cirranello et al. (2016) in recognizing the three lower premolars in bats as p2, p3, p4, with p1 miss- ing. In describing a petrosal, we use the terminol- ogy of Staněk (1933), Henson (1970), Simmons and Geisler (1998), and Giannini et al. (2006). The terminology for chiropteran postcranial elements follows Vaughan (1959), Smith (1972), Czaplews- ki et al. (2008), and Gaudioso et al. (2020). We estimated body mass in grams of the extinct taxa of emballonurid bats using equations generated by (Gunnell et al., 2009). All other measurements are in mm. We compared the I-75, Brooksville 2, Buda, and Thomas Farm emballonurid fossils to teeth, dentaries, and postcranial skeletons of at least one species in each of the 14 genera of extant Embal- lonuridae (list of modern comparative material examined in Appendix 1). We also compared the Florida Cenozoic emballonurids to five extinct genera of emballonurids from Europe and Africa, Afrillonura, Dhofarella, Pseudovespertiliavus, Ta- chypteron, and Vespertiliavus, primarily from the literature. Most of our morphological comparisons of the Florida fossil emballonurids are at the ge- neric level. All Tertiary sites from Florida that have pro- duced bats were collected using standard screen- washing techniques for microvertebrates. Before the mid 1970s, screens finer than standard window screen (16 mesh, 1.5 mm opening) were not in gen- eral use, and thus isolated teeth of small bats may have been lost from sites screenwashed during this time period (e.g., I-75, Buda, early Thomas Farm collections). All Florida sites collected from the early 1980s to the present, including later Thomas Farm collections and Brooksville 2, were screen- washed through both standard window screen and fine mesh brass screen (24 mesh, 1.0 mm opening). After 1980, no identifable bat specimens were lost from Florida Tertiary sites during the screenwash- ing process. The I-75, Brooksville 2, Buda, and Thomas Farm emballonurid fossil specimens are from the vertebrate paleontology collection of the Florida Museum of Natural History, University of Florida, Gainesville (UF). Modern comparative skeletons of Emballonuridae are from the mammal collec- tions of the American Museum of Natural History, New York (AMNH); Museum of Southwestern Bi- ology, Department of Biology, University of New Mexico, Albuquerque (MSB); Oklahoma Museum of Natural History, University of Oklahoma, Nor- man (OMNH); Florida Museum of Natural History, University of Florida (UF-M); and U. S. National Museum of Natural History, Smithsonian Institu- tion, Washington, DC (USNM). Other abbrevia- tions are: LF (Local Fauna); Ma (Mega-annum or millions of years); MNI (minimum number of indi- viduals); NISP (number of identifiable specimens); NALMA (North American land mammal age); SALMA (South American land mammal age). The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature (ICZN), and hence the new genus and species names contained herein are available under that Code. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank Publication number for this issue is: D72FD99C-656E-4FF5- 9813-2F1C71A53B9. GEOLOGIC SETTING AND CHRONOLOGY Prior to the Oligocene, the region that is now the Florida peninsula was entirely submerged, consist- ing of shallow tropical reefs and carbonate banks. Consequently, much of northern peninsular Flor- ida is now underlain by Eocene and Oligocene marine limestones. Sometime in the latter part of the early Oligocene (Rupelian) about 30 million years ago (Ma), Florida first emerged above sea level based on the presence of the I-75 land ver- tebrate fauna correlated with the Whitneyan NA- LMA (30–32 Ma). Throughout the remainder of the Cenozoic, the Florida peninsula fluctuated be- tween submerged and emergent, depending upon changes in relative sea level. During this time pe- 138 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) riod, the highly soluble Paleogene limestones of northern Florida were acted upon by both chemi- cal and physical erosional processes to develop one of the most extensive karst terrains in North America (Lane, 1986; Florea, 2008). From a pale- ontological standpoint, the most significant aspect of the north Florida karst geomorphic province is the abundance of terrestrial vertebrate deposits oc- curring in current or former caves, fissures, sink- holes, and other paleokarst features (Morgan and Hulbert, 2008). The limestones in northern Florida are mined commercially, which has resulted in the discovery of most of the fossil deposits. More than 150 vertebrate fossil sites of karst origin are known from the Florida peninsula, including many depos- its that contain bats (Morgan and Hulbert, 2008; Morgan and Czaplewski, 2012). Nowhere else in North America are karst-derived vertebrate fossil deposits of Cenozoic age found in such abundance. Florida vertebrate faunas of karst origin range in age from the Oligocene through the Pleistocene, with the oldest being the early Oligocene I-75 LF and the largest concentration of sites from the late Pleistocene (Rancholabrean NALMA). The common occurrence of bat fossils in karst deposits from northern peninsular Florida strongly indicates that many of these sites represent the remnants of former cave systems. The rich- est pre-Pleistocene bat faunas from Florida (I-75, Brooksville 2, Thomas Farm) were derived from sediment-filled fissures or sinkholes in Paleogene limestones. In addition to bats, these karst depos- its often contain large samples of small, non-volant mammals such as marsupials, lipotyphlans (‘in- sectivores’), rodents, and lagomorphs, as well as other small vertebrates, including frogs, toads, sal- amanders, lizards, snakes, and birds. Despite their original deposition in caves, pre-Pleistocene karst faunas in Florida do not occur in what are currently caves. The same physical and chemical erosional processes that originally formed the caves eventu- ally destroyed them. With the collapse of the caves, their contained sediments became buried, and the caves were no longer accessible from surface en- trances. We suspect that only a small percentage of these buried karst deposits have been uncovered within the past century or so by limestone mining operations, as heavy equipment exposed deeply buried clay and sand deposits containing fossils. This process of cave formation and destruction ap- pears to have occurred fairly rapidly in a geologic sense, because all vertebrate fossils so far recov- ered from surficial deposits in Florida caves are late Pleistocene (Rancholabrean) in age. There are numerous karst-derived vertebrate faunas of early to middle Pleistocene age (Blancan and Irving- tonian NALMAs) from peninsular Florida, all of which are from previously buried cave or sinkhole deposits (Morgan and Hulbert, 2008). Because Florida karst deposits usually con- sist of isolated pockets of sediment that cannot be directly correlated stratigraphically, their signifi- cance for biochronology has often been overlooked or downplayed. However, the abundance in many Florida sites of small mammals and age-diagnostic large mammals, including carnivores, horses, and artiodactyls, often allows precise biochronologic comparisons with faunas of similar age in west- ern North America that have associated geochro- nological data such as radioisotopic dates or geo- magnetic polarity stratigraphy (e.g., Frailey, 1978, 1979; Pratt and Morgan, 1989; Morgan, 1993; Al- bright, 1998; Hayes, 2000, 2005; MacFadden and Morgan, 2003; Tedford et al., 2004; Albright et al., 2008; Morgan and Hulbert, 2008; Czaplewski and Morgan, 2015). Moreover, Florida faunas are often more diverse and have larger and better- preserved fossil samples than faunas of equivalent age in western North America. Most Florida Ce- nozoic vertebrate assemblages are ‘local faunas’ in the sense of Tedford (1970), with the follow- ing definition from Woodburne (2004, xiii): “Lo- cal Fauna: An aggregate of fossil vertebrate spe- cies that have a limited distribution in time from a number of closely grouped localities in a limited geographic area…A local fauna could be based on taxa from a single locality.” Indeed, most local faunas from Florida have been described based on the vertebrate fossils from a single locality, includ- ing I-75 and Thomas Farm (Patton, 1969a; Pratt, 1989, 1990). Brooksville 2 and Buda are derived from several separate karst sediment pockets in a MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 139 limited geographic area, specifically a limestone quarry (Frailey, 1979; Hayes, 2000). Since Florida Cenozoic local faunas mostly occur in a single lo- cality and in a narrow stratigraphic interval, there are minimal complications from ecological mixing and time averaging. Vertebrate local faunas or fau- nas from western North America often occur over a considerably larger geographic area and through a substantial stratigraphic interval and, as such, time averaging and ecological mixing can become sub- stantial error factors in evaluating the biochronol- ogy and/or paleoecology of these sites. Our biochronology follows the North Ameri- can land mammal ages (NALMA), which are bio- chronologic units based on mammalian evolution in North America, specifically an association of fossil mammals that represents a particular interval of geologic time. Most NALMA are defined based on a specific association of mammalian genera. For the faunal definitions and age of the NALMAs rep- resented in Florida Tertiary paleokarst deposits that contain emballonurid bats, we follow Prothero and Emry (2004) for the Whitneyan NALMA (early Oligocene) and Tedford et al. (2004) for the Ari- kareean and Hemingfordian NALMAs (late Oligo- cene and early Miocene). We also incorporate the updated Arikareean biostratigraphy of Albright et al. (2008). FLORIDA FOSSIL SITES CONTAINING EMBALLONURIDAE I-75 The I-75 Local Fauna (LF) was discovered in 1965 during the construction of Interstate High- way 75 (‘I-75’) through the southwestern edge of Gainesville, Alachua County, in northern pen- insular Florida (Fig. 1). The site was located in a roadcut at the southwest corner of the intersection of I-75 and Florida State Route 121 but was de- stroyed by road building activities shortly after its discovery. Mervin Kontrovitz of the University of Florida initially discovered the I-75 site, which was collected by Florida State Museum (now Florida Museum of Natural History) field crews. The fos- siliferous sediments in the I-75 site consisted of massive, slightly sandy, dark brown to black clays, deposited in a small karst solution feature 5 m in diameter and 2 m deep, developed in Eocene ma- rine limestone (Patton, 1969a). Considering the small size of the fossiliferous deposit, the I-75 site has a diverse vertebrate fau- na composed of about 45 species. Patton (1969a) published a preliminary vertebrate faunal list and Holman (1999) and Holman and Harrison (2000, 2001) described the snake fauna. Hayes (2000) updated the mammalian faunal list from Patton (1969a). The herpetofauna includes anurans (both bufonids and scaphiopodids), an indeterminate sirenid salamander, a small land tortoise, a pond turtle, the lizard Peltosaurus, and an impressive snake fauna consisting of nine species of boids and colubrids (Holman, 1999; Holman and Harrison, 2000, 2001). Inexplicably, birds are absent from the I-75 LF. The fauna of large mammals consists of (from Patton, 1969a; Hayes, 2000): two species of carnivorans, the amphicyonid Daphoenus and the small mustelid Palaeogale; the equid Miohip- pus sp.; a tayassuid; two oreodonts; and the small leptomerycid artiodactyl Leptomeryx sp., the most Figure 1. Outline map of the southeastern United States showing Florida, with locations of Oligocene (I-75, Brooksville 2, and Buda) and Miocene (Thomas Farm) sites containing fossils of Emballonuridae. 140 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) abundant mammal in the fauna. Small mammals include: the marsupial Herpetotherium cf. merria- mi (Hayes, 2005), the large lipotyphlan Centetodon cf. wolffi; the lagomorph Palaeolagus sp.; four spe- cies of rodents, Eutypomys sp., a heteromyid, and two eomyids; as well as seven species of bats (de- scribed in more detail below). Marine vertebrates also were recovered from I-75, including sharks, rays, and bony fish (Patton, 1969a; Tessman, 1969). Because of the association of marine, freshwater, and terrestrial taxa, Patton (1969a) suggested there may have been some re- working of the I-75 deposit. In most other Florida Tertiary sites containing a mixture of marine and nonmarine vertebrates, the marine vertebrates far outnumber the terrestrial component of the fauna, and most teeth of small mammals are isolated and show some evidence of transport and water wear (Morgan, 1993). However, terrestrial vertebrates dominate the I-75 fauna and there are several par- tial mandibles and maxillae with teeth represent- ing bats and other small mammals. Furthermore, most of the isolated teeth and limb bones are well preserved and show little evidence of water wear. The most plausible explanation would seem to be that the terrestrial vertebrates from the I-75 site were deposited subaerially in a small cave or fis- sure. Shortly thereafter, with a rise in sea level the marine vertebrates were deposited, with minor re- working. Florida had only recently emerged above sea level in the late early Oligocene and probably consisted of a short, narrow peninsula of low relief. A slight rise in sea level would have been sufficient to inundate most of the land present in Florida at that time, including the cave/fissure represented by the I-75 site. I-75 is the oldest land vertebrate fauna known from Florida, referred to the Whitneyan NALMA by Patton (1969a) and most subsequent workers (Savage and Russell, 1983; Emry et al., 1987; Pro- thero and Emry, 2004). The Whitneyan represents a restricted interval of time in the early Oligocene (~30-32 Ma), known primarily by faunas from the Northern Great Plains in Nebraska, South Dakota, and Wyoming. The following is a brief discussion of the most biochronologically diagnostic land mammals present in the I-75 LF. Four isolated teeth of a brachydont horse from I-75 are identified as Miohippus sp., although Patton (1969a) referred these teeth to Mesohippus. These teeth compare more closely with Miohippus than Mesohippus on the basis of the characters discussed by Prothero and Shubin (1989), including larger size and well- developed hypostyles on the upper molars. Miohip- pus occurs from the Chadronian through the early Arikareean but is most typical of Whitneyan and Arikareean faunas (Emry et al., 1987; Prothero and Shubin, 1989). The I-75 Miohippus teeth appear to be most similar in size and morphology to M. inter- medius from the late Whitneyan of South Dakota. A large sample of isolated teeth is tentatively referred to the small leptomerycid artiodactyl Leptomeryx, although Patton (1969a) and Hayes (2005) referred these teeth to Nanotragulus. Leptomeryx occurs from the Chadronian through the early Arikareean, whereas Nanotragulus first appears in the early Arikareean (Prothero and Emry, 2004; Tedford et al., 2004). The geolabidid lipotyphlan Centetodon is represented in the I-75 LF by a partial upper molar and three complete lower molars. The morphology and size of the I-75 Centetodon teeth are similar to C. wolffi from the Orellan and Whitneyan of South Dakota, the largest species in the genus (Lil- legraven et al., 1981). Hayes (2005) identified two upper molars from I-75 as the marsupial Herpeto- therium cf. merriami, originally described from the early Arikareean John Day Formation of Oregon. A single upper cheektooth of a small leporid is tentatively referred to Palaeolagus, the most com- mon rabbit in Orellan and Whitneyan faunas. Pal- aeolagus went extinct in the early Arikareean. The large beaver-like rodent Eutypomys is represented by about 20 teeth from the I-75 fauna. Eutypomys is typical of Whitneyan and Orellan faunas in the Great Plains, but survived into the early Arikareean (Macdonald, 1963, 1970; Tedford et al., 1996; Pro- thero and Emry, 2004). The presence of Centetodon, Palaeolagus, Eutypomys Miohippus, and Leptomeryx establishes an age of early late Arikareean or older (older than 24 Ma) for the I-75 LF, as none of these genera are MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 141 known to survive into the latest Arikareean (Ar4; Lillegraven et al., 1981; Tedford et al., 1996, 2004; Albright et al., 2008). All five of these genera also occur in both the Orellan and Whitneyan NALMAs. A post-Orellan age is suggested by the presence of an advanced species of Miohippus near M. inter- medius, and an age no younger than Whitneyan is indicated by the occurence of Centetodon wolffi. The absence of mammals clearly indicative of the Arikareean (e.g., Nanotragulus) favors a Whit- neyan age, probably late Whitneyan (~30-31 Ma; Prothero and Emry, 2004; Albright et al., 2008), which is several million years older than the late early Arikareean (Ar2; ~26-28 Ma) Brooksville 2 LF. The similarity of bats from I-75 and Brooks- ville 2 suggests these two faunas are fairly close in age. Although their chiropteran faunas are similar, the remainder of the mammalian assemblages from these two sites have few genera and no species in common (Hayes, 2000). I-75 provides one of the earliest records of the taxonomic diversity and community structure of North American Late Paleogene chiropteran faunas. About 40 specimens representing seven species of bats are known from I-75, including: Oligopteryx floridanus and O. hamaxitos (de- scribed in this paper), the oldest members of the Emballonuridae in the New World; Koopmanycte- ris palaeomormoops, the oldest known member of the Mormoopidae (Morgan et al., 2019); Speonyc- teris aurantiadens and S. naturalis (type locality) in the extinct noctilionoid family Speonycteridae (Czaplewski and Morgan, 2012); an indeterminate genus and species representing the oldest known member of the Natalidae (Morgan and Czaplewski, 2003); and a large indeterminate genus and spe- cies of Vespertilionidae (Morgan and Czaplewski, 2012). The most abundant bats in the I-75 fauna are Oligopteryx floridanus and Koopmanycteris palaeomormoops. BrooksvIlle 2 The Brooksville 2 site was discovered in 1994 in an abandoned limerock quarry of the Flori- da Rock Industries company, about 8 km northwest of Brooksville, Hernando County, central Florida (Fig. 1). Brooksville 2 consists of clays and sands filling five small karst solution features in the ma- rine lower Oligocene Suwannee Limestone (Hayes, 2000). See field photos of the Brooksville 2 Quarry in Figure 2. FLMNH crews visited the Brooksville Quarry on numerous field trips between April 1994 and February 1995. Although some specimens of larger vertebrates were found on the surface, the fossils were collected primarily by screenwash- ing because of the abundance of microvertebrates. Approximately 500 kg of sediments were scre- enwashed from solution features in the quarry in which the Brooksville 2 site was located (Hayes, 2000). The vertebrate assemblage from Brooksville 2 is composed predominantly of small terrestrial vertebrates, including frogs, lizards, snakes, and a diverse fauna of small mammals, but no birds. Mead (2013) reported three vertebrae of a tiny sco- lecophidian snake from Brooksville 2. Larger mam- mals are represented primarily by isolated teeth. Hayes (2000) reported 27 species of mammals from Brooksville 2. He described the lipotyphlans, carnivorans, lagomorphs, and selected taxa of ro- dents, and briefly reviewed the ungulates. Hayes (2005) identified the marsupial Herpetotherium fu- gax from Brooksville 2. The large sample of bats is described in more detail below. The ungulates and carnivorans include (Hayes, 2000): the horse Miohippus; a phenacocoeline oreodont; the camel Nothokemas waldropi; the tiny artiodactyl Nano- tragulus loomisi; and six species of carnivorans, Palaeogale minuta, the musteloids Acheronictis webbi (type locality) and Arikarictis chapini (type locality), and the canids Enhydrocyon cf. pahinsin- tewakpa, Osbornodon wangi (type locality), and Phlaocyon taylori (type locality). In addition to the small didelphid marsupial Herpetotherium fugax and five species of bats, the small mammal fauna also includes: two lipotyphlans, the geolabidid Centetodon magnus and the erinaceid Parvericius montanus; the lagomorph Megalagus abaconis (type locality); and numerous rodents, including sciurids, eomyids, two heteromyids, the castorid Agnotocastor sp., and an undescribed entoptychine geomyoid (Hayes, 2000, 2005; this paper). 142 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 2. Field photographs of the late Oligocene (early Arikareean) Brooksville 2 site (as it appeared in the mid 1990s), located in an abandoned limerock mine northwest of Brooksville, Hernando County, Florida. The Brooksville 2 Local Fauna is the type locality of the emballonurid bats Oligopteryx floridanus and O. hamaxitos. A. Overview of the Brooksville Quarry showing exposures of the marine lower Oligocene Suwannee Limestone. B. View of the quarry wall showing a karst deposit (indicated by black arrow), within the Suwannee Limestone. C. Close-up view of the same karst deposit shown in B. D. Even closer view of the same karst deposit in B and C, showing the fossiliferous laminated red and gray clays and fine sands (photos courtesy of Glynn Hayes). MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 143 Hayes (2000) determined the age of the Brooksville 2 LF through biochronologic com- parisons with other faunas from Florida and the Gulf Coast representing the Arikareean NALMA (Albright, 1998), and with Arikareean faunas from western North America. Preservation of the Brooksville 2 fossils in isolated karst solution fea- tures precludes direct stratigraphic correlation with described Oligocene and Miocene stratigraphic units from northern Florida. We follow Albright et al. (2008) for the updated calibration and subdivi- sions of the Arikareean NALMA. The maximum age of Brooksville 2 is constrained by the presence of the erinaceid Parvericius and entoptychine ro- dents, both of which appear at the beginning of the late early Arikareean (Ar2, ~28 Ma; Tedford et al., 1996, 2004; Albright et al., 2008). The minimum age of the site is restricted to early late Arikareean (Ar3) by the occurrence of Miohippus. Albright et al. (2008) recorded the range of Miohippus as ex- tending into the early late Arikareean in the John Formation of Oregon, disappearing at about 24 Ma. Hayes (2000) placed the Brooksville 2 LF in the ‘medial’ Arikareean (late Oligocene; between 24 and 28 Ma). With more recent changes in the boundaries of the subdivisions within the Arika- reean (Albright et al., 2008), the age range Hayes (2000) suggested for Brooksville 2 would now place this fauna in either the late early Arikareean (Ar2) or the early late Arikareean (Ar3). The co-oc- currence of Parvericius and entoptychine rodents with Miohippus, together with the strong similarity between the chiropteran faunas from Brooksville 2 and the Whitneyan I-75 LF, supports a late early Arikareean age (~26-28 Ma) for the Brooksville 2 LF. An analysis of the mammalian biochronology of Brooksville 2 at the species level, rather than the generic level, would probably yield a more precise age for this fauna. The chiropteran sample from Brooksville 2 consists of about 200 fossils representing five spe- cies: the emballonurids Oligopteryx floridanus and O. hamaxitos (both described in this paper), with Brooksville 2 as the type locality for both species; the mormoopid Koopmanycteris palaeomormoops (type locality); Speonycteris aurantiadens (type locality) in the extinct family Speonycteridae; and an indeterminate genus and species of molossid (Czaplewski et al., 2003a; Czaplewski and Mor- gan, 2012; Morgan and Czaplewski, 2012; Morgan et al., 2019). With the exception of the molossid, all of these species are shared with I-75. Brooks- ville 2 has the second largest bat sample from any Florida Tertiary site, after Thomas Farm, and the third most diverse bat fauna after Thomas Farm and I-75. As with the I-75 LF, the most common bats from Brooksville 2 are two species from fami- lies currently restricted to the tropics, the embal- lonurid Oligopteryx floridanus and the mormoopid Koopmanycteris palaeomormoops. It seems rather remarkable that the chirop- teran fauna from Brooksville 2 shares four of its five species with I-75, Koopmanycteris palaeo- mormoops, Oligopteryx floridanus, O. hamaxitos, and Speonycteris aurantiadens, whereas the non- volant mammalian faunas from these two sites have no species in common and share only three genera (Centetodon, Herpetotherium, and Mio- hippus). Both the Brooksville 2 and I-75 sites are fissure deposits in which the fossils were origi- nally deposited in caves, which is supported by the abundance of cave-dwelling bats, including the two species of Emballonuridae described here, Oligopteryx floridanus and O. hamaxitos, and a previously described member of the Mormoopi- dae, Koopmanycteris palaeomormoops (Morgan et al., 2019). The similar chiropteran faunas in these two sites primarily consist of species with tropi- cal affinities (Emballonuridae, Mormoopidae), in- dicating Florida had a tropical/subtropical climate during their deposition. Apparently, the chiropteran fauna was well-adapted to the climate of peninsu- lar Florida in the Oligocene and underwent only minimal evolutionary change over a period of sev- eral million years. The total lack of overlap at the species level between the non-volant mammalian faunas from Brooksville 2 and I-75 is indicative of both evolutionary changes within genera (e.g., Centetodon, Herpetotherium, and Miohippus) and overall faunal change associated with the age dif- ference between Whitneyan and early Arikareean faunas (e.g., the small ruminant Leptomeryx and 144 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) eutypomyine rodents in I-75 compared to the tiny ruminant Nanotragulus, entoptychine rodents, and erinaceid Parvericius in Brooksville 2). The differ- ences between the non-volant mammalian faunas in these two sites appear to be primarily related to their difference in age, with I-75 about 2 million years older. Buda The Buda Quarry is an abandoned lime- stone mine located about 8 km southwest of High Springs, Alachua County, northern peninsular Florida (Fig. 1). The fossiliferous deposit, long since destroyed by mining operations, consisted of three shallow, clay-filled fissures from 1 to 3 m in diameter that probably shared a common open- ing, eroded into Eocene marine limestone. Frailey (1979) reported 12 taxa of large mammals from the Buda LF: five carnivorans, the amphicyonid Da- phoenodon notionastes, the canids Bassariscops achoros and Cynarctoides sp., a mustelid, and a nimravid; two perissodactyls, the small chalicoth- ere Moropus sp. and an indeterminate equid; and five artiodactyls, a small tayassuid, a phenacocoe- line oreodont, two camelids, and the tiny ruminant Nanotragulus loomisi, the most abundant mammal in the fauna. Wang et al. (1999) restudied the Buda canid fauna, transferring Bassariscops achoros to the genus Phlaocyon, referring Cynarctoides to the species C. lemur, and identifying the species Cormocyon cf. copei. The Buda LF also has a sig- nificant small mammal fauna that remains mostly unstudied. Rich and Patton (1975) reported the eri- naceid Amphechinus from Buda, later reidentified as Parvericius by Hayes (2000). Czaplewski and Morgan (2015) described the last surviving apate- myid (Apatotheria) Sinclairella simplicidens from Buda. Other small mammals include the geolabi- did lipotyphlan Centetodon cf. magnus, and three rodents, the jimomyid Texomys sp., the eomyid Arikareeomys sp., and a heteromyid, as well as the emballonurid bat Karstopteryx gunnelli described here (Hayes, 2000; Czaplewski and Morgan, 2015). Frailey (1979) regarded the Buda LF as late Arikareean, and Albright (1998) and Hayes (2000) placed this fauna in the early late Arikareean (Ar3). The most compelling evidence for a late Arika- reean age is the presence of the amphicyonid Da- phoenodon, one of the defining genera for the late Arikareean, and the small chalicothere Moropus cf. oregonensis (Coombs et al., 2001; Tedford et al., 2004). According to Albright et al. (2008), Da- phoenodon and Moropus first appeared in the John Day Formation in Oregon in the early late Arika- reean at about 25 Ma (early Ar3). However, several other mammals from Buda are more typical of early Arikareean faunas (Ar1 and/or Ar2), including Cy- narctoides lemur and Cormocyon cf. copei (Wang et al., 1999; Albright et al., 2008). Two small mam- mal genera, Centetodon and Arikareeomys, are also more typical of the early Arikareean but oc- cur in the early late Arikareean. We follow Albright (1998), Hayes (2000), and Czaplewski and Morgan (2015) in placing the Buda LF in the early late Ari- kareean (early Ar3; ~24-26 Ma; latest Oligocene), based on the co-occurrence of Daphoenodon and Moropus that first appear in the Ar3, together with Centetodon and Arikareeomys that last occur in the Ar3. The Buda LF chiropteran fauna consists of a single tooth (M1), described below as a new genus and species, Karstopteryx gunnelli. We suspect the sparse bat sample from Buda may be due to a col- lecting bias, because only minimal screenwashing for microvertebrates was conducted at this site and nothing finer than window screen (1.5 mm open- ing) was used. Dental differences between Kar- stopteryx and the two species of Oligopteryx from the somewhat older Brooksville 2 LF confirm that significant morphological changes occurred in the Florida emballonurid lineage in the late Oligocene, supporting the younger early late Arikareean age of the Buda LF proposed by other authors (Frai- ley 1979; Hayes, 2000; Czaplewski and Morgan, 2015). Thomas Farm The Thomas Farm site is located 12 km north of Bell in Gilchrist County, northern penin- sular Florida (Fig. 1). Thomas Farm has produced the best known and most diverse early Miocene (early Hemingfordian NALMA; He1) vertebrate MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 145 fauna in eastern North America. The site consists of clays and sands filling a sinkhole developed in Eocene marine limestone. See field photos of the Thomas Farm site in Figure 3. Pratt (1989, 1990) concluded that Thomas Farm consisted of a 30 m deep, vertical-walled sinkhole surrounded by a for- ested habitat. There is a large sample of bat fossils in a 1 m thick lime sand near the top of the section that is also rich in other small vertebrates. Most of the small vertebrates appear to have been depos- ited in a former cave system developed in the wall of the sinkhole, either as a coprocoenosis derived from the scat of small mammalian carnivores or the pellets of raptorial birds, or through the natural ac- cumulation of carcasses on a cave floor in the case of the bats (Pratt, 1989). The Thomas Farm LF consists of more than 90 species of vertebrates (Webb, 1981; Pratt, 1989, 1990), primarily composed of terrestrial forms but also including a freshwater component of frogs, salamanders, pond turtles, alligators, and aquatic birds. Thomas Farm has 23 species of large mammals, the most abundant of which is the horse Parahippus leonensis (Hulbert, 1984), and Figure 3. Field photographs of the early Miocene (early Hemingfordian) Thomas Farm Local Fauna, north of Bell, Gilchrist County, Florida. Thomas Farm is the type locality of the emballonurid bat Floridopteryx poyeri. A. View of a portion of the Thomas Farm site in the early 1980s showing the meter square grid system used to plot the location of larger fossils; from left to right, Steve Emslie, Ann Pratt, and Gary Morgan. B. Excavating sediment for screenwashing in 2001; from left to right, Nick Czaplewski, Art Poyer, and Gary Morgan. C. Close-up of a sample of the limesand layer, the richest sediment at Thomas Farm for bat fossils; from left to right, Nick Czaplewski (with sediment sample in hand), Art Poyer, and Gary Morgan. D. Wall of meter square showing thin layers of limesand sediment. Photos are courtesy of Richard Hulbert (A) and Erika Simons (B-D). 146 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) a diverse small vertebrate fauna of nearly 70 spe- cies, including frogs, toads, salamanders, lizards, snakes, birds, bats, shrews, and rodents. Small mammals include (Pratt, 1989; Pratt and Morgan, 1989): the soricid Limnoecus sp.; nine species of bats (see below); three sciurid rodents, Petaurist- odon pattersoni (type locality), Nototamias hulber- ti (type locality), and cf. Miospermophilus sp.; two heteromyid rodents, Proheteromys floridanus and P. magnus; and the eomyid rodent Pseudotherido- mys sp. The biochronology of artiodactyls (Patton, 1969b), equids (Forstén, 1975; Hulbert and Mac- Fadden, 1991), carnivorans (Tedford and Frailey, 1976; Wang et al., 1999), and sciurids (Pratt and Morgan, 1989) indicates a late early Hemingford- ian age for Thomas Farm (late He1; ~17.5–18 Ma). The early Hemingfordian is defined in part by the first appearance of several genera of Eurasian im- migrant carnivorans, including the amphicyonid Amphicyon, ursid Hemicyon (= Phoberocyon), and mustelid Leptarctus, as well as the first occurrence of the camelid Floridatragulus and the large fly- ing squirrel Petauristodon, and the last occurrence of the amphicyonid Cynelos and the rhinocerotid Menoceras (Tedford and Frailey, 1976; Tedford et al., 1987, 2004; Pratt and Morgan, 1989). These genera are all present at Thomas Farm. Amphicyon and the equids Anchitherium, Archaeohippus, and Parahippus from Thomas Farm are very similar to species of these same genera from early Heming- fordian faunas in the Runningwater Formation of Nebraska. Other correlative early Hemingfordian faunas from the western United States are the Flint Hill LF of South Dakota and the Martin Canyon LF of Colorado (Tedford et al., 1987, 2004). Thomas Farm has the largest bat sample from any Tertiary fossil deposit in North America, with more than 3,000 specimens (3,180 bat specimens as of 17 November 2022; UF/FLMNH vertebrate paleontology database). Most of the bat fossils con- sist of isolated teeth or ends of limb bones, but there are also numerous mandibles, maxillary fragments, and complete limb bones. The Thomas Farm chi- ropteran fauna is composed of at least nine species, including four species belonging to families with Neotropical affinities: one species of Emballon- uridae, Floridopteryx poyeri, described here; one species of Natalidae, Primonatalus prattae, with Thomas Farm as the type locality (Morgan and Czaplewski, 2003); and two undescribed species of Molossidae similar to Tadarida (Czaplewski et al., 2003a). The other five species belong to the Ves- pertilionidae, three of which have been described, with Thomas Farm as the type locality, Miomyo- tis floridanus and Suaptenos whitei (Lawrence, 1943) and Karstala silva (Czaplewski and Morgan, 2000). Two other undescribed species of vespertil- ionids occur in the Thomas Farm LF based on dif- ferences in the morphology of the distal humerus with the three described vespertilionids from the site. One of these species is similar to tree bats of the genus Lasiurus and a second species is similar to big-eared bats of the genus Corynorhinus. Ves- pertilionids dominate the chiropteran fauna, with more than 75% of all bats belonging to Suaptenos whitei. SYSTEMATIC PALEONTOLOGY Order CHIROPTERA Blumenbach, 1779 Family EMBALLONURIDAE Gervais, 1855 OLIGOPTERYX new genus Type Species. —Oligopteryx floridanus sp. nov. Included Species.—Type species and Oli- gopteryx hamaxitos. Diagnosis.—The m1 and m2 are nyctalo- dont, with the talonid significantly broader than the trigonid, paraconid and metaconid very close to one another, metaconid located anterior to the protoconid, postcristid at about a 45° angle to the long axis of tooth, entocristid blade-like and dis- tinctly V-shaped with the apex of V oriented labi- ally, and with a triangular-shaped process that proj- ects posteroventrally from the posterior end of the mandibular symphysis below p4. M1 with a greatly reduced parastylar region, parastyle absent or very small, preparacrista very short, oriented posteriorly and running parallel to the postparacrista, parac- ingulum well developed, talon triangular in shape with a prominent posterolingual extension, metac- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 147 ingulum with a deep notch just posterior and lin- gual to the base of the metacone; in the M1 and M2 the talon basin is deeply concave, and in lingual view there is a deep V-shaped notch in the post- protocrista separating the protocone and the promi- nent, triangular hypocone. Etymology.—Oligo (Greek), few, scanty, in reference to the Oligocene epoch, and pteryx (Greek), wing, a common ending for generic names in the family Emballonuridae. OLIGOPTERYX FLORIDANUS new species Fig. 4-10 Holotype.—UF 157769, right dentary frag- ment with p2, p4-m1, Brooksville 2 Local Fauna, late Oligocene (late early Arikareean), Hernando County, Florida. Paratypes.—UF 157784, RM1; UF 182780, RM2; UF 182773, RM3 in maxillary fragment; UF 157790, left dentary fragment with m1-m2, p4 broken off at base of crown, and alveolus for p3; UF 157771, left dentary fragment with m2- m3; UF 179958, edentulous dentary with nearly complete horizonal ramus from anterior tip to m2 and all alveoli from c1-m2; UF 179904, left proxi- mal humerus; UF 179964, left distal humerus; UF 179911, right proximal radius. All paratypes are from the Brooksville 2 Local Fauna, late Oligocene (late early Arikareean), Hernando County, Florida. Referred Specimens.—Brooksville 2 Local Fauna.–Left M1 (7): UF 157776, 157779, 157785, 182777, 182779, 182858, 182896; right M1 (4): UF 157773, 157778, 182787, 182866; left M2 (2): UF 157780; 182781; right M2 (2): UF 157777, 182873; left M3 (6): UF 157775, 179991, 182774, 182775, 182778, 182899; right M3 (3): UF 157781, 182776, 182898; UF 157772, 157787, right dentary frag- ments with m3; UF 157789, right dentary fragment with m2; UF 182855, right dentary fragment with p4 and alveolus with root of p3; UF 182874, right dentary fragment with m2 and alveoli for p4 and m1; UF 157788, left dentary fragment with m2; left m1/m2 (9): UF 157782, 182814, 182861, 182862, 182864, 182868, 182871, 182891, 182893; right m1/m2 (13): UF 156289, 157783, 179987, 182809, 182811, 182813, 182819, 182859, 182860, 182865, 182867, 182870, 182892; UF 182872, left m3; right m3 (2): UF 182857, 182895; left C1: UF 182884; right c1: UF 182802; left P4: UF 182907, 182783; left proximal humerus (5): UF 179935, 179959, 179961, 179962, 179984; right proximal humerus (3): UF 179903, 179936, 179963; left distal humerus (5): UF 179909, 179965-179968; right distal humerus (2): UF 179910, 179939; left proximal radius (3): UF 179942, 179969, 179971; right proximal radius (6) UF 179912, 179913, 179940, 179972, 179981, 179982; left distal radius (2), UF 209956, 209957; right distal radius (4), UF 179926, 179944, 179974, 179975; left proximal femur, UF 182788. The minimum number of indi- viduals (MNI) in the Brooksville 2 sample of Oli- gopteryx floridanus is seven based on the presence of seven left M1s and seven proximal ends of the right radius. The number of identifiable specimens (NISP) is 106. I-75 Local Fauna.—UF 121701, 121702, right M1; UF 121704, left M2; UF 121703, right M2; UF 121724, partial right dentary with poste- rior half of p4 and alveoli for c1, p3, m1-m2; UF 16882, edentulous left dentary with alveoli for m1- m3; UF 121725, edentulous left dentary with alve- oli for p3-m2; UF 121707, 121708, left m1/m2 (2); UF 121705, 121706, right m1/m2 (2); UF 121710, right distal humerus; UF 121711, 121712, left proximal radius (2). The MNI in the I-75 sample of Oligopteryx floridanus is two individuals based on the presence of two right M1s, two partial left dentaries, and two proximal ends of the left radius. The NISP is 14. Type Locality and Age.—Brooksville 2 Lo- cal Fauna, late Oligocene (late early Arikareean; Ar3), Hernando County, Florida. Occurrence.—Known only from the early Oligocene (Whitneyan) I-75 LF, Alachua County, Florida and the late Oligocene (early Arikareean) Brooksville 2 LF, Hernando County, Florida. Etymology.—floridanus; in reference to the state of Florida, where all known specimens of this species have been collected. Diagnosis.—Same as for the genus, with cer- tain additional characters, mostly consisting of fea- 148 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) tures that cannot be observed in the smaller species Oligopteryx hamaxitos because of the more limited sample of the latter (e.g., no lower premolars and M3/m3 of O. hamaxitos are known). The p2 is sin- gle-rooted, comparatively large, diamond-shaped, laterally compressed, and has a blade-like central cusp; the p3 is present, tiny, single-rooted, and lo- cated along the lingual margin of the mandibular toothrow; the p4 is tall, almost caniniform, with a conical central cusp, and triangular occlusal outline; the m3 with the talonid about the same breadth as the trigonid or slightly narrower, the cristid obliqua meets the trigonid near the labial base of the meta- conid; the mandibular symphysis of the dentary is narrow anteriorly; M3 with a prominent hooked parastyle, metacone bulbous, posteriorly oriented, and more labially placed than the paracone, lacks a postmetacrista, metastyle, and hypocone, paraloph present. morphologIcal descrIpTIons Upper dentition.–Among both fossil and re- cent emballonurids, the M1 is one of the most di- agnostic teeth. The sample of M1s of Oligopteryx floridanus from Brooksville 2 consists of 11 com- plete teeth (7 left, 4 right; measurements in Table 1). The following description is primarily based on UF 157784 (paratype; Fig. 4 E-F), a right M1 from Brooksville 2. However, the entire sample of M1s was used to evaluate dental variation. The overall shape of the M1 is shorter anteroposteri- orly and more transversely elongated than in most other emballonurids. The M1 of the Florida fossils is somewhat rectangular with the long axis oriented Figure 4. Right upper molars of Oligopteryx floridanus from Brooksville 2 LF (A-F) and Karstopteryx gunnelli from Buda LF (G-H). All molars in top row are occlusal views (A, C, E, G), all molars in bottom row are lingual views (B, D, F, H). A-B, Oligopteryx floridanus, UF 182773, M3 (paratype); C-D, O. floridanus, UF 182780, M2 (paratype); E-F, O. floridanus, UF 157784, M1 (paratype); G-H, Karstopteryx gunnelli, UF 97386, M1 (holotype). MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 149 Table 1. Measurements of the upper molars of Oligocene and Miocene Emballonuridae from Florida, including: Oligopteryx floridanus and O. hamaxitos from the Oligocene I-75 LF (Whitneyan) and Brooksville 2 LF (early Arikareean), Karstopteryx gunnelli from the late Oligocene Buda LF (late Arikareean), and Floridopteryx poyeri from the early Miocene Thomas Farm LF (early Hemingfordian). All measurements are in mm. Missing measurement indicated by “—”. Abbreviations for sample statistics: N (number of specimens); M (mean); OR (observed range). Statistics are calculated if there are three or more speci- mens for a particular tooth position. Species, tooth position, anteroposterior transverse fauna, and catalog length width number Oligopteryx floridanus M1 Brooksville 2 UF 157773 1.82 2.42 UF 157776 1.75 2.37 UF 157778 1.90 2.45 UF 157779 1.77 2.17 UF 157784 (paratype) 1.92 2.50 UF 157785 1.85 2.47 UF 182777 1.77 2.22 UF 182779 1.55 2.40 UF 182787 2.15 2.42 UF 182896 1.65 2.37 I-75 UF 121701 1.65 2.42 N 11 11 M 1.80 2.38 OR 1.55–2.15 2.17–2.50 M2 Brooksville 2 UF 157777 1.77 2.57 UF 157780 1.77 2.72 UF 182780 (paratype) 1.80 2.47 UF 182873 1.67 2.45 I-75 UF 121703 1.70 2.42 UF 121704 1.72 2.62 N 6 6 M 1.74 2.54 OR 1.67–1.80 2.42–2.72 M3 Brooksville 2 UF 157775 1.17 2.37 UF 157781 0.95 2.12 UF 179991 1.10 2.35 UF 182773 (paratype) 1.22 2.22 UF 182774 1.27 2.42 UF 182778 1.05 2.37 UF 182898 1.00 2.22 UF 182899 1.12 2.27 N 8 8 M 1.11 2.29 OR 0.95–1.27 2.12–2.42 Oligopteryx hamaxitos M1 Brooksville 2 UF 182808 (paratype) 1.37 1.65 M2 Brooksville 2 UF 157774 1.22 1.57 UF 157786 (paratype) 1.35 1.85 Karstopteryx gunnelli M1 Buda UF 97386 (holotype) 1.75 2.27 Floridopteryx poyeri M1 Thomas Farm UF 121134 0.85 – Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 150 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) labiolingually, while in most other emballonurids the M1 is squarish. The anterolabial region of the M1 labial to the paracone is highly reduced, almost lacking in some specimens. A parastyle is absent in most M1s from Brooksville but is present in sev- eral specimens consisting of a tiny cuspule at the anterolabial termination of the paracingulum (= precingulum). The only portion of the tooth pres- ent anterior to the mesostyle consists of the short preparacrista, paracone, and the postparacrista; the parastylar shelf labial to these structures is virtually absent. The parastylar region of the M1 is more re- duced in Oligopteryx than in any living emballon- urid. There is some variation in the length of the preparacrista in the sample of M1s from Brooks- ville, ranging from very short to moderate in length. In those teeth with a somewhat longer preparacrista (from a third to half the length of the postparacris- ta), this crest is oriented posteriorly, almost parallel to the postparacrista. In several M1s, the labialmost extension of the preparacrista is distinctly curved posteriorly. Posterior to the mesostyle, there is a shallow V-shaped notch in the metafossa labial to the metacone. The paracingulum is relatively broad, extending from the anterolabial base of the protocone to about the middle of the paracone, end- ing abruptly just anterior to the labial termination of the preparacrista. The protocone is located on the anterolingual margin and is oriented anteriorly, whereas the paracone and metacone are more verti- cal. About halfway between the protocone and the hypocone, there is a shallow but distinct triangular- shaped notch in the postprotocrista separating the two cusps. The hypocone is about half the height of the protocone and consists of a distinct low, rounded cusp. A weak paraloph extends from the tip of the protocone to the base of the paracone. A metaloph is absent. The trigon basin has a rather shallow, somewhat elliptical-shaped pit in its cen- ter that is deeper than the remainder of the basin but does not have sharply defined edges. The trigon basin is not nearly as deep or ‘pocketed’ as in some other emballonurids. The talon basin is deeper and better defined than the trigon basin, with the deepest portion posterior to the base of the meta- cone. The posterolabial portion of the talon basin is the posteriormost portion of the tooth, extend- ing slightly farther posteriorly than the metastyle. The talon constitutes the posterolingual extension of the tooth, consisting of a well-developed, trian- Table 1. (Cont.) Species, tooth position, anteroposterior transverse fauna, and catalog length width number Oligopteryx floridanus M1 Brooksville 2 UF 157773 1.82 2.42 UF 157776 1.75 2.37 UF 157778 1.90 2.45 UF 157779 1.77 2.17 UF 157784 (paratype) 1.92 2.50 UF 157785 1.85 2.47 UF 182777 1.77 2.22 UF 182779 1.55 2.40 UF 182787 2.15 2.42 UF 182896 1.65 2.37 I-75 UF 121701 1.65 2.42 N 11 11 M 1.80 2.38 OR 1.55–2.15 2.17–2.50 M2 Brooksville 2 UF 157777 1.77 2.57 UF 157780 1.77 2.72 UF 182780 (paratype) 1.80 2.47 UF 182873 1.67 2.45 I-75 UF 121703 1.70 2.42 UF 121704 1.72 2.62 N 6 6 M 1.74 2.54 OR 1.67–1.80 2.42–2.72 M3 Brooksville 2 UF 157775 1.17 2.37 UF 157781 0.95 2.12 UF 179991 1.10 2.35 UF 182773 (paratype) 1.22 2.22 UF 182774 1.27 2.42 UF 182778 1.05 2.37 UF 182898 1.00 2.22 UF 182899 1.12 2.27 N 8 8 M 1.11 2.29 OR 0.95–1.27 2.12–2.42 Oligopteryx hamaxitos M1 Brooksville 2 UF 182808 (paratype) 1.37 1.65 M2 Brooksville 2 UF 157774 1.22 1.57 UF 157786 (paratype) 1.35 1.85 Karstopteryx gunnelli M1 Buda UF 97386 (holotype) 1.75 2.27 Floridopteryx poyeri M1 Thomas Farm UF 121134 0.85 – Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Species, tooth position, anteroposterior transverse fauna, and catalog length width number Oligopteryx floridanus M1 Brooksville 2 UF 157773 1.82 2.42 UF 157776 1.75 2.37 UF 157778 1.90 2.45 UF 157779 1.77 2.17 UF 157784 (paratype) 1.92 2.50 UF 157785 1.85 2.47 UF 182777 1.77 2.22 UF 182779 1.55 2.40 UF 182787 2.15 2.42 UF 182896 1.65 2.37 I-75 UF 121701 1.65 2.42 N 11 11 M 1.80 2.38 OR 1.55–2.15 2.17–2.50 M2 Brooksville 2 UF 157777 1.77 2.57 UF 157780 1.77 2.72 UF 182780 (paratype) 1.80 2.47 UF 182873 1.67 2.45 I-75 UF 121703 1.70 2.42 UF 121704 1.72 2.62 N 6 6 M 1.74 2.54 OR 1.67–1.80 2.42–2.72 M3 Brooksville 2 UF 157775 1.17 2.37 UF 157781 0.95 2.12 UF 179991 1.10 2.35 UF 182773 (paratype) 1.22 2.22 UF 182774 1.27 2.42 UF 182778 1.05 2.37 UF 182898 1.00 2.22 UF 182899 1.12 2.27 N 8 8 M 1.11 2.29 OR 0.95–1.27 2.12–2.42 Oligopteryx hamaxitos M1 Brooksville 2 UF 182808 (paratype) 1.37 1.65 M2 Brooksville 2 UF 157774 1.22 1.57 UF 157786 (paratype) 1.35 1.85 Karstopteryx gunnelli M1 Buda UF 97386 (holotype) 1.75 2.27 Floridopteryx poyeri M1 Thomas Farm UF 121134 0.85 – Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 151 gular-shaped process, ranging from sharply trian- gular to more broadly or bluntly triangular in some specimens. The narrow metacingulum (= postcin- gulum) extends from the base of the metacone to the metastyle. A deep, V-shaped notch separates the metacingulum from the more posteriorly and lingually placed talon basin. A single right M1 of a large emballon- urid from I-75 (UF 121701) is very similar to the Brooksville M1s. This tooth fits in the middle of the range of variation in most dental characters and size compared to the Brooksville sample (measure- ments in Table 1). A parastyle is absent, although there is a slight anterior bulge at the anterolabial termination of the paracingulum. The preparacrista is short, about one-third the length of and parallel to the postprotocrista. There is a very shallow in- dentation in the metafossa labial to the metacone. A weak paraloph is present and the metaloph is ab- sent. There is a deep, V-shaped notch in the post- protocrista separating the protocone from the well- developed hypocone. The deeply concave talon ba- sin extends posteriorly to the level of the metastyle. There is a V-shaped indentation in the metacingu- lum at the posterior base of the metacone. There are seven M2s from Brooksville 2 (4 left, 3 right; measurements in Table 1). The follow- ing description is primarily based on UF 182780 (paratype; Fig. 4 C-D), a right M2. However, the entire sample of M2s was examined to evaluate dental variation. Unlike M1, the M2 of Oligopter- yx floridanus has a well-developed parastyle that is hooked or curved. The preparacrista is also much better developed and longer than on M1, connect- ing the paracone to the parastyle. The paracingu- lum extends along the anterior margin of the tooth from the base of the protocone to the tip of the parastyle. There is a rather deep V-shaped notch in the labial margin of the parafossa labial to the paracone and a shallow notch in the metafossa la- bial to the metacone. The protocone and hypocone are separated by a rather deep, V-shaped notch in the postprotocrista. The hypocone is prominent and is positioned slightly more lingually than the pro- tocone. A weak but distinct paraloph connects the protocone to the base of the paracone. A metaloph is absent in most specimens, but a weak metaloph connecting the posterior edge of the protocone to the base of the metacone is present in UF 157777. The trigon basin is relatively shallow for an em- ballonurid. The talon basin has a deep pit between the base of the metacone and the metacingulum. The talon is more squared off than in the M1 in which the talon is triangular. The metacingulum has a notch posterior to the base of the metacone, although this notch is not as deep as in the M1. Two M2s of large emballonurids from I-75 (UF 121703, 121704) easily fit within the range of variation of M2s of Oligopteryx floridanus from Brooksville 2 (measurements in Table 1). The I-75 M2s have a strong, hooked parastyle. The paracin- gulum is broad and well developed. There is a fairly deep V-shaped notch in the parafossa labial to the paracone and a weak indentation in the metafossa labial to the metacone. There is a V-shaped notch between the protocone and hypocone. The hypo- cone is lingually offset. A weak paraloph is present, but no metaloph. The talon basin is deeply concave and the talon has a square posterior margin. There are ten M3s from Brooksville 2 (6 left, 4 right; measurements in Table 1). The following description is primarily based on UF 182773 (para- type; Fig. 4 A-B), a right M3. The M3 of Oligop- teryx is not as reduced as in some other emballon- urids, in particular taphozoines that have a highly reduced M3. The only structures on the ectoloph of M3 that are lacking compared to the M2 are the postmetacrista and metastyle. There is a prominent hooked parastyle. A well-developed paracingulum extends from the base of the protocone to the para- style. The paracone and metacone are about the same size and height, although the metacone is ori- ented slightly posteriorly. The paracone is sharply triangular in shape, whereas the metacone is more inflated or bulbous. The metacone is slightly more labially placed than the paracone, whereas the me- sostyle is about a third of the distance between the parastyle and paracone, closer to the parastyle. By comparison, the mesostyle and parastyle are lo- cated at about the same level near the labial mar- gin on M2. The premetacrista is noticeably shorter than the postparacrista because of the more labial 152 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) position of the metacone. By comparison, the post- paracrista and premetacrista are approximately the same length on M2. The metacone extends farther posteriorly than the posterior margin of the trigon basin. The talon and its basin are absent in M3, whereas the talon is the posteriormost portion of the tooth on M1 and M2. The protocone is low and rounded, located about midway along the lingual margin. A fairly strong paraloph extends from the protocone to the base of the paracone. A hypocone is absent, as is a metaloph. The trigon basin is rath- er shallow. Isolated chiropteran canines are difficult to identify, and we have based our identification on a combination of size and similarity in morphology with other New World emballonurids. A C1 (UF 182884; Fig. 5 A-C) from the Brooksville 2 LF re- sembles the C1s of many other emballonurids in having anterior and posterior secondary cusps. It is larger than C1s referred to Oligopteryx hamaxitos described below. The secondary cusps are miss- ing enamel and thus appear relatively weak, with the anterior one merged into the anterior crest of the main cusp, while the posterior one is separate yet connected with the posterior crest of the main cusp. In life, these secondary cusps would have been more prominent. The tooth lacks a labial cin- gulum and has a relatively weak lingual cingulum. The crown is rather hemiconical and labiolingually narrow, longer than wide, with a concave lingual face and a convex labial face. The root angles back from the crown base, giving a procumbent orienta- tion to the tooth. We refer two isolated P4s (UF 182783 and 182907; Fig. 6) from the Brooksville 2 local fauna to Oligopteryx floridanus. The P4s are three-root- ed, with one root each supporting the central cusp (protocone), the posterolabial crest and style, and the lingual lobe. UF 182907 has a curved postero- labial crest, while UF 182783 has a slightly more sinuous posterolabial crest (but the latter might be due to greater wear in UF 182907). Both P4s have a relatively small lingual lobe (talon) with a shal- low talon basin surrounded by a prominent cingu- lum. The anterior cingulum surrounding the base of the protocone is separated from the talon by an Figure 5. Canines referred to Oligopteryx floridanus from Brooksville 2 LF. A-C, UF 182884, left C1 in occlusal (A), lingual (B), and labial (C) views. D-F, UF 182802, right c1 in labial (D), lingual (E), and occlusal (F) views. anterolingual notch, with anterolingual cingular cusps on either side of the notch. UF 182783 has a small swelling along the labial margin that is ab- sent in UF 182907. Lower Dentition.–None of the dentary frag- ments of Oligopteryx floridanus from Brooksville 2 or I-75 contains the canine, although these sites have a significant sample of isolated canines in the MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 153 size range to belong to this species. An edentulous dentary from Brooksville 2 with alveoli for most of the lower teeth (UF 179958, paratype) is the only specimen in the sample that contains the alveolus for the lower canine. The c1 assigned to Oligopty- eryx floridanus (UF 182802; Fig. 5 D-F) from the Brooksville 2 LF has a tall, narrow main cusp that is aligned with the root but appears strongly canted forward relative to the basal cingulum and cingular cusps. The cingulum is absent labially but strong lingually, with prominent secondary cusps at the anterolingual and posterior ends. The base of the tooth is anteroposteriorly elongated and transverse- ly narrowed, and the root is much more robust than the main cusp. The main cusp bears a longitudinal crest on its posterolingual face that does not con- nect with the lingual cingulum. The tooth is larger than the c1s of O. hamaxitos described below. The presence of three lower premolars in Oligopteryx is the primitive condition in bats. The Eocene emballonurids Tachypteron and Vespertil- iavus also have three lower premolars. However, extant emballonurids possess only two lower pre- molars; the intermediate premolar (p3) is absent. As discussed above under Methods and Materials, we identify the anteriormost lower premolar in bats that possess three lower premolars, including Oli- gopteryx, as the p2. We follow Miller (1907), Hand et al. (2015b), and Ciranello et al. (2016) in recog- nizing the three lower premolars in bats as p2, p3, p4, with p1 missing. There are two specimens of the p2 of Oligopteryx floridanus, one is preserved in the holotype dentary (UF 157769; Fig. 7 A-C) and the second is an isolated left p2. The p2 is rela- tively large and single-rooted, and laterally com- pressed with a large, blade-like central cusp, here regarded as the protoconid. It is sharply triangular in labial and lingual views and diamond-shaped in occlusal view. There is a weak lingual cingulum. The occlusal area is similar to that of p4, but in lat- eral view the protoconid of p2 is noticeably shorter. The posterior margin of p2 and anterior margin of p4 are in contact in the holotype, the only specimen that preserves both teeth. Three dentaries of Oligopteryx floridanus from Brooksville 2 (UF 157790, 179958 [Fig. 8], 182855) and two from I-75 (UF 121723, 121724) preserve the tiny, round alveolus of a single-rooted p3, located along the lingual margin of the toothrow, wedged between the posterior edge of the alveolus for p2 and the anterior alveolus of p4. Several of these specimens have the root of p3, but none pre- serve the tooth crown. In one dentary (UF 182855), the p3 alveolus causes a slight indentation in the posterolingual margin of the p2 alveolus. No living emballonurid possesses a p3, although this tooth is present in the extinct Eocene genera Tachypteron Figure 6. Left P4s referred to Oligopteryx floridanus from Brooksville 2 LF. A-C, UF 182907, left P4 in occlusal (A), lingual (B), and labial (C) views; D-F, UF 182783, left P4 in occlusal and slightly posterolingual (D), lingual and slightly anterior (E), and labial and slightly posterior (F) views. 154 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) and Vespertiliavus, as noted above. Two partial dentaries from Brooksville 2 contain the p4 (UF 157769–holotype [Fig. 7A-C], 182855) and a partial p4 is present in a dentary fragment from I-75 (UF 121724). The p4 is double- rooted and has a very tall, conical central cusp that is somewhat caniniform in shape. The central cusp is noticeably taller than the m1 in the only speci- men that preserves both teeth (UF 157769–holo- type; Fig. 7 A-C). The occlusal outline is that of a rounded triangle, with the rounded apex of the triangle anterior and the posterior edge horizontal. The anterior edge of the conical central cusp slopes slightly anteriorly and has a distinct ridge extend- ing from the apex almost to the anterior margin. The posterior edge of the central cusp is almost vertical. In occlusal view, there is a shallow but distinct ba- sin between the central cusp and the posterior mar- gin. There are well-developed labial, anterior, and posterior cingula. A small cuspid is present on the Figure 7. Lower teeth of Oligopteryx floridanus from Brooksville 2 LF. A-C, UF 157769 (holotype) right dentary fragment with p2, p4, and m1 in labial (A), occlusal (B), and lingual (C) views; D-E, UF 157771 left dentary fragment with m2-m3 in occlusal (D) and lingual (E) views. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 155 anterior cingulum, forming the anteriormost exten- sion of p4. There is a distinct emargination or in- dentation along the anterolingual margin of p4 just posterior to the anterolingual cuspid. The posterior cingulum bears a cuspid at the posterolingual cor- ner of the tooth. The m1 and m2 of Oligopteryx floridanus are almost identical in size and morphology and can- not be reliably separated based on isolated teeth. Among the specimens from Brooksville 2, one dentary preserves both the m1 and m2 (UF 157790) and several other dentaries preserve either the m1 (UF 157769–holotype; Fig. 7 A-C) or the m2 (UF 157771; Fig. 7 D-E). There is also a significant sample of isolated m1s or m2s from Brooksville and several from I-75. Since the m1 and m2 are so similar, the following description pertains to both teeth. The m1/m2 are nyctalodont, the postcristid connects the hypoconid to the hypoconulid, with the latter cusp at the lingual margin of the tooth. The talonid is distinctly broader than the trigonid, and the talonid is also slightly longer than the tri- gonid anteroposteriorly. The paraconid and meta- conid are located very close to one another along the lingual edge of the tooth, resulting from the anteroposterior compression of the trigonid. The metaconid is positioned noticeably anterior to the protoconid. The protocristid is oriented at an angle to the long axis of the toothrow, extending from the protoconid lingually and anteriorly to the metaconid. All four lingual cusps are rather low; the metaconid is the tallest, followed by the ento- conid and paraconid, while the hypoconulid is the lowest. All except the hypoconulid are broadly conical, triangular-shaped, with rather blunt cusps. In lingual view, the metaconid and entoconid are oriented vertically, while the paraconid is oriented at about a 45° angle anteriorly. The labial cusps are taller than the lingual cusps, with the protoco- nid the tallest cusp on the m1 and m2, followed by the hypoconid. The cristid obliqua connects to the trigonid near the lingual base of the protoco- nid and lingual to the notch in the protocristid. In several specimens (e.g., UF 157771), the cristid obliqua meets the trigonid somewhat more labially, about midway between the protoconid and meta- conid. The high, sharp entocristid in occlusal view has a V-shaped notch about halfway between the metaconid and entoconid, with the apex oriented labially. The small hypoconulid is located on the posterolingual corner of the tooth directly poste- rior to the larger entoconid. There is a basal labial cingulum extending from near the anterior margin around the protoconid along the base of the crown on the labial side of the tooth to the base of the hypoconid. The anterior labial cingulum is a well- developed but narrow shelf, terminating in an an- terolabial cuspid located just anterior and labial to the paraconid. The cingulum labial to the protoco- nid and hypoconid is not as well developed as the anterior cingulum, especially compared to some other emballonurids. There is a weak postcingulum extending from the hypoconid to the base of the hypoconulid. A lingual cingulum is lacking. Two dentaries from Brooksville 2 contain m3 (UF 157771 [Fig. 7 D-E], 157772) and there are also three isolated m3s from this site. Apart from its smaller size, the m3 is generally similar to the m1/ m2, except for the following differences. The talon- id on m3 is about the same breadth as the trigonid or is slightly narrower. The cristid obliqua forms a sharper angle with the long axis of the toothrow, meeting the trigonid farther lingually than on m1/ m2, near the labial base of the metaconid. The en- tocristid is more broadly rounded labially, not as sharply V-shaped. The hypoconulid is smaller. Oligopteryx floridanus is similar in size to the largest living New World emballonurid, Diclidu- rus ingens. It also compares well in size to the Old World Taphozous melanopogon, one of the larger extant emballonurids. The m1 and M1 lengths and humeral midshaft diameter of specimens (Tables 1, 2) provide estimates of its body weight of about 15.3 g to 24.1 g (by method of Gunnell et al., 2009). Dentary.–The characters of the dentary of Oligopteryx floridanus are based primarily on an edentulous mandible from Brooksville 2 that pre- serves the alveoli for most of the lower teeth ex- cept the m3 (UF 179958–paratype; Fig. 8), as well as three edentulous dentary fragments from I-75. The ventral margin of the horizontal ramus is fairly straight below the molars and posterior to the sym- 156 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 8. Edentulous partial horizontal ramus of dentary of Oligopteryx floridanus from Brooksville 2 LF. UF 179958, in labial (A), occlusal (B), and lingual (C) views. physis. The anterior portion of the dentary below the incisors, canine, and p2 is shallow dorsoven- trally. The mandibular symphysis extends poste- riorly to about the level of p2. Along the ventral margin of the dentary there is a sharp, transversely narrow ridge or flange extending posteroventrally from the posterior edge of the symphysis to the level of p4. This ridge is triangular in lateral view with the apex of the triangle ventral to the anterior root of p4. The mental foramen is round and fairly small, located about halfway between the alveolar and ventral margins below the p2. The incisor al- veoli are somewhat damaged, so it is not possible to determine the number of incisors present. The alveolus for the lower canine is large, about the same width as the alveolus for the p2 but somewhat longer. The canine alveolus is not round or ellipti- cal as in most bats but is figure 8-shaped or kid- ney-shaped, consisting of a broader, more rounded anterior portion and a shorter, narrower posterior portion along the lingual margin of the toothrow. The anterior portion of the canine alveolus extends across the entire breadth of the toothrow, whereas the narrower posterior portion is lingually offset. The rounded alveolus for the single-rooted p2 is large, as broad as the canine alveolus but not as long. The tiny, round alveolus of the single-rooted p3 is located along the lingual margin, wedged be- tween the posterior edge of the p2 alveolus and the anterior alveolus of the double-rooted p4. A dentary fragment from I-75 (UF 16682) is the only specimen that preserves a portion of the ascending ramus posterior to the toothrow. The an- terior edge of the coronoid process, the only portion of this process preserved, is located directly poste- rior to the m3, not lateral to the toothrow as in some other bats. Posterior to m3, the ascending ramus rises vertically such that the anterior edge of the coronoid process forms an angle of approximately 45° with the alveolar margin of the toothrow. Petrosal.–A single left petrosal (UF 179902; Fig. 9A-D) is available from the Brooksville 2 Quarry and is tentatively referred to Oligopteryx floridanus. The petrosal is fairly intact except for the damage to the crista parotica, which is almost completely broken away exposing part of the later- al semicircular canal, and to the bone over the junc- tion of the lateral and anterior semicircular canals. The fossil petrosal was compared with petrosals of modern Neotropical bats, in particular specimens of the emballonurids Balantiopteryx plicata, Di- clidurus albus, Peropteryx macrotis, and Saccop- teryx bilineata as representatives of the subfamily Emballonurinae, as well as a petrosal of a modern Afrotropical species, Taphozous mauritianus as a representative of the subfamily Taphozoinae, and of Nycteris thebaica as a representative of Nycteri- dae, the sister family to Emballonuridae (O’Leary et al., 2013; Teeling et al., 2018). Fig. 9 compares the petrosals of O. floridanus and Peropteryx mac- rotis. The petrosal of Oligopteryx (UF 179902) is about the size of that in Diclidurus albus and Ta- phozous mauritianus, and larger than the petrosals of Peropteryx macrotis, Saccopteryx bilineata, and Balantiopteryx plicata. The specimen is referable to the family Emballonuridae by virtue of its close morphological resemblance to petrosals of extant emballonurids. In particular, the fossil petrosal shows a complete lamina between the three semi- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 157 Table 2. Measurements of the lower premolars and molars of Oligocene and Miocene Emballonuridae from Florida, including: Oligopteryx floridanus and O. hamaxitos from the early Oligocene (Whitneyan) I-75 LF and the late Oligocene (early Arikaree- an) Brooksville 2 LF and Floridopteryx poyeri from the early Miocene (early Hemingfordian) Thomas Farm LF. For the lower premolars (p2 and p4), only two measurements were taken, anteroposterior length and maximum transverse width (recorded in the second column). For the lower molars (m1–m3), three measurements were taken, anteroposterior length, transverse width of the trigonid, and transverse width of the talonid. We were not able to positively identify the tooth position of isolated m1s or m2s, listed here as m1/m2. All measurements are in mm. Missing measurement indicated by “—”. Abbreviations for sample statistics: N (number of specimens); M (mean); OR (observed range). Statistics are calculated if there are three or more speci- mens for a particular tooth position. Species, fauna, anteroposterior transverse transverse tooth position, and length width width catalog number trigonid talonid Oligopteryx floridanus Brooksville 2 p2 UF 157769 (holotype) 1.45 0.75 – p4 UF 157769 (holotype) 1.30 0.85 – UF 182855 1.47 0.97 – m1 UF 157769 (holotype) 1.77 1.20 1.32 UF 157790 1.67 1.15 1.25 m2 UF 157771 1.80 1.30 1.40 UF 157788 1.72 1.22 1.30 UF 157789 1.67 1.27 1.45 UF 157790 1.62 1.20 1.27 UF 182874 1.75 1.22 1.30 N 5 5 5 M 1.71 1.24 1.34 OR 1.62–1.80 1.20–1.30 1.27–1.45 m1/m2 UF 156289 1.80 1.22 1.30 UF 157782 1.70 1.22 1.35 UF 157783 1.80 1.12 1.30 UF 179987 1.75 1.30 1.37 UF 182809 1.75 1.12 1.27 UF 182811 1.80 1.22 1.30 UF 182813 1.75 1.20 1.25 UF 182814 1.72 1.10 1.35 UF 182819 1.75 1.15 1.40 UF 182859 1.82 1.30 1.37 UF 182860 1.85 1.07 1.32 UF 182861 1.75 1.10 1.35 UF 182862 1.67 1.17 1.30 UF 182864 1.70 1.22 1.30 UF 182865 1.75 1.20 1.27 UF 182867 1.77 1.07 1.32 UF 182868 1.75 1.17 1.35 UF 182870 1.77 1.20 1.45 UF 182871 1.75 1.17 1.35 UF 182891 1.70 1.15 1.27 UF 182892 1.65 1.15 1.25 UF 182893 1.70 1.25 1.32 N 22 22 22 M 1.75 1.18 1.32 OR 1.65–1.85 1.07–1.30 1.25–1.45 Oligopteryx floridanus (cont.) I-75 m1/m2 UF 121705 1.70 1.10 1.10 UF 121706 1.77 1.30 1.42 UF 121707 1.72 1.05 1.10 UF 121708 1.72 1.20 1.35 N 4 4 4 M 1.73 1.16 1.24 OR 1.70–1.77 1.05–1.30 1.10–1.42 Brooksville 2 m3 UF 157771 1.60 1.22 1.05 UF 157772 1.57 1.17 1.05 UF 157787 1.60 1.02 1.00 UF 182857 1.60 1.15 1.02 UF 182872 1.55 1.15 1.02 UF 182895 1.65 1.22 1.05 N 6 6 6 M 1.60 1.16 1.03 OR 1.55–1.65 1.02–1.22 1.00–1.05 Oligopteryx hamaxitos Brooksville 2 m1 UF 157768 (holotype) 1.10 0.75 0.92 m2 UF 157768 (holotype) 1.12 0.85 0.90 m1/m2 UF 182817 1.25 0.80 0.90 UF 182869 1.27 0.82 0.92 Floridopteryx poyeri Thomas Farm m1 UF 121132 (holotype) 1.52 0.92 1.12 m1/m2 UF 121133 1.40 0.85 1.05 m3 UF 108664 1.32 0.97 0.87 Species, fauna, anteroposterior transverse transverse tooth position, and length width width catalog number trigonid talonid Oligopteryx floridanus Brooksville 2 p2 UF 157769 (holotype) 1.45 0.75 – p4 UF 157769 (holotype) 1.30 0.85 – UF 182855 1.47 0.97 – m1 UF 157769 (holotype) 1.77 1.20 1.32 UF 157790 1.67 1.15 1.25 m2 UF 157771 1.80 1.30 1.40 UF 157788 1.72 1.22 1.30 UF 157789 1.67 1.27 1.45 UF 157790 1.62 1.20 1.27 UF 182874 1.75 1.22 1.30 N 5 5 5 M 1.71 1.24 1.34 OR 1.62–1.80 1.20–1.30 1.27–1.45 m1/m2 UF 156289 1.80 1.22 1.30 UF 157782 1.70 1.22 1.35 UF 157783 1.80 1.12 1.30 UF 179987 1.75 1.30 1.37 UF 182809 1.75 1.12 1.27 UF 182811 1.80 1.22 1.30 UF 182813 1.75 1.20 1.25 UF 182814 1.72 1.10 1.35 UF 182819 1.75 1.15 1.40 UF 182859 1.82 1.30 1.37 UF 182860 1.85 1.07 1.32 UF 182861 1.75 1.10 1.35 UF 182862 1.67 1.17 1.30 UF 182864 1.70 1.22 1.30 UF 182865 1.75 1.20 1.27 UF 182867 1.77 1.07 1.32 UF 182868 1.75 1.17 1.35 UF 182870 1.77 1.20 1.45 UF 182871 1.75 1.17 1.35 UF 182891 1.70 1.15 1.27 UF 182892 1.65 1.15 1.25 UF 182893 1.70 1.25 1.32 N 22 22 22 M 1.75 1.18 1.32 OR 1.65–1.85 1.07–1.30 1.25–1.45 Oligopteryx floridanus (cont.) I-75 m1/m2 UF 121705 1.70 1.10 1.10 UF 121706 1.77 1.30 1.42 UF 121707 1.72 1.05 1.10 UF 121708 1.72 1.20 1.35 N 4 4 4 M 1.73 1.16 1.24 OR 1.70–1.77 1.05–1.30 1.10–1.42 Brooksville 2 m3 UF 157771 1.60 1.22 1.05 UF 157772 1.57 1.17 1.05 UF 157787 1.60 1.02 1.00 UF 182857 1.60 1.15 1.02 UF 182872 1.55 1.15 1.02 UF 182895 1.65 1.22 1.05 N 6 6 6 M 1.60 1.16 1.03 OR 1.55–1.65 1.02–1.22 1.00–1.05 Oligopteryx hamaxitos Brooksville 2 m1 UF 157768 (holotype) 1.10 0.75 0.92 m2 UF 157768 (holotype) 1.12 0.85 0.90 m1/m2 UF 182817 1.25 0.80 0.90 UF 182869 1.27 0.82 0.92 Floridopteryx poyeri Thomas Farm m1 UF 121132 (holotype) 1.52 0.92 1.12 m1/m2 UF 121133 1.40 0.85 1.05 m3 UF 108664 1.32 0.97 0.87 158 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Table 2. Cont. Species, fauna, anteroposterior transverse transverse tooth position, and length width width catalog number trigonid talonid Oligopteryx floridanus Brooksville 2 p2 UF 157769 (holotype) 1.45 0.75 – p4 UF 157769 (holotype) 1.30 0.85 – UF 182855 1.47 0.97 – m1 UF 157769 (holotype) 1.77 1.20 1.32 UF 157790 1.67 1.15 1.25 m2 UF 157771 1.80 1.30 1.40 UF 157788 1.72 1.22 1.30 UF 157789 1.67 1.27 1.45 UF 157790 1.62 1.20 1.27 UF 182874 1.75 1.22 1.30 N 5 5 5 M 1.71 1.24 1.34 OR 1.62–1.80 1.20–1.30 1.27–1.45 m1/m2 UF 156289 1.80 1.22 1.30 UF 157782 1.70 1.22 1.35 UF 157783 1.80 1.12 1.30 UF 179987 1.75 1.30 1.37 UF 182809 1.75 1.12 1.27 UF 182811 1.80 1.22 1.30 UF 182813 1.75 1.20 1.25 UF 182814 1.72 1.10 1.35 UF 182819 1.75 1.15 1.40 UF 182859 1.82 1.30 1.37 UF 182860 1.85 1.07 1.32 UF 182861 1.75 1.10 1.35 UF 182862 1.67 1.17 1.30 UF 182864 1.70 1.22 1.30 UF 182865 1.75 1.20 1.27 UF 182867 1.77 1.07 1.32 UF 182868 1.75 1.17 1.35 UF 182870 1.77 1.20 1.45 UF 182871 1.75 1.17 1.35 UF 182891 1.70 1.15 1.27 UF 182892 1.65 1.15 1.25 UF 182893 1.70 1.25 1.32 N 22 22 22 M 1.75 1.18 1.32 OR 1.65–1.85 1.07–1.30 1.25–1.45 Oligopteryx floridanus (cont.) I-75 m1/m2 UF 121705 1.70 1.10 1.10 UF 121706 1.77 1.30 1.42 UF 121707 1.72 1.05 1.10 UF 121708 1.72 1.20 1.35 N 4 4 4 M 1.73 1.16 1.24 OR 1.70–1.77 1.05–1.30 1.10–1.42 Brooksville 2 m3 UF 157771 1.60 1.22 1.05 UF 157772 1.57 1.17 1.05 UF 157787 1.60 1.02 1.00 UF 182857 1.60 1.15 1.02 UF 182872 1.55 1.15 1.02 UF 182895 1.65 1.22 1.05 N 6 6 6 M 1.60 1.16 1.03 OR 1.55–1.65 1.02–1.22 1.00–1.05 Oligopteryx hamaxitos Brooksville 2 m1 UF 157768 (holotype) 1.10 0.75 0.92 m2 UF 157768 (holotype) 1.12 0.85 0.90 m1/m2 UF 182817 1.25 0.80 0.90 UF 182869 1.27 0.82 0.92 Floridopteryx poyeri Thomas Farm m1 UF 121132 (holotype) 1.52 0.92 1.12 m1/m2 UF 121133 1.40 0.85 1.05 m3 UF 108664 1.32 0.97 0.87 Species, fauna, anteroposterior transverse transverse tooth position, and length width width catalog number trigonid talonid Oligopteryx floridanus Brooksville 2 p2 UF 157769 (holotype) 1.45 0.75 – p4 UF 157769 (holotype) 1.30 0.85 – UF 182855 1.47 0.97 – m1 UF 157769 (holotype) 1.77 1.20 1.32 UF 157790 1.67 1.15 1.25 m2 UF 157771 1.80 1.30 1.40 UF 157788 1.72 1.22 1.30 UF 157789 1.67 1.27 1.45 UF 157790 1.62 1.20 1.27 UF 182874 1.75 1.22 1.30 N 5 5 5 M 1.71 1.24 1.34 OR 1.62–1.80 1.20–1.30 1.27–1.45 m1/m2 UF 156289 1.80 1.22 1.30 UF 157782 1.70 1.22 1.35 UF 157783 1.80 1.12 1.30 UF 179987 1.75 1.30 1.37 UF 182809 1.75 1.12 1.27 UF 182811 1.80 1.22 1.30 UF 182813 1.75 1.20 1.25 UF 182814 1.72 1.10 1.35 UF 182819 1.75 1.15 1.40 UF 182859 1.82 1.30 1.37 UF 182860 1.85 1.07 1.32 UF 182861 1.75 1.10 1.35 UF 182862 1.67 1.17 1.30 UF 182864 1.70 1.22 1.30 UF 182865 1.75 1.20 1.27 UF 182867 1.77 1.07 1.32 UF 182868 1.75 1.17 1.35 UF 182870 1.77 1.20 1.45 UF 182871 1.75 1.17 1.35 UF 182891 1.70 1.15 1.27 UF 182892 1.65 1.15 1.25 UF 182893 1.70 1.25 1.32 N 22 22 22 M 1.75 1.18 1.32 OR 1.65–1.85 1.07–1.30 1.25–1.45 Oligopteryx floridanus (cont.) I-75 m1/m2 UF 121705 1.70 1.10 1.10 UF 121706 1.77 1.30 1.42 UF 121707 1.72 1.05 1.10 UF 121708 1.72 1.20 1.35 N 4 4 4 M 1.73 1.16 1.24 OR 1.70–1.77 1.05–1.30 1.10–1.42 Brooksville 2 m3 UF 157771 1.60 1.22 1.05 UF 157772 1.57 1.17 1.05 UF 157787 1.60 1.02 1.00 UF 182857 1.60 1.15 1.02 UF 182872 1.55 1.15 1.02 UF 182895 1.65 1.22 1.05 N 6 6 6 M 1.60 1.16 1.03 OR 1.55–1.65 1.02–1.22 1.00–1.05 Oligopteryx hamaxitos Brooksville 2 m1 UF 157768 (holotype) 1.10 0.75 0.92 m2 UF 157768 (holotype) 1.12 0.85 0.90 m1/m2 UF 182817 1.25 0.80 0.90 UF 182869 1.27 0.82 0.92 Floridopteryx poyeri Thomas Farm m1 UF 121132 (holotype) 1.52 0.92 1.12 m1/m2 UF 121133 1.40 0.85 1.05 m3 UF 108664 1.32 0.97 0.87 Species, fauna, anteroposterior transverse transverse tooth position, and length width width catalog number trigonid talonid Oligopteryx floridanus Brooksville 2 p2 UF 157769 (holotype) 1.45 0.75 – p4 UF 157769 (holotype) 1.30 0.85 – UF 182855 1.47 0.97 – m1 UF 157769 (holotype) 1.77 1.20 1.32 UF 157790 1.67 1.15 1.25 m2 UF 157771 1.80 1.30 1.40 UF 157788 1.72 1.22 1.30 UF 157789 1.67 1.27 1.45 UF 157790 1.62 1.20 1.27 UF 182874 1.75 1.22 1.30 N 5 5 5 M 1.71 1.24 1.34 OR 1.62–1.80 1.20–1.30 1.27–1.45 m1/m2 UF 156289 1.80 1.22 1.30 UF 157782 1.70 1.22 1.35 UF 157783 1.80 1.12 1.30 UF 179987 1.75 1.30 1.37 UF 182809 1.75 1.12 1.27 UF 182811 1.80 1.22 1.30 UF 182813 1.75 1.20 1.25 UF 182814 1.72 1.10 1.35 UF 182819 1.75 1.15 1.40 UF 182859 1.82 1.30 1.37 UF 182860 1.85 1.07 1.32 UF 182861 1.75 1.10 1.35 UF 182862 1.67 1.17 1.30 UF 182864 1.70 1.22 1.30 UF 182865 1.75 1.20 1.27 UF 182867 1.77 1.07 1.32 UF 182868 1.75 1.17 1.35 UF 182870 1.77 1.20 1.45 UF 182871 1.75 1.17 1.35 UF 182891 1.70 1.15 1.27 UF 182892 1.65 1.15 1.25 UF 182893 1.70 1.25 1.32 N 22 22 22 M 1.75 1.18 1.32 OR 1.65–1.85 1.07–1.30 1.25–1.45 Oligopteryx floridanus (cont.) I-75 m1/m2 UF 121705 1.70 1.10 1.10 UF 121706 1.77 1.30 1.42 UF 121707 1.72 1.05 1.10 UF 121708 1.72 1.20 1.35 N 4 4 4 M 1.73 1.16 1.24 OR 1.70–1.77 1.05–1.30 1.10–1.42 Brooksville 2 m3 UF 157771 1.60 1.22 1.05 UF 157772 1.57 1.17 1.05 UF 157787 1.60 1.02 1.00 UF 182857 1.60 1.15 1.02 UF 182872 1.55 1.15 1.02 UF 182895 1.65 1.22 1.05 N 6 6 6 M 1.60 1.16 1.03 OR 1.55–1.65 1.02–1.22 1.00–1.05 Oligopteryx hamaxitos Brooksville 2 m1 UF 157768 (holotype) 1.10 0.75 0.92 m2 UF 157768 (holotype) 1.12 0.85 0.90 m1/m2 UF 182817 1.25 0.80 0.90 UF 182869 1.27 0.82 0.92 Floridopteryx poyeri Thomas Farm m1 UF 121132 (holotype) 1.52 0.92 1.12 m1/m2 UF 121133 1.40 0.85 1.05 m3 UF 108664 1.32 0.97 0.87 circular canals. This lamina is lacking in most fam- ilies of bats in the Western Hemisphere except for Emballonuridae and some members of the Molos- sidae; it is shared with the Eastern Hemisphere Nycteridae. The fossil petrosal also has a complete lamina closing off the posterior semicircular canal with a few minuscule circular perforations (and a few tiny broken areas). The posterior semicircular canal is also completely laminated in Peropteryx, Saccopteryx, and Balantiopteryx. By contrast there is no lamina within the posterior semicircular canal in Diclidurus and Taphozous, and only a periph- eral flange of lamina with a large central opening in Nycteris. The posterior semicircular canal is much smaller than either of the other two semicircular canals in the fossil and modern specimens. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 159 The fossil cochlea is phanerocochlear (hav- ing the petrosal wall thinly ossified such that the cochlear labyrinth is barely visible externally (No- vacek 1985; 1991), as in all of the modern taxa examined. On the posteroventral surface of the cochlea is a small circular tubercle with a central hollow (Fig. 9, labeled “tu”); this tubercle is well developed also in Balantiopteryx, Peropteryx, Di- clidurus, and Taphozous but is weaker in Saccop- teryx. It is absent in Nycteris. The fenestra cochleae in UF 179902 in posterior view is wider than high, rounded ventrally and flat along its dorsal edge; it Figure 9. Petrosal bones of Emballonuridae. A-D, Oligopteryx floridanus from Brooksville 2, UF 179902, left petrosal with labeled interpretative drawings above and photographs below, in (A) lateral view, oriented with the lateral semicircular canal parallel to the horizon; (B) dorsal (endocranial) view; (C) ventrolateral view; and (D) posterior view. E-H, Peropteryx macrotis, modern specimen from Tikal, El Petén, Guatemala, UF Mammalogy 6935, left petrosal in views matching those of A-D. The fossa for the stapedius muscle is filled with organic matrix in the modern specimen. Orientation arrows indicate approximate anterior (ant.), dorsal (dors.), lateral (lat.), and medial (med.) directions. Abbreviations: ant, anterior; asc, anterior semicircular canal; avs, area vestibularis superior; cc, cochlear canaliculus (opening of cochlear aqueduct or perilymphatic duct); cp, crista parotica (damaged and mostly absent in the fossil); cr, common crus; dors, dorsal; fc, fenestra cochleae; fn, facial canal (for cranial nerve VII); fs, facial sulcus (semicanal for facial nerve); fv, fenestra vestibuli for footplate of stapes; lam, laminar cap between semicircular canals; lat, lateral; lsc, lateral semicircular canal; med, medial; op, process overhanging fenestra cochle- ae; pf, prefacial commissure; pl, platelike structure at lateral end of posterior semicircular canal; pr, promontorium of cochlea; psc, posterior semicircular canal; sa, subarcuate fossa; sf, fossa for stapedius muscle; tsf, spiral foraminous tract within the internal acoustic meatus; tu, small tubercle anteroventral to fenestra cochleae; va, opening of vestibular aqueduct; vent, ventral. 160 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) is larger than the fenestra vestibuli, as is also true in the other emballonurids examined except for Di- clidurus, in which the two fenestrae are about the same size. The fenestra cochleae is more elongate in Balantiopteryx, Saccopteryx, and Taphozous, and especially elongate in Peropteryx. The fenestra cochleae is oval and higher than wide in Nycteris, about the same size as the fenestra vestibuli. Over- hanging the fenestra cochleae in the fossil petrosal and the petrosals of all the recent emballonurids examined, there is a pointed process or flange of bone (Fig. 9 labeled “op”); this process occurs at the junction of the posterior and lateral semicircu- lar canals. It includes a flangelike extension that runs anteriorly to form a partial floor beneath the stapedial fossa in all the emballonurids; this exten- sion might enlarge the surface area for origin of the stapedius muscle. In the fossil, and in Peropteryx, Saccopteryx and Balantiopteryx the portion over- hanging the fenestra cochleae occurs as a pointed or V-shaped process, whereas in Diclidurus it oc- curs as a long thin flange, and in Taphozous it is broader and rounded instead of pointed. Just dorsal and posterior to this process at the lateral base of the posterior semicircular canal, the fossil bears a relatively large, posteriorly facing, oval, flat plate- like area that is slightly recessed (Fig. 9 labeled “pl”); this platelike area is about the same size and shape in Taphozous but twice the size in the fossil as in any of the other recent emballonurids. In Per- opteryx, Saccopteryx, and Balantiopteryx it is situ- ated similarly, but in Diclidurus it occurs behind a curved thin flange of bone. The petrosal of Nycteris completely lacks the process, flange, and platelike area seen in the emballonurids. The prefacial commissure is relatively thick and transmits a moderate length of facial canal in UF 179902, Taphozous, and Saccopteryx, is thin and spans a short facial canal in Balantiopteryx and Peropteryx, and is thick and encloses a very long facial canal in Diclidurus. The area vestibularis su- perior occurs as a small, finely perforated, subcir- cular plate adjacent to the dorsal (endocranial) end of the facial canal in the fossil and all recent em- ballonurids. The facial canal is smaller in Nycteris, and the area vestibularis superior is even smaller. The cochlear canaliculus is a small pore, much smaller than the fenestra vestibuli. The co- chlear canaliculus is small in the recent emballon- urids examined, too, and is situated closer to the fenestra cochleae in Peropteryx but about the same distance from the fenestra cochleae in the fossil as in Balantiopteryx, Saccopteryx, Diclidurus, and Taphozous. In the fossil the external aperture of the vestibular aqueduct (Fig. 9 labeled “va”) is large and rather slitlike, with an angled opening hav- ing its dorsal rim extending beyond the common crus while the posterior edge does not extend be- yond the common crus. The structure is similar in Peropteryx, Balantiopteryx, Saccopteryx, and Ta- phozous. In Diclidurus both edges of the aperture extend equally beyond the common crus, and to- gether extend somewhat farther than in the other emballonurids as a flattened tubelike structure, and the extension bears an adjacent flange of thin bone extending posteriad from the posterior side of the tubelike structure. In Nycteris, the aperture of the vestibular aqueduct is similarly slit-like but lacks a spout-like or funnel-like extension. A number of important features occur on the crista parotica of bats (e.g., Morgan et al., 2019); unfortunately, these features cannot be examined or compared because of breakage of this structure in UF 179902. Sulser et al. (2022) studied neuroanatomy of the cochlea in bats including two extant emballon- urids (Taphozous nudiventris and Coleura afra). Their work examined the Rosenthal’s canal wall for the spiral ganglion, which is perforated to varying degrees for the passage of fascicles of the cochle- ar nerve. The canal wall can be (1) perforated by many small holes for the nerve bundles (in which case it is called a “foraminal wall” or “tractus fo- raminosus” [and termed “spiral foraminous tract” in our Fig. 9B, labeled “tsf”]; this is the primitive condition of the entire tract in most mammals and non-echolocating bats), or (2) perforated by fewer larger holes (in which case it is called a “fenestral wall” or “tractus fenestralis”), or (3) widely open along its length (in which case it is called “wall- less”). These authors (Sulser et al., 2022) further MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 161 defined the tracts along the spiral of the canal at the base (basal ½ turn, near the open portion of the in- ternal acoustic meatus), middle (at the 1.5 cochlear turn), and apex (deep within the internal acoustic meatus near the apical turns). Among the two ex- tant emballonurids examined, Sulser et al. (2022) found the tract in C. afra to be fenestrated in the base and middle, and wall-less nearer the apex. In T. nudiventris, they found a tractus foraminosus in the basal turn, tractus fenestralis in the middle, and wall-less condition in the apical portion. In the Oligocene fossil petrosal that we attribute to Oli- gopteryx floridanus, we note that the basal turn of the canal wall is foraminated (possibly partly dam- aged), whereas deeper turns are not visible with optical inspection and might require CT-scanning to properly characterize them. The only other isolated petrosal available from an Oligocene bat in North America is that of the mormoopid Koopmanycteris palaeomormoops (UF 179901), described and illustrated by Morgan et al (2019), from the same site that produced the petrosal of Oligopteryx floridanus, the Brooksville 2 LF. Both Oligopteryx and Koopmanycteris have a phanerocochlear petrosal. The most obvious differ- ence between the petrosal of O. floridanus and that of Koopmanycteris is the complete lamina between the semicircular canals that is absent in K. palaeo- mormoops (and other mormoopids). The petrosal of O. floridanus also has a lamina cap within the posterior semicircular canal (so that the only open- ing into the subarcuate fossa is through the anterior semicircular canal); in Koopmanycteris the posteri- or semicircular canal lacks a laminar cap or flanges. As noted above, O. floridanus has a Rosenthal’s ca- nal wall with a tractus foraminosus in its basal half turn; in Koopmanycteris, Rosenthal’s canal wall is similarly foraminated in its basal and additionally in its middle turns, but the condition in the apical turns is not visible. Koopmanycteris lacks the small circular tubercle (Fig. 9, “tu”) on the posteroven- tral surface of the cochlea; it also lacks the large pointed or V-shaped process that overhangs the fe- nestra cochleae in Oligopteryx. While the cochlear canaliculus in O. floridanus is much smaller than the fenestra vestibuli, the same opening in Koop- manycteris is about the same size as the fenestra vestibuli. While the external aperture of the ves- tibular aqueduct in O. floridanus (Fig.9, “va”) is on a rather spoutlike tiny projection from the common crus, that in Koopmanycteris is not on a bony pro- jection; instead, the aperture in Koopmanycteris is a small, curved slit along the common crus. Humerus.–There are nine proximal ends and eight distal ends of the humerus of Oligopteryx floridanus from Brooksville 2 and a single distal humerus from I-75, although there are no complete specimens. The humeri are not directly associated with dental material. The association is based on the presence of large (and small) humeri that com- pare closely to humeri of modern members of the Emballonuridae, occurring together with teeth and dentaries that also represent large (and small) spe- cies belonging to the same family. The following description is based primarily on four well-pre- served specimens, a proximal end of a left humerus (UF 179904, paratype), a proximal right humerus (UF 179936), a distal end of a left humerus (UF 179964, paratype; Fig. 10), and a distal right hu- merus (UF 179910) with more of the shaft pre- served than UF 179964 but with some damage to the articular surface. The remainder of the sample was examined and used to help determine varia- tion. All well-preserved proximal and distal ends of the humerus of O. floridanus were measured (Table 3). In posterior view, the head on the proximal end of the humerus is elliptical in shape, somewhat transversely flattened, rounded distally, and more pointed or triangular proximally. The humeral head projects farther proximally than either the greater or lesser tuberosities. The head is not in the cen- ter of the shaft but is shifted noticeably laterally, located closer to the greater tuberosity than to the lesser tuberosity. The head is oriented at a slight angle to the shaft, canted distally toward the great- er tuberosity. The greater tuberosity is rather elon- gated, rounded at its proximal end, and oriented at about a 45° angle to the shaft. The lesser tuberosity is considerably larger than the greater tuberosity, forming a large, triangular-shaped process with a squared-off proximal end. The greater and lesser 162 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Table 3. Measurements of the humerus and radius of Oligopteryx floridanus and O. hamaxitos from the Oligocene Brooksville 2 LF (Arikareean) and I-75 LF (Whitneyan) of Florida. All measurements are in mm. Missing measurements are indicated by “–”. Abbreviations for sample statistics: N (number of specimens); M (mean); OR (observed range). Statistics were calculated if there were three or more specimens for a particular limb element. Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 163 Table 3. Cont. tuberosities project about the same distance proxi- mally. In proximal view, the pointed proximal por- tion of the head extends anteriorly to the base of the pectoral ridge. The larger size of the lesser tuberos- ity compared to the greater tuberosity is even more evident when viewed proximally. The lesser tuber- osity is a prominent, triangular-shaped process ex- tending medially, with a rounded anterior edge and straight posterior edge. It is about three times larger than the greater tuberosity. In this same view, the greater tuberosity is a narrow process with rounded edges, somewhat elongated in the anteroposterior dimension, with a shallow indentation about half- way along its length. The supraglenoid fossa is very shallow. In lateral view, the pectoral ridge is rather short but broad, triangular in shape, broader proximally. There is some variation in the shape of the pectoral ridge in the sample of proximal humeri from Brooksville 2; in most specimens this ridge is triangular but in several it is more rectangular. In medial view, the pectoral ridge bears a distinct, raised ridge that is oriented proximodistally, divid- ing the pectoral ridge into two approximately equal sections. There is a weak medial ridge on the shaft distal to the lesser tuberosity. The distal articular surface of the humerus is slightly offset laterally from the shaft (Fig. 10). In anterior view, the lateral edge of the humeral shaft is in line with the weak groove separating the medial and lateral ridges of the capitulum, with the lateral ridge of the capitulum located lateral to the shaft. The lateral ridge of the capitulum is narrow transversely, occupying about one-fourth the breadth of the distal articular surface, but is deep in the proximo-distal dimension, extending proximally well beyond the medial ridge of the capitulum and trochlea. The medial ridge of the capitulum is large, bulbous, and somewhat spheri- cal, composing about half the breadth of the distal articular surface. The medial and lateral ridges of the capitulum are separated by a shallow groove. The trochlea occupies about one-fourth the width of the distal articular surface and forms about a 45° angle with the medial ridge of the capitulum. The medial edge of the trochlea is essentially ver- tical and extends slightly farther distally than the capitulum. The medial ridge of the capitulum and trochlea are separated by a much deeper groove than the groove that separates the two parts of the capitulum. The medial epicondyle (epitrochlea), including the medial process of the epitrochlea and distal spinous process, is separated from the medial edge of the trochlea by a deep, rounded notch. Al- most the entire medial epicondyle extends medial to the medial edge of the shaft. The medial process is well developed and rounded medially. The tip of the spinous process is sharply triangular and ex- tends distally about half the distance between the proximal and distal edges of the articular surface, although this process is somewhat shorter in sever- Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 164 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) al specimens. The radial fossa is weak and shallow in most specimens, but in several humeri, including UF 179910, the radial fossa is deeper with distinct edges proximal to the trochlea. In posterior view, the small olecranon fossa varies from shallow to almost nonexistent. There are no ridges that extend distally from the medial and lateral margins of the shaft onto the distal articular surface and medial epicondyle. In medial view, the posterior edge of the spinous process is vertical or parallel to the shaft, whereas the anterior edge of this process an- gles posteriorly at about 45° toward the distal tip. The humeri of Oligopteryx floridanus from Brooksville 2 and I-75, as well as several very similar humeri of the smaller O. hamaxitos from the same two sites, are identified as emballonurids by the elliptical and transversely flattened humeral head that angles distally toward the greater tuber- osity, distal articular surface slightly offset laterally from the shaft, well-developed medial epicondyle bearing a strong distal spinous process separated from the trochlea by a deep rounded notch, and the large bulbous medial ridge of the capitulum. The distal articular surface of the humerus is more strongly offset laterally from the shaft in most other New World chiropteran families, including Mormoopidae, Phyllostomidae, Furipteridae, Thy- ropteridae, and Natalidae (Smith, 1972). In these families, the lateral ridge of the capitulum and at least a portion of the medial ridge of the capitulum are lateral to the humeral shaft, whereas in most emballonurids only the narrow lateral ridge is lat- eral to the shaft. In the Noctilionidae, the lateral ridge of the capitulum is more noticeably offset from the shaft and the central ridge of the capit- ulum is deeper but not as broad or as bulbous as in emballonurids. The distal articular surface is in line with the shaft in Vespertilionidae and Molos- sidae and the medial ridge of the capitulum is much narrower and almost ridge-like compared to the broader more bulbous medial ridge in emballon- urids. Vespertilionids have a reduced distal spinous process compared to emballonurids. Some molos- sids have a well-developed distal spinous process but it is connected directly to the medial edge of the trochlea, not separated from the trochlea by a Figure 10. Distal portion of humerus of Oligopteryx floridanus from Brooksville 2 LF. UF 179964, in anterior (A), posterior (B), lateral (C), distal (D), and medial (E) views. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 165 distinct notch as in emballonurids. Radius.—There are 12 specimens of the proximal radius of Oligopteryx floridanus, ten from Brooksville 2 and two from I-75, as well as six distal radii from Brooksville. All of the com- plete proximal and distal ends of the radius were measured (Table 3). The proximalmost extension of the proximal radius has a gently rounded cur- vature, rather than consisting of a distinctly trian- gular-shaped process as in most other bats. The proximal articular surface consists of three slightly concave to nearly flat articular facets, correspond- ing to the three articular facets on the distal end of the humerus, the lateral and medial ridges of the capitulum and the trochlea. The facet for the me- dial ridge of the capitulum is a large, rounded con- cavity that occupies almost half of the proximal ar- ticular surface of the radius. The articular facet for the lateral ridge of the capitulum is shallower and narrower transversely but is taller or deeper in the proximodistal direction. The articular facet for the trochlea is nearly flat and consists of a narrow, me- dially directed process that is blunt or squared-off on its medial surface. Distal to the articular surface for the trochlea is a deep pit, the flexor fossa. There is a prominent ridge or flange along the medial edge of the shaft on the posterior surface distal to the flexor fossa. Two proximal radii from Brooks- ville 2 (UF 179911, paratype, 179912) preserve a considerably longer portion of the shaft than do the other specimens of the proximal radius. Both specimens consist of about the proximal one-third of the radius, preserving enough of the shaft to ex- hibit the very sharp bend or bowing of the shaft that is characteristic of emballonurids. In the complete radius of living emballonurids, this sharp bend oc- curs about one-third the distance between the prox- imal and distal ends. A sharp, narrow, raised ridge on the posterior surface of the shaft in UF 179912 marks the point where the thin, thread-like shaft of the ulna becomes fused with the radius, about halfway along the shaft but slightly closer to the proximal end. These two specimens also have a strong ridge along the medial edge of the shaft that extends from the flange distal to the flexor fossa distally beyond the break in the shaft. This ridge is strongest at midshaft. The distal end of the radius is rather simple, exhibiting few morphological characters. We iden- tified four distal radii of Oligopteryx floridanus from Brooksville 2, as well as a single distal radius of the smaller O. hamaxitos (described below). In anterior view, the distal edge of the articular sur- face is essentially straight, nearly horizontal to the shaft. There is a slight convexity representing the styloid process on the anterolateral edge of the ar- ticular surface. In posterior view, the distal edge of the articular surface forms a 45° angle to the shaft, trending distally from lateral to medial. The distal articular surface is deeply concave where it articu- lates with the lunar bone of the carpus. Just proxi- mal to the lateral edge of the distal articular surface on the posterior surface of the shaft is a prominent, triangular-shaped, laterally oriented process that corresponds to the distalmost portion of the ulna. The most obvious characters associating these radii with the Emballonuridae are the gently rounded proximalmost portion of the proximal ar- ticular surface and the strongly bent shaft about a third of the distance from the proximal end. In most other bats, the proximal extension of the radius is distinctly triangular in shape and the shaft is more gently curved. Except for their greater size, the fos- sil radii are very similar in morphology to the ra- dius of living Neotropical emballonurids, such as Peropteryx and Saccopteryx. The only noticeable difference is the larger size and greater distal exten- sion of the ridge or flange on the medial edge of the shaft just distal to the flexor fossa. Femur.—A single proximal end of a femur of Oligopteryx floridanus was identified from the Brooksville 2 LF (UF 182788). This femur is easily distinguished from the proximal femur of the other common bat from Brooksville 2, the mormoopid Koopmanycteris palaeomormoops, by its larger size and much better development of the greater and lesser trochanters. The trochanters are highly reduced in mormoopids. Except for its larger size, the Brooksville emballonurid femur compares fa- vorably with the femur of Saccopteryx. The femo- ral head of Oligopteryx is located in the center of the shaft, is generally spherical in shape although 166 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) slightly flattened proximally, and bears a large centrally positioned fovea. The lesser trochanter is slightly larger than the greater trochanter and is somewhat triangular in shape, the greater trochan- ter is more pointed. The shaft is slightly bent later- ally at the level of the medial ridge on the medial edge of shaft distal to the lesser trochanter. comparIsons wITh oTher emBallonurIdae Considering the geographic proximity and general similarity in age, the most important com- parisons of Oligopteryx floridanus from the early Oligocene (Whitneyan) I-75 LF and the late Oli- gocene (late early Arikareean) Brooksville 2 LF are with other Oligocene and early Miocene Em- ballonuridae from Florida, including O. hamaxi- tos from the same two localities as O. floridanus, Karstopteryx gunnelli from the latest Oligocene (early late Arikareean) Buda LF, and Floridopteryx poyeri from the early Miocene (early Hemingford- ian) Thomas Farm LF. Next, we compare O. flori- danus to the five extinct genera of Emballonuridae described from the Eocene, Oligocene, and Mio- cene of the Old World, Afrillonura, Dhofarella, Pseudovespertiliavus, Tachypteron, and Vespertil- iavus. Table 4 lists comparative dental characters for the eight extinct genera of Emballonuridae, five from the Old World listed in the previous sentence (Ravel et al., 2016; Rosina and Pickford, 2021) and three from the New World (Florida) described here, Oligopteryx, Karstopteryx, and Floridopteryx. Al- though originally referred to the Emballonuridae by Hooker (1996), Eppsinycteris from the early Eocene of England is almost certainly not an em- ballonurid (Storch et al., 2002); it was placed in the Onychonycteridae by Smith et al. (2012). Finally, we compare O. floridanus to each of the 14 living genera of Emballonuridae (see list of modern com- parative material examined in Appendix 1). Six extant species of emballonurids are also included in our analysis of dental characters in Table 4, in- cluding: two Old World species, Taphozous mela- nopogon (Taphozoinae) and Coleura afra (Embal- lonurinae: Emballonurini); and four New World species, Balantiopteryx plicata, Diclidurus albus, Peropteryx macrotis, and Saccopteryx bilineata (Emballonurinae: Diclidurini). Comparisons with other extinct Emballonur- idae from Florida.—Among the three other species of extinct Emballonuridae described here from the Oligocene and early Miocene of Florida, Oligop- teryx floridanus is most similar to O. hamaxitos, both of which occur in the Oligocene Brooksville 2 and I-75 LFs, with the type specimens and larg- est samples of both species from Brooksville 2. O. floridanus differs from O hamaxitos primarily in its larger size (Tables 1–3). The mean anteroposterior length (1.80 mm) and transverse width (2.38 mm) of a sample of 11 M1s of O. floridanus are approxi- mately 25-30% larger than those same two mea- surements in a paratype M1 of O. hamaxitos (1.37 mm, 1.65 mm, respectively). Except for the dispar- ity in size, these two species are very similar in the morphology of the M1 and M2, with the main dif- ference being the presence of a weak paraloph and absence of a metaloph in O. floridanus compared to a well developed paraloph and metaloph on both M1 and M2 in O. hamaxitos. There are also minor differences between these two species in the upper and lower canines and distal humerus that are dis- cussed in more detail under the species account of O. hamaxitos. Only the M1 of Oligopteryx floridanus can be compared to Karstopteryx gunnelli from the latest Oligocene Buda LF (Table 1), because the latter species is known from a single M1 (see be- low). O. floridanus and K. gunnelli are similar in size and in the reduction of the parastylar region of the M1, including the reduced parafossa labial to the paracone and the short preparacrista. How- ever, O. floridanus can be readily distinguished from K. gunnelli in several other characters of the M1, including the absence of a parastyle, labial or posterior orientation of the preparacrista, rather strong V-shaped indentation in the labial margin of the metafossa, presence of a paraloph, location of the protocone on the anterolabial margin, promi- nent hypocone separated from the protocone by a distinct V-shaped notch in the postprotocrista, and an enlarged talon with a well developed, triangu- lar-shaped, posterolingual extension. Other dif- ferences between the M1s of O. floridanus and K. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 167 Ta bl e 4 . C om pa ra tiv e d en ta l c ha ra ct er s i n se le ct ed g en er a a nd sp ec ie s o f E m ba llo nu rid ae , i nc lu di ng fi ve ex tin ct sp ec ie s f ro m th e E oc en e, O lig oc en e, an d M io ce ne o f t he O ld W or ld , t hr ee e xt in ct sp ec ie s f ro m th e O lig oc en e an d M io ce ne o f t he N ew W or ld (F lo rid a) , a nd si x ex ta nt sp ec ie s, tw o fr om th e O ld W or ld a nd fo ur fr om th e N ew W or ld . T he fi ve e xt in ct O ld W or ld g en er a ar ea li st ed fi rs t, in o rd er fr om o ld es t/m os t p rim iti ve o n th e le ft (T ac hy pt er on ) t o yo un ge st o n th e rig ht (A fr ill on ur a) , a nd n ex t ar e th e th re e ex tin ct N ew W or ld g en er a, a ls o lis te d fr om o ld es t ( O lig op te ry x) to y ou ng es t ( Fl or id op te ry x) . T he e xt en de d po rti on o f T ab le 4 in cl ud es s ix li vi ng g en er a, Ta ph oz ou s a nd C ol eu ra fr om th e O ld W or ld a nd B al an tio pt er yx , D ic lid ur us , P er op te ry x, a nd S ac co pt er yx fr om th e N ew W or ld . C ha ra ct er s 1 -6 5 an d ch ar ac te r s ta te s i n th is ta bl e fo r T ac hy pt er on , V es pe rt ili av us , P se ud ov es pe rt ili av us , a nd D ho fa re lla a re fr om R av el e t a l. (2 01 6) . C ha ra ct er st at es fo r A fr ill on ur a ar e fr om R os in a an d Pi ck - fo rd (2 02 1) a nd c ha ra ct er st at es fo r O lig op te ry x, K ar st op te ry x, a nd F lo ri do pt er yx a re fr om th is st ud y. F or th e ne w c ha ra ct er s i nt ro du ce d in th is st ud y (C ha ra ct er s 6 6- 85 ), w e ha ve sc or ed th e fiv e ex tin ct O ld W or ld g en er a ba se d on th e lit er at ur e (e .g ., M ai tre , 2 01 4; R av el e t a l., 2 01 6; R os in a an d Pi ck fo rd , 2 02 1) . A m on g ou r n ew c ha ra ct er s, se ve ra l c an no t b e ev al ua te d in th e ex tin ct O ld W or ld g en er a be ca us e th es e ch ar ac te rs a re n ot il lu st ra te d or d es cr ib ed in th e lit er at ur e. C ha ra ct er s f or th e si x liv in g sp ec ie s ar e ba se d on e xa m in at io n of m us eu m sp ec im en s i n th is st ud y. S ee A pp en di x 2 fo r a m or e de ta ile d ex pl an at io n of th e ch ar ac te rs a nd c ha ra ct er st at es li st ed in th is ta bl e. Th e ch ar ac te r s ta te s f or e ac h ch ar ac te r a re li st ed a fte r t he v er ba l d es cr ip tio n of e ac h ch ar ac te r i n pa re nt he se s ( e. g. , 0 , 1 , 2 ). If a c ha ra ct er is n ot p re se rv ed in a p ar tic ul ar sp ec ie s be ca us e of in co m pl et e fo ss il m at er ia l, th e ch ar ac te r i s lis te d as “ ?” (e .g ., th e M 3 of F lo ri do pt er yx p oy er i i s no t p re se rv ed , s o ch ar ac te rs o f t he M 3, c ha ra ct er nu m be rs 6 2- 65 , a re li st ed a s “ ?” ). If a c ha ra ct er is a bs en t f ro m a p ar tic ul ar sp ec ie s, th e ch ar ac te r i s l is te d as “ –” (e .g ., Fl or id op te ry x po ye ri a nd a ll liv in g sp ec ie s o f E m - ba llo nu rid ae la ck th e p3 , s o ch ar ac te rs o f t he p 3, n um be rs 1 3 an d 14 , a re li st ed a s “ –“ ). A bb re vi at io ns : E ., Ea rly , M ., M id dl e, L ., La te ; N ., no rth ; S , s ou th ; S E, so ut he as t; pe n. , p en in su la ; a nt ., an te rio r/a nt er io rly ; p os t., p os te rio r/p os te rio rly ; l ab ., la bi al /la bi al ly ; l in g. , l in gu al /li ng ua lly . S ym bo ls : < le ss th an ; > g re at er th an ; = e qu al to . A ge , l oc at io n, a nd Ex tin ct G en er a of E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ch yp te ro n Ve sp er til ia vu s Ps eu do - D ho fa re lla Af ril lo nu ra O lig op te ry x Ka rs to pt er yx Fl or id op te ry x fra nz en i1 wi ng ei 2 ve sp er til ia vu s sig ei /D . t ha le ri 3 na m ib en sis flo rid an us 4 gu nn el li po ye ri pa rv a A ge M . E oc en e M . E oc en e- E. -M . E oc en e L. E oc en e/ M . M io ce ne E. -L . O lig oc en e L. O lig oc en e E. M io ce ne E. O lig oc en e E. O lig oc en e Lo ca tio n Eu ro pe Eu ro pe N . A fr ic a Eg yp t/ S. A fr ic a U SA U SA U SA G er m an y Fr an ce A lg er ia O m an N am ib ia Fl or id a Fl or id a Fl or id a C ha ra ct er s f ro m R av el e t a l. (2 01 6) D en ta ry 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) ? ? ? ? ? lo w (1 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) ? ? ? ? ? sh ar p (0 ) 3. H or iz on ta l r am us gr ac ile (0 ro bu st (1 ) ? ? ? ro bu st (1 ) ? ro bu st (1 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) or ie nt ed p os t. (1 ) ? ? ? or ie nt ed p os t. (1 ) ? or ie nt ed p os t. (1 ) Lo we r d en tit io n 5. L ow er in ci so r n um be r th re e (0 ) th re e (0 ) ? ? th re e (0 ) ? ? ? 6. L ow er p re m ol ar n um be r th re e (0 ) th re e (0 ) ? ? ? th re e (0 ) ? tw o (1 ) 7. L ow er in ci so r l ob es bi lo be d (1 ) ? ? ? ? ? ? ? 8. L ow er in ci so r r el at iv e siz e eq ui va le nt (1 ) ? ? ? ? ? ? ? 9. c 1 la bi al c in gu lu m str on g (1 ) str on g (1 ) ? ? str on g (1 ) ab se nt (0 ) ? ? 10 . c 1 ro ot /tu be rc le str ai gh t ( 0) str ai gh t ( 0) ? ? str ai gh t ( 0) str ai gh t ( 0) ? ? 11 . c 1 po ste rio r b as in re du ce d (1 ) ex te nd ed p os t. (0 ) ? ? ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) ? ? 12 . p 2 siz e re du ce d (0 ) no t r ed uc ed (1 ) ? ? no t r ed uc ed (1 ) no t r ed uc ed (1 ) ? ? 13 . p 3 ro ot s tw o (1 ) tw o (1 ) ? ? ? on e (0 ) ? – 14 . p 3 cr ow n sm al l ( 1) sm al l ( 1) ? ? ? sm al l ( 1) ? – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) ? ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) ? ? 17 . p 4 pa ra co ni d pr es en t ( 0) ab se nt (1 ) ? ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ? ? 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ? ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ? ? 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) ? hi gh (0 ) 20 . m 1 tri go ni d ? op en li ng ua lly (0 ) co m pr es se d (1 ) op en li ng ua lly (0 ) co m pr es se d (1 ) co m pr es se d (1 ) ? co m pr es se d (1 ) 21 . m 2 tri go ni d ? co m pr es se d (1 ) co m pr es se d (1 ) ? co m pr es se d (1 ) co m pr es se d (1 ) ? co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id d ist an ce ? m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) ? ? U pp er d en tit io n (c on t.) 63 . M 3 m et ac on e sm al l ( 0) sm al l ( 0) ? ? w el l-d ev el op ed (1 ) w el l-d ev el op ed (1 ) ? ? 64 . M 3 lin gu al c in gu lu m pr es en t ( 0) ab se nt (1 ) ? ? ab se nt (1 ) ab se nt (1 ) ? ? 65 . M 3 pr em et ac ris ta pr es en t ( 0) pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m ? w el l d ev el op ed (0 ) ? ? w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) ? ? 67 . c 1 an te ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 68 . c 1 po ste ro lin gu al c us p ? pr es en t ( 0) ? ? pr es en t ( 0) pr es en t ( 0) ? ? 69 . p 2 sh ap e ro un de d (0 ) el on ga te d (1 ) ? ? ro un de d (0 ) el on ga te d (1 ) ? ? 70 . p 4 sh ap e ro un de d (0 ) el on ga te d (1 ) ? el on ga te d (1 ) ro un de d (0 ) el on ga te d (1 ) ? ? 71 . m 1/ m 2 m or ph ol og y ? sim ila r, op en (0 ) ? di ff er en t ( 2) si m ila r, co m pr es se d (1 ) si m ila r, co m pr es se d (1 ) ? si m ila r, co m pr es se d 72 . m 1 tri go ni d ? op en li ng ua lly (0 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) m od . c om pr es se d (1 ) st ro ng ly c om pr es se d (2 ) ? st ro ng ly c om pr es se d (2 ) 73 . m 1/ m 2 pr ot oc on id /m et ac on id p la ce m en t ? pr ot oc on id a nt . t o m et ac on id a nt . t o sa m e le ve l ( 2) sa m e le ve l ( 2) m et ac on id a nt . t o ? m et ac on id a nt . t o m et ac on id (0 ) pr ot oc on id (1 ) pr ot oc on id (1 ) pr ot oc on id (1 ) 74 . m 1/ m 2 pr ot oc ris tid ? po st. -.l in g. in cl in ed tra ns ve rs e (0 ) tra ns ve rs e (0 ) tra ns ve rs e (0 ) an t.- lin g. -in cl in ed ? an t.- lin g. -in cl in ed fr om p ro to co ni d (1 ) fr om p ro to co ni d (2 ) fr om p ro to co ni d (2 ) 75 . m 1/ m 2 en to co ni d ? p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 171 Ta bl e 4. (E xt en de d) . A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) 172 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Ta bl e 4. (E xt en de d) . A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) A ge , l oc at io n, a nd Ex ta nt S pe ci es o f E m ba llo nu rid ae ch ar ac te rs O ld W or ld N ew W or ld Ta ph oz ou s Co le ur a Ba la nt io pt er yx D ic lid ur us Pe ro pt er yx Sa cc op te ry x m el an op og on af ra pl ic at a al bu s m ac ro tis bi lin ea ta A ge Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng Li vi ng G eo gr ap hi c di str ib ut io n In di a, S E A sia A fr ic a N . M ex ic o to S. M ex ic o to S. M ex ic o to S. M ex ic o to Ph ili pp in es Co sta R ic a So ut h A m er ic a So ut h A m er ic a So ut h A m er ic a C ha ra ct er s f ro m R av el e t a l. (2 01 6) M an di bu la r c ha ra ct er s 1. C or on oi d pr oc es s h ei gh t hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 2. C or on oi d pr oc es s a pe x sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) sh ar p (0 ) 3. H or iz on ta l r am us ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) ro bu st (1 ) gr ac ile (0 ) 4. C or on oi d pr oc es s a ng le ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) ve rti ca l ( 0) Lo we r d en tit io n 5. L ow er in ci so r n um be r tw o (1 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) th re e (0 ) 6. L ow er p re m ol ar n um be r tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) tw o (1 ) 7. L ow er in ci so r l ob es tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) tri lo be d (1 ) 8. L ow er in ci so r s iz e al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) al l e qu iv al en t ( 0) 9. c 1 la bi al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) th in /a bs en t ( 1) th in /a bs en t ( 1) 10 . c 1 ro ot str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) str ai gh t ( 0) 11 . c 1 po ste rio r b as in ex te nd ed p os t. (0 ) ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) re du ce d (1 ) 12 . p 2 siz e >p 4 (2 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) re du ce d (0 ) 13 . p 3 ro ot s – – – – – – 14 . p 3 cr ow n – – – – – – 15 . p 4 siz e w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 16 . p 4 ta lo ni d ex te nd ed p os t. (0 ) re du ce d (1 ) re du ce d (1 ) ex te nd ed p os t. (0 ) re du ce d (1 ) ex te nd ed p os t. (0 ) 17 . p 4 pa ra co ni d ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 18 . p 4 m et ac on id ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 19 . l ow er m ol ar tr ig on id c us ps hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) hi gh (0 ) 20 . m 1 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) 21 . m 2 tri go ni d lin g. o pe n (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 22 . m 1 pa ra co ni d/ m et ac on id >m et ac on id / m et ac on id / ½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) >½ w id th M 2 (0 ) 63 . M 3 m et ac on e ab se nt (2 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) 64 . M 3 lin gu al c in gu lu m ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) ab se nt (1 ) 65 . M 3 pr em et ac ris ta ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) N ew c ha ra ct er s ( th is st ud y) Lo we r d en tit io n 66 . c 1 lin gu al c in gu lu m w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) w el l d ev el op ed (0 ) 67 . c 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) ab se nt (1 ) pr es en t ( 0) 68 . c 1 po ste ro lin gu al c us p pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) pr es en t ( 0) 69 . p 2 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) el on ga te d (1 ) 70 . p 4 oc cl us al sh ap e el on ga te d (1 ) ro un d (0 ) ro un d (0 ) el on ga te d (1 ) ro un d (0 ) ro un d (0 ) 71 . m 1/ m 2 m or ph ol og y sim ila r- tri go ni d sim ila r- tri go ni d sim ila r- tri go ni d di ff er en t ( 2) sim ila r- tri go ni d sim ila r- tri go ni d op en (0 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) co m pr es se d (1 ) 72 . m 1 tri go ni d op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) op en li ng . ( 0) co m pr es se d (1 ) co m pr es se d (1 ) 73 . m 1/ m 1 pr ot oc on id /m et ac on id p la ce m en t pr ot oc on id a nt . ( 0) sa m e le ve l ( 2) m et ac on id a nt . ( 1) pr ot oc on id a nt . ( 0) m et ac on id a nt . ( 1) m et ac on id a nt . ( 1) 74 . m 1/ m 2 pr ot oc ris tid po st. -li ng . tra ns ve rs e (0 ) an t.- lin g. po st. -li ng . an t.- lin g. an t.- lin g. in cl in ed (1 ) in cl in ed (2 ) in cl in ed (1 ) in cl in ed (2 ) in cl in ed (2 ) 75 . m 1/ m 2 en to co ni d/ m et ac on id en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) en to co ni d sm al le r ( 0) eq ua l ( 1) 76 . m /m 2 hy pc on ul id sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) sm al l ( 1) w el l d ev el op ed (0 ) 77 . m 1/ m 1 hy po co ni d/ en to co ni d pl ac em en t hy po co ni d an t. (0 ) sa m e le ve l ( 2) sa m e le ve l ( 2) hy po co ni d an t. (0 ) en to co ni d an t. (1 ) sa m e le ve l ( 2) 78 . m 1/ m 2 po stc ris tid po st. -li ng . tra ns ve rs e (0 ) tra ns ve rs e (0 ) po st. -li ng . an t.- lin g. tra ns ve rs e (0 ) in cl in ed (1 ) in cl in ed (1 ) in cl in ed (2 ) 79 . m 1/ m 2 la bi al c in gu lu m th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) th in , c on tin uo us (1 ) 80 . m 3 ta lo ni d ve ry n ar ro w (0 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) br oa d (2 ) 81 . m 3 en to co ni d ab se nt (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) w el l d ev el op ed (1 ) U pp er d en tit io n 82 . C 1 an te ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 83 . C 1 po ste ro lin gu al c us p ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) ab se nt (1 ) pr es en t ( 0) pr es en t ( 0) 84 . M 1 pa ra sty la r r eg io n un re du ce d (0 ) re du ce d re du ce d re du ce d re du ce d re du ce d pa ra sty le a bs en t ( 2) pa ra sty le a bs en t ( 2) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) pa ra sty le p re se nt (1 ) 85 . M 1 pr ep ar ac ris ta w ea kl y re du ce d (0 ) ab se nt (2 ) ab se nt (2 ) ab se nt (2 ) sh or t ( 1) sh or t ( 1) MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 173 gunnelli are discussed below in the account of the latter species. Oligopteryx floridanus differs in many char- acters from the new genus and species of embal- lonurid Floridopteryx poyeri, described below from the early Miocene Thomas Farm LF. Ele- ments shared by these two extinct emballonurid species are the M1, m1–m3, dentary, and proximal femur. Perhaps the most important difference be- tween these two species is the presence of a small, single-rooted p3 in O. floridanus and the absence of a p3 in F. poyeri. As discussed below, all living genera in the Emballonuridae also lack a p3. The most obvious difference between these two species is that O. floridanus is much larger. A sample of 11 M1s of O. floridanus have a mean anteroposterior length of 1.80 compared to a single M1 of F. poyeri which has an anteroposterior length of only 0.85 mm, less than half that of the larger species. This difference in size is over-emphasized by the much shorter, truncated talon of F. poyeri compared to O. floridanus that has a well-developed, postero- lingual extension of the talon. Other differences that characterize the M1 of Floridopterx compared to Oligopteryx are the well-developed parastyle, weak paracingulum, bulbous anteriorly directed mesostyle, and reduced hypocone. Floridopteryx shares two of these characters with Karstopteryx, the more prominent parastyle and reduced hypo- cone. See the accounts of K. gunnelli and F. poyeri for more detailed comparisons between those two species. The difference in size between O. florida- nus and F. poyeri is also observed in the m1 and m2 but it is not as great as the difference in size between the M1s of these two species. The m1 and m2 (samples of these two teeth combined here) of O. floridanus are about 20% larger (mean from a sample of 22 m1/m2s, see Table 2), anteroposte- rior length, 1.75 mm; maximum transverse width (talonid), 1.32 mm, compared to two m1/m2s of F. poyeri, with a mean anteroposterior length of 1.46 and maximum transverse width (talonid) of 1.09 mm. Features of the m1 and m2 of Oligop- teryx are similar to those two teeth in Floridopteryx but there are minor differences. In Oligopteryx, the talonid is comparatively narrower, the metaconid is more posteriorly placed, and the protocristid is not as sharply angled from the protoconid to the metaconid. On the m3, the metaconid is located farther anteriorly in O. floridanus and accordingly the postcristid forms a more obtuse angle to the long axis of the tooth, whereas in Floridopteryx the metaconid is farther posterior and the protocristid is at a right angle to the long axis of the tooth. Comparisons with extinct genera of Embal- lonuridae from the Old World.—Dental characters of all eight extinct genera and six living genera of Emballonuridae are summarized in Table 4 and Ap- pendix 2. Oligopteryx differs in a number of char- acters from Tachypteron franzeni, one of the old- est known emballonurids, from the middle Eocene Messel site in Germany. Characters of Tachypteron are from descriptions and illustrations in Storch et al. (2002) and Smith et al. (2012). The parastylar region of M1 in Tachypteron is much better devel- oped than in Oligopteryx, which either lacks or has a very small parastyle and has a much reduced pre- paracrista. The M1 and M2 of Tachypteron have a transversely elongated (wider than long) occlu- sal outline, whereas the M1 of Oligopteryx has a more squarish occlusal outline and M2 is longer anteroposteriorly. M1 and M2 of Tachypteron lack a hypocone, whereas both teeth have a well-devel- oped hypocone in Oligopteryx. Both Oligopteryx and Tachypteron have a p3, distinguishing these two genera from all living emballonurids that lack a p3. However, this tooth is much larger and bet- ter developed in Tachypteron, with two roots and intermediate in size between the smaller p2 and larger p4; the premolars becoming progressively larger from p2 to p4. In contrast to Tachypteron, the p3 in Oligopteryx is a tiny, single-rooted tooth that is much smaller than p2. Labial cingula are very strong on the three lower premolars and three mo- lars in Tachypteron (Storch et al., 2002), much bet- ter developed than the labial cingula on the lower teeth of Oligopteryx. The M1 and M2 of Vespertiliavus, from the Eocene of Europe and northern Africa (Barghoorn, 1977; Maitre, 2014; Ravel et al., 2016), are simi- lar to one another in size and shape, whereas the M1 and M2 of Oligopteryx have distinctly differ- 174 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) ent morphologies, with M1 more quadrate in oc- clusal outline and M2 more transversely elongated. The M1 of Vespertiliavus has a well-developed parastyle and preparacrista, whereas the parasty- lar region is greatly reduced in Oligopteryx with the parastyle small to absent and the preparacrista very short. The M1 and M2 of Vespertiliavus have a very small or incipient hypocone that lacks a dis- tinct apex, whereas the hypocone of Oligopteryx is a prominent cusp separated from the protocone by a deep cleft in the postprotocrista. Vespertiliavus also differs from Oligopteryx in having a double- rooted p3 with obliquely oriented roots. Pseu- dovespertiliavus from the early to middle Eocene of Algeria (Ravel et al., 2016) is similar in most features to the upper molars of Vespertiliavus and thus differs similarly from Oligopteryx (Table 4). Pseudovespertiliavus further differs from Oligop- teryx in having M1 and M2 talons rounded distally rather than pointed, in lacking a connecting ridge between postprotocrista and hypoconal crest, and in having a stronger paraloph on M1. Sigé et al. (1994) described a new genus and species of emballonurid, Dhofarella thaleri, from the early Oligocene Taqah fauna in Oman on the Arabian Peninsula, and Gunnell et al. (2008) de- scribed a second, smaller species of Dhofarella, D. sigei, from a late Eocene fauna in the Fayum Depression of Egypt. Both species of Dhofarella are represented by small samples, including half a dozen isolated teeth of D. thaleri and a single man- dible with m1–m3 of D. sigei. Dhofarella thaleri is much smaller than Oligopteryx floridanus but similar in size to O. hamaxitos, whereas D. sigei is considerably smaller than both Florida species of Oligopteryx. Neither species of Dhofarella pre- serves the dentary anterior to the m1, so the pres- ence or absence of p3 in this genus cannot be deter- mined. The M1 of Oligopteryx differs from that of D. thaleri in the more strongly reduced parastylar region, shorter preparacrista, better-developed hy- pocone, and prominent triangular-shaped (rather than rounded) talon basin (Sigé et al., 1994). Com- pared to D. thaleri, the m1 of Oligopteryx has a more lingually placed paraconid, more anteriorly placed metaconid causing the protocristid to be an- gled anterolingually from protoconid to metaconid (protocristid is horizontal in D. thaleri), and more labially inflected entocristid. Besides its larger size, Oligopteryx differs from D. sigei in the similarity of m1 and m2; the m1 in D. sigei is narrower than the m2, especially the trigonid, and both the para- conid and metaconid are located more anteriorly (Gunnell et al., 2008). The entocristid is strongly inflected labially in Oligopteryx on the lower mo- lars, whereas the entocristid is rather straight in D. sigei. Gunnell et al. (2008) noted that among living emballonurids D. sigei is most similar to Coleura, the only extant emballonurine genus in Africa. Rosina and Pickford (2019) reported a large and diverse sample of fossil bats from the middle Miocene (~12–13 Ma) Otavi Mountain karst depos- its from northern Namibia in southwestern Africa. Remains of emballonurids are common in the Mio- cene chiropteran assemblage from the Berg Aukas I site of the Otavi Mountain karst, including both taphozoines and emballonurines (Rosina and Pick- ford, 2020). A small species of emballonurid from the Berg Aukas I site in Namibia was recently de- scribed as a new genus and species in the subfam- ily Emballonurinae, Afrillonura namibensis (Ro- sina and Pickford, 2021). The M1 of Afrillonura is considerably smaller (length, 1.40 mm; width, 1.80 mm) than the M1 of Oligopteryx floridanus (mean of 11 M1s: length, 1.80 mm; width, 2.38 mm), but is similar in size to O. hamaxitos (M1 paratype: length, 1.37 mm; width, 1.65 mm). The most no- table difference between the M1s of Afrillonura and Oligopteryx is that Afrillonura has a prominent rounded cusp or process at the anterolabial mar- gin of the tooth lacking in Oligopteryx. We iden- tify this cusp as the parastyle in our descriptions of emballonurid M1s, whereas Rosina and Pickford (2021, p. 6) identify this feature as a “pronounced cingulum expansion” where the ectoloph meets the precingulum (= paracingulum in our descriptions) at the anterobuccal (= anterolabial) margin of the M1. Other differences between the M1s of these two genera are the presence of a distinct metaloph, larger, more bulbous and more labial position of the metastyle, longer postmetacrista, and deeper V- shaped indentation in the metafossa in Afrillonura. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 175 Differences in the lower dentition include the more elongated p2 and p4 in Oligopteryx compared to Afrillonura in which these two premolars are short- er and have a more rounded occlusal outline and a better developed talonid on p4 in Oligopteryx. Comparisons with living genera of Embal- lonuridae.—Oligopteryx floridanus is similar in size to several living species in the Old World gen- era Taphozous and Saccolaimus (subfamily Tapho- zoinae), especially Taphozous melanopogon, but differs from taphozoines in many dental characters. The M1 and M2 of Taphozous and Saccolaimus are squarish in occlusal outline, but are more rectangu- lar in Oligopteryx, especially the M2, shorter an- teroposteriorly and wider transversely. Taphozoines have a well-developed parastyle and preparacrista on M1, unlike Oligopteryx in which the parastylar region of M1 is greatly reduced, with the preparac- rista shorter and the parastyle is small to absent. Taphozous and Saccolaimus lack a paracingulum and a distinct hypocone on M1 and M2; both fea- tures are well developed in Oligopteryx. The M3 is more reduced in taphozoines compared to Oli- gopteryx and to living New World emballonurines, consisting only of a small portion of the ectoloph including the parastyle, preparacrista, and para- cone and with a greatly reduced trigon basin and weak to absent protocone. The M3 of Oligopteryx is not nearly so reduced, with a postparacrista, me- sostyle, premetacrista, and metacone, lacking only the postmetacrista and metastyle on the ectoloph. In Taphozous and Saccolaimus, the lower molars are more elongated than in Oligopteryx, especially the m1, the paraconid and metaconid are well separated resulting from a lack of com- pression of the trigonid, and the metaconid is po- sitioned posterior to the protoconid, rather than anterior to the protoconid as in Oligopteryx. The protocristid on the m1 and m2 of taphozoines is perpendicular to the long axis of the tooth or may be directed slightly posteriorly from the protoconid to the metaconid, whereas in Oligopteryx the pro- tocristid is directed noticeably anteriorly from the protoconid to more anteriorly placed metaconid. In taphozoines, the talonid is slightly broader than the trigonid on m1 and m2 and very narrow and reduced on m3. In Oligopterx, the talonid is much broader than the trigonid on m1 and m2, and on m3 the talonid is not nearly so reduced as in taphozo- ines, only slightly narrower than the trigonid. The taphozoines have a straight entocristid on the lower molars that is not as high and lacks the deep, labial- ly oriented, V-shaped notch found in Oligopteryx. The differences in both the upper and lower molars between Oligopteryx and the living genera of taphozoines Taphozous and Saccolaimus are also observed in teeth of these two genera reported from several Late Cenozoic fossil sites in Africa and one in Australia. Two extinct species of Sacco- laimus have been described from Kenya: S. incog- nita from the early Miocene Koru fauna is known from a poorly preserved partial skull with broken P4 and M2 (Butler and Hopwood, 1957) and S. ke- nyensis from the early Pliocene Kanapoi fauna in the Turkana basin, which is known from a single M1 (Gunnell and Manthi, 2020). The M1 of Sac- colaimus kenyensis differs from that of Oligopteryx in the longer preparacrista, poorly developed para- cingulum, crestiform and posteriorly elongated mesostyle, and short postprotocrista (Gunnell and Manthi, 2020). M1s from the middle Miocene Berg Aukas 1 site in Namibia referred to both Saccolai- mus and Taphozous differ from Oligopteryx in their unreduced parastylar regions, prominent parastyle, longer preparacrista, nearly straight ectoloph lack- ing indentations in the parafossa and metafossa, lack of a hypocone, and broadly rounded talon basin (Rosina and Pickford, 2020). An M3 of Saccolai- mus from Berg Aukas 1 has a much more reduced M3 than Oligopteryx, lacking the postparacrista, premetacrista, and metacone. A mandible of Sac- colaimus from Berg Aukas 1 differs from Oligop- teryx in lacking a p3, having the m1 trigonid more elongated with the paraconid and metaconid more widely separated, the m1 and m2 with a straight entocristid, and the m3 with a greatly reduced, nar- row talonid (Rosina and Pickford, 2020). Both extant taphozoine genera, Taphozous and Saccolaimus, have been identified from the early Pleistocene Rackham’s Roost Site at the Riv- ersleigh World Heritage Area in Australia (King et al., 2020). Several M1s from Rackham’s Roost 176 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) were referred to the living Australian species Ta- phozous georgianus, differing from Oligopteryx in the better developed parastylar region, including the presence of a parastyle and longer preparacris- ta, absence of a hypocone, and rounded posterior margin of the talon basin (King et al., 2020). Sev- eral fossil mandibles of T. georgianus from Rack- ham’s Roost differ from Oligopteryx in lacking a p3 and having the m1 with a more elongated tri- gonid, a greater separation between the paraconid and metaconid, and metaconid located posterior to the protoconid, and a very narrow talond on m3. A single M2 referred to Saccolaimus from Rack- ham’s Roost differs from the M2 of Oligopteryx in the squarish overall shape, lack of a well-devel- oped ‘hooked’ parastyle, lack of a hypocone, and rounded posterior margin of the talon basin. Oligopteryx floridanus is larger than any liv- ing species in the Old World emballonurine genera Coleura, Emballonura, Mosia, and Paremballon- ura. Although these four genera differ from Oli- gopteryx in a number of dental characters, they are more similar to the Florida genus than are Ta- phozous and Saccolaimus, supporting the basic di- chotomy within the Emballonuridae between the Taphozoinae and Emballonurinae. Like Oligop- teryx but unlike taphozoines, the Old World embal- lonurines have a reduced parastylar region on M1. Compared to Oligopteryx, the Old World embal- lonurines have a better-developed parastylar shelf labial to the paracone and a much stronger paras- tyle at the anterolabial termination of the paracin- gulum but lack a preparacrista (the ectoloph ends abruptly at the paracone). The M1 and M2 of the Old World emballonurines are more squarish in oc- clusal outline and the trigon basin is compressed transversely. The M1 and M2 are transversely broader in Oligopteryx, primarily because of the broader trigon basin. The Old World emballon- urines have a distinct hypocone, but it is smaller and has a shallower notch in the postprotocrista separating it from the protocone than Oligopteryx. The lower premolars (p2 and p4, p3 is absent) in the Old World emballonurines are more rounded in occlusal outline, whereas the premolars are lateral- ly compressed, elongated and blade-like in Oligop- teryx. The trigonid on the lower m1 and m2 is not as anteroposteriorly compressed in Coleura, Em- ballonura, Mosia, and Paremballonura as it is in Oligopteryx. The Old World emballonurines have a deeper anterior portion of the dentary and lack the triangular-shaped flange extending posteriorly from the mandibular symphysis. With the excep- tion of Pleistocene records of Coleura from Ethio- pia and Paremballonura from Madagascar (Rosina and Pickford, 2021), the only other fossil record of a living genus of emballonurine from Africa is a lower m1 described as the extinct species Coleura muthokai from the late Pliocene of Ethiopia (Wes- selman, 1984). Like the living species C. afra, the trigonid of m1 in C. muthokai is not as anterposte- riorly compressed as in Oligopteryx. There is considerable dental variation among the eight genera of New World emballonurids in the monophyletic subfamily Diclidurinae. We compared each of the modern genera in this group to Oligopteryx; first to genera in the subtribe Di- clidurina (Balantiopteryx, Cormura, Cyttarops, Diclidurus, and Peropteryx) and then genera in the subfamily Saccopterygina (Centronycteris, Rhyn- chonycteris, and Saccopteryx). Characters for four of these genera, Balantiopteryx, Diclidurus, Per- opteryx, and Saccopteryx, are included in Table 4. The dental and mandibular features discussed for each genus are only those characters that dif- fer from Oligopteryx. In addition to their overall smaller size, especially when compared to Oligop- teryx floridanus, most living New World emballon- urids differ from Oligopteryx in having: the labial margins of the ectoloph on the upper molars with deep V-shaped notches (ectoflexi) in the the para- fossa labial to the paracone and the metafossa la- bial to the metacone; thin, high, and sharp cristae; deeply concave or pocketed trigon and talon basins with well-defined edges; and in lacking p3. Both living species of Balantiopteryx are smaller than Oligopteryx floridanus, whereas B. plicata is similar in size to the smaller species, O. hamaxitos. On M1 and M2, Balantiopteryx has a distinct paraloph and metaloph, O. floridanus has a weak paraloph and lacks a metaloph, and O. hamaxitos has a paraloph and metaloph. The M1 MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 177 of Balantiopteryx is squarish with a narrow tri- gon basin compared to Oligopteryx which is wider transversely with a broad trigon basin. The talon on M1 in Balantiopteryx is shorter, more rounded, and lacks the triangular-shaped, posterolingual ex- tension typical of Oligopteryx. The p2 and p4 are comparatively smaller and more rounded in oc- clusal outline in Balantiopteryx compared to the larger, laterally compressed lower premolars of Oligopteryx. The mandibular symphysis of Balan- tiopteryx is broader anteriorly and the mandibular flange extending posteriorly from the symphysis is weaker. The only species of Cormura, C. breviros- tris, is intermediate in size between the two spe- cies of Oligopteryx. The upper molars of Cormura and Oligopteryx are similar in certain characters; in both genera the parastylar region of M1 is re- duced, with a tiny preparacrista that curves poste- riorly, paralleling the postparacrista. Cormura dif- fers from Oligopteryx in having M1 and M2 with a weak to absent hypocone lacking a notch in the postprotocrista separating it from the protocone, and a rounded talon on M1 that lacks a posterolin- gual expansion. Compared to Oligopteryx, the p2 and p4 of Cormura are rounded in occlusal outline, the paraconid and metaconid on m1/m2 are located farther apart, the entocristid is less blade-like and not as sharply inflected labially, and the mandibu- lar flange below p4 is weaker. Cyttarops alecto, the only species in this genus, is similar in most of its dental features to Diclidurus. Compared to Oligopteryx, the M1 of Cyttarops has an anteriorly oriented preparacrista and lacks a prominent, triangular posterolabial ex- tension of the talon. Cyttarops differs from Oligop- teryx in having small and rounded p2 and p4, the lower molars with the paraconid and metaconid widely separated, especially on m1, and the meta- conid located posterior to the protoconid such that the protocristid is directed posterolingually from the protoconid to the metaconid. Diclidurus contains the largest living species of New World emballonurids. The largest species, D. ingens, is similar in size to Oligopteryx flori- danus; all species of Diclidurus are larger than O. hamaxitos. On M1 of Diclidurus, the parastylar shelf is better developed than in Oligopteryx, the preparacrista is anteriorly oriented, a small but dis- tinct parastyle is present, and the talon is rounded and lacks a posterolingual expansion. The hypo- cone on M1 and M2 of Diclidurus is smaller and is separated from the protocone by a weaker notch in the postprotocrista. Similar to Oligopteryx, the lower premolars of Diclidurus are transversely flattened and blade-like, but the p2 is relatively smaller in Diclidurus and the p4 has a prominent posterolabial cusp that is absent in Oligopteryx. The m1 and m2 of Diclidurus are more elongated, especially m1, the paraconid and metaconid are well-separated, and the metaconid is posterior to the protoconid. The mandibular symphysis of Di- clidurus is narrow anteriorly and has a tubercle at its posterior terminus. Compared to Oligopteryx, the parastylar shelf, parastyle, and preparacrista of M1 are better developed in Peropteryx, and the talon is squared off and lacks a triangular, posterolingual expan- sionThe p2 and p4 of Peropteryx are rounded, not narrow and blade-like as in Oligopteryx. The meta- conid in Peropteryx occupies an even more ante- rior position on m1 and m2 than in Oligopteryx, and the protocristid forms an even more oblique angle to the tooth, oriented anterolingually from the protoconid to the metaconid. The mandibular symphysis of Peropteryx is typical for emballon- urids, with minimal anterior narrowing and a small posterior flange. Compared to Oligopteryx, the M1 and M2 of Centronycteris have the ecotoloph on the labial margin deeply emarginated labial to the paracone (parafossa) and metacone (metafossa), a distinct metaloph, and a tall and sharp hypocone. A meta- loph is present on the M1 and M2 in O. hamaxitos. On the M1 of Centronycteris, the preparacrista is labially oriented and a parastyle is present, where- as the preparacrista is posteriorly oriented and the parastyle is absent or tiny in Oligopteryx. The p2 and p4 of Centronycteris are rounded in occlusal outline, not flattened and blade-like as in Oligop- teryx. The trigonids are compressed on the m1 and m2 in Centronycteris and the paraconid and meta- 178 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) conid are very close to one another, as close or even closer than in Oligopteryx. The mandibular ramus is very slender in Centronycteris, the symphysis is not shallower anteriorly, and there is a small flange extending posteriorly from the symphysis. Rhynchonycteris is tiny, much smaller than either species of Oligopteryx. Compared to Oli- gopteryx, the crests and cusps on the ectoloph of the upper molars are tall, sharp, and blade-like in Rhynchonycteris, the ectoloph along the labial margins of M1 and M2 is deeply notched labial to the paracone and metacone, the hypocone is partic- ularly tall and sharp, and the M1 has a large paras- tyle. On the lower m1 and m2 of Rhynchonycteris, the trigonid is compressed and the paraconid and metaconid are placed even closer together than in Oligopteryx. The horizontal ramus of the mandible in Rhynchonycteris is very slender and the sym- physis is relatively deep anteriorly. Compared to Oligopteryx, the M1 of Sac- copteryx has a better developed parastylar shelf and parastyle and the talon lacks a posterolingual expansion, the ectoloph on the M2 has deep, V- shaped emarginations in the parafossa and meta- fossa, and the paraloph and metaloph on the upper molars are distinct. A metaloph absent on M1 and M2 in O. floridanus, present in O. hamaxitos. The p2 of Saccopteryx is compressed and blade-like, the p4 is rounded; both premolars are comparative- ly smaller than in Oligopteryx. The paraconid and metaconid on m1 and m2 are slightly farther apart in Saccopteryx than in Oligopteryx and the pro- tocristid is nearly horizontal to the long axis of the tooth. The entocristid is U-shaped in Saccopteryx, not as high and sharply V-shaped as in Oligopteryx. The mandibular symphysis of Saccopteryx is deep- er anteriorly and has a small posterior flange. Remarks on Systematic Relationships.–Ac- cording to Barghoorn (1977), a reduced parastylar region on M1 is one of the derived characters unit- ing the four genera of Old World emballonurines (tribe Emballonurini), Coleura, Emballonura, Mo- sia, Paremballonura, and all New World members of the family (tribe Dicliurini), and together com- prising the subfamily Emballonurinae and separat- ing them from Taphozous and Saccolaimus (sub- family Taphozoinae). Oligopteryx floridanus is similar to emballonurines in the strongly reduced parastylar region on M1. Most genera of smaller emballonurids have some expression of a metaloph running lingually from the base of the metacone towards the protocone, thus separating the deep- ly concave trigon and talon basins. According to Barghoorn (1977), the presence of a metaloph di- viding the trigon and talon basins on the M1 and M2 is a primitive character. One of the most sig- nificant dental differences between the two species of Oligopteryx is in the development of the paralo- ph and metaloph on the M1 and M2. O. floridanus has a weak paraloph and lacks a metaloph on these two teeth compared to O. hamaxitos, which has both the paraloph and metaloph well developed. Karstoptyerx gunnelli lacks both a paraloph and metaloph on the only known M1. The hypocone has two distinct forms in the Emballonuridae. In the Emballonurinae, as well as Oligopteryx, the hy- pocone forms a distinct cusp at the termination of the postprotocrista. Taphozous, Saccolaimus, and Vespertiliavus do not exhibit the distinct apex of the hypocone, a derived feature according to Barg- hoorn (1977). OLIGOPTERYX HAMAXITOS new species Fig. 11-15 Holotype.—UF 157768, left dentary frag- ment with m1-m2. Brooksville 2 Local Fauna, late Oligocene (late early Arikareean), Florida. Paratypes.—UF 182808, right M1; UF 157786, right M2; UF 182792, distal end of hu- merus; UF 179983, proximal end of radius. All paratypes are from the Brooksville 2, LF, Florida. Referred Specimens.—Brooksville 2 Local Fauna.– UF 182880, right C1; UF 182911, right P4; UF 157774, right M2; UF 182803, left c1; UF 182817, 182869, left m1/m2 (2); UF 179936, proximal humerus; UF 179909, distal humerus; UF 179914, 179915, proximal radius (2); UF 179977, distal radius. MNI is 2 based on two left m1. NISP is 15. I-75 Local Fauna.—UF 121715, right C1; UF 16861, 121716, left C1 (2); UF 121714, left MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 179 distal humerus. MNI is 2 based on two left upper canines (C1). NISP is 4. Type Locality and Age.—Brooksville 2 Lo- cal Fauna, Hernando County, Florida, late early Arikareean (Ar2), late Oligocene. Occurrence.–Known only from the early Oligocene (Whitneyan) I-75 LF, Alachua County, Florida and the late Oligocene (early Arikareean) Brooksville 2 LF, Hernando County, Florida. Etymology.—hamaxitos (Greek), road, high- way; in reference to the original discovery of this species during the construction of Interstate High- way 75. Diagnosis.—Oligopteryx hamaxitos is very similar to Oligopteryx floridanus in most morpho- logical characters (see generic diagnosis above), and is separated from the latter species primarily by its much smaller size. O. hamaxitos can also be distinguished from O. floridanus by the pres- ence of a metaloph on M1 and M2. O. hamaxitos can be separated from Karstopteryx gunnelli by its smaller size, lack of a parastyle, and larger hy- pocone separated from the protocone by a distinct notch in the postprotocrista. Oligopteryx hamaxi- tos can be separated from all other emballonurids by the following combination of characters. M1 with parastyle reduced to absent, preparacrista very short, well-developed paracingulum, talon triangu- lar in shape with prominent posterolingual exten- sion; M1 and M2 labial margin of metafossa with shallow V-shaped indentation labial to metacone, deep V-shaped notch in postprotocrista separating protocone and hypocone, hypocone distinct and triangular-shaped, talon basin deeply concave and separated from metacingulum by distinct notch just posterior and lingual to base of metacone; M2 with well-developed hooked parastyle at anterolabial termination of paracingulum, shallow V-shaped indentation in labial margin of parafossa; m1 and m2 nyctalodont, talonid significantly broader and slightly longer than trigonid, trigonid compressed with paraconid and metaconid located close to- gether along lingual margin, metaconid anterior to protoconid, protocristid at angle to long axis of toothrow, entocristid blade-like and distinctly V- shaped with apex oriented labially and open lin- gually; dentary with triangular-shaped process pro- jecting ventrally below p4. morphologIcal descrIpTIons Oligopteryx hamaxitos from the Brooksville 2 and I-75 local faunas is much rarer than the larg- er O. floridanus from these same two sites. There are only 15 specimens of O. hamaxitos, 11 from Brooksville 2 (MNI of 2) and 4 from I-75 (MNI of 2). Several teeth present in O. floridanus are not represented in the sample of O. hamaxitos, in- cluding all lower premolars and M3/m3. There is a large enough sample to confirm that O. hamaxi- tos represents a distinct species, very closely re- lated to O. floridanus. The following descriptions and comparisons are not as detailed as those for O. floridanus because so many of the dental fea- tures are identical between the two species, except for the smaller size of O. hamaxitos. The M1 and m1 lengths and humeral midshaft diameter (Tables 1, 2) provide estimates of its body weight ranging from 7.3 to 11.7 g (by method of Gunnell et al., 2009). The comparisons above of O. floridanus with other fossil and modern emballonurids are at the generic level, and therefore differences from O. floridanus also pertain to O. hamaxitos. Upper dentition.–Compared to Oligopter- yx floridanus, the only upper tooth position not represented in the sample of O. hamaxitos is the M3. There is a single M1 of O. hamaxitos from Brooksville 2 (UF 182808, paratype; Fig. 11 C-D). The M1 of O. hamaxitos is very similar in over- all morphology to the large sample of M1s of O. floridanus from Brooksville. The parastylar region anterior and labial to the paracone is highly re- duced. A parastyle is absent and the preparacrista is extremely short, consisting of a barely discern- ible ridge, oriented labially from the paracone. The paracingulum (= precingulum) is well developed, with a slight convex bulge, not a cusp, at its an- terolabial termination. There is a very shallow, V- shaped emargination in the metafossa along the ectoloph on the labial margin. The paracone and metacone are oriented vertically, the protocone has an anterior orientation. The low, rounded proto- cone is located on the anterolingual margin. There 180 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) is a shallow notch in the postprotocrista, about two thirds of the distance between the protocone and hypocone, isolating the low, rounded hypocone. The hypocone is lingual to the protocone and pos- terior to the metacone. A paraloph and metaloph are well-developed. The metaloph separates the tri- gon and talon basins. The trigon basin is concave but rather shallow compared to other New World emballonurids. The talon basin is much deeper and better defined, being deepest posterior and lingual to the metacone. The prominent, triangular-shaped talon forms the posterolingual extension of the M1. The talon extends farther posteriorly than the metacingulum and is separated from it by a distinct V-shaped notch posterior to the base of the meta- cone. The narrow metacingulum (= postcingulum) extends from the base of the metacone to the meta- style. There are two M2s of Oligopteryx hamaxi- tos from Brooksville 2. UF 157786 (paratype; Fig. 11 A-B) is a complete tooth, UF 157774 is broken in half and lacks the apex of the metacone. The following description focuses on the differences between the M2 and M1. Characters not listed are identical to the M1 described in the previ- ous paragraph. The two M2s of O. hamaxitos are very similar, except for smaller size, to the larger sample of M2s of O. floridanus from Brooksville 2. One important difference is the presence in O. hamaxitos of a well-developed metaloph on the M2, extending anterolingually from the base of the metacone ending before reaching the postpro- tocrista and separating the trigon and talon basins. A metaloph is absent on the M2 in O. floridanus. A well-developed paraloph is present on the M2 in O. hamaxitos, extending from the base of the paracone to the base of the protocone. O. florida- nus also has a fairly well-developed paraloph on M2. The parastylar region of M2 in O. hamaxitos is much better developed than on the M1. There is a strong, curved parastyle that forms the anterola- bial corner of the tooth. The parastyle is located at the anterolabial termination of the well-developed paracingulum. The preparacrista is complete, con- necting the paracone to the parastyle. The ectoloph on the labial margin has shallow, sharply V-shaped notches in the parafossa labial to the paracone and metafossa labial to the metacone. The metacone is slightly taller than the paracone, the protocone is much lower. There is a deep, V-shaped notch in the postprotocrista, about halfway between the proto- cone and the well-developed hypocone. This notch is also present in the M1 but is not as prominent. The hypocone is a distinct, sharply pointed cusp about half the height of, and well lingual to, the protocone. The elliptical trigon basin is concave but shallow for an emballonurid. The talon basin is more rounded and much deeper, deepest posterior and lingual to the posterior base of the metacone. The talon basin extends farther posteriorly than the metacingulum and is separated from it by a shallow indentation that is not as prominent as the deep, V- shaped notch on M1. The talon is more rounded than on M1 and does not extend as far lingually. The metacingulum is narrow and indistinct. There is one upper canine (C1) of Oligop- Figure 11. Right upper molars of Oligopteryx hamaxitos from Brooksville 2 LF. A-B, UF 157786, M2, in occlusal (A) and lingual (B) views; C-D, UF 182808, M1, in occlusal (C) and lingual (D) views. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 181 teryx hamaxitos from Brooksville 2 (UF 182880; Fig. 12A-C). There are three C1s of a small embal- lonurid from I-75, the most abundant element of O. hamaxitos in this fauna. These canines are laterally compressed and lack an internal (lingual) cingulum. Basal cingula are almost entirely lacking. They have prominent anterior and posterior secondary cusps that are similar in size. The cusps are located on the anterolingual and posterolingual margins of the canines, respectively. No smaller cusps are present. These canines are similar to those of other emballonurids in overall shape (rounded or convex on the labial surface and flattened on the lingual surface) and possession of prominent spike-like anterior and posterior secondary cusps. In cross- section the canines form a hemicircle with the con- vex portion labial. The fossils differ from modern emballonurids in the total lack of an internal cin- gulum, lack of tiny secondary cusps, and a more triangular shape. These upper canines are smaller than those of O. floridanus described above, with a lesser angle between the root and crown, weaker lingual cingulum, and more distinct anterior sec- ondary cusp. The only other upper tooth locus represented in our sample of Oligopteryx hamaxitos is a P4 (UF 182911; Fig. 13) from the Brooksville 2 LF. The P4 of O. hamaxitos is similar to two P4s assigned to O. floridanus (Fig. 6), except for its smaller over- all size and relatively smaller talon. In labial and lingual views, the central cusp or protocone of the P4 is tall, narrow, and sharply triangular, almost caniniform in shape. The anterior margin is near- ly vertical, whereas the posterior margin is more gently curved. Prominent basal cusps are present at the anterolingual and posterolabial margins. The anterolingual cusp is slightly smaller, trian- gular, and in a more ventral position whereas the posterolabial cusp is larger, more rounded, and lo- cated dorsal to the anterolingual cusp. In occlusal view, the protocone is rounded, especially anteri- orly, and descends along its anterior margin almost vertically to the anterolingual cusp. Posteriorly, the protocone is connected to a narrow, blade-like crest that terminates at the posterolabial cusp. The P4 has a small lingual lobe or talon with a shallow Figure 12. Canines referred to Oligopteryx hamaxitos from Brooksville 2 LF. A-C, UF 182880, right C1 in occlusal (A), labial (B), and lingual (C) views. D-F, UF 182803, left c1 in lingual (D), labial (E), and occlusal (F) views. talon basin surrounded by a cingulum. The anterior cingulum surrounding the base of the protocone is separated from the talon by an anterolingual notch, with anterolingual cingular cusps on either side of 182 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 13. Right P4 referred to Oligopteryx hamaxitos from Brooksville 2 LF. UF 182911, right P4 in occlusal (A), lin- gual (B), and labial (C) views. the notch. The well-developed lingual cingulum descends gently from the anterolingual cusp to the posterior edge of the talon and is confluent with a posterior cingulum that ascends at a sharper angle from a notch at the posterior edge of the talon to the posterolabial cusp. In occlusal view, the pos- terior cingulum curves gently anterior labial to the talon and then curves back posteriorly to the pos- terolabial cusp A labial cingulum is absent. The P4 is three-rooted, with one root each ventral to the protocone, posterolabial crest, and talon. Lower dentition.—Among the tooth posi- tions in the lower dentition, only the m1, m2, and canine (c1) are represented in the sample of Oli- gopteryx hamaxitos from Brooksville 2. There are no lower teeth of O. hamaxitos from I-75. The type specimen of O. hamaxitos (UF 157768; Fig. 14) is a partial left dentary with the m1 and m2, as well as the alveolus for the posterior root of p4 and alveoli for both roots of m3. The m1 and m2 are very simi- lar in morphology, and when they occur as isolated teeth cannot be reliably separated. Therefore, the following description applies to both the m1 and m2 and is based primarily on UF 157768 (Fig. 14), together with data on dental variation from two ad- ditional isolated m1/m2s from Brooksvile 2 (UF 182817, 182869). Measurements of the m1 and m2 of O. hamaxitos are presented in Table 2. The tal- onid is broader than the trigonid on both m1 and m2. Because of the anteroposterior compression of the trigonid, the talonid is also slightly longer than the trigonid. Both teeth are nyctalodont, the postcristid connects the hypoconid and the much smaller hypoconulid. The paraconid and metaconid are located close together along the lingual mar- gin. The metaconid is located anterior to the pro- toconid and thus the protocristid is oriented at an angle to the long axis of the tooth row, extending from the more posterior protoconid anterolingually to the more anterior metaconid. The four lingual cusps are rather low, the metaconid is the tallest, followed by the entoconid and paraconid, with the much smaller hypoconulid lowest. The metaconid and entoconid are vertical, the paraconid is ori- ented slightly anteriorly. The labial cusps are taller than the lingual cusps, with the protoconid the tall- est cusp on the m1 and m2, followed by the hypo- conid. The cristid obliqua meets the trigonid at the lingual base of the protoconid. The high, sharp, an- gular entocristid has a distinct notch about halfway between the metaconid and entoconid. The apex of MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 183 the notch in the entocristid is oriented labially and is open lingually. The entocristid on the m1 of the type mandible penetrates somewhat more deeply into the talonid basin than does the entocristid on m2 on this same specimen. The small hypoconulid is located at the posterolingual corner of the tooth and is separated from the much larger entoconid immediately anterior on the lingual margin by a distinct notch or groove. The labial cingulum ex- tends from the paraconid to the hypoconid, but is weak compared to other emballonurids, especially labial to the protoconid and hypoconid. The labial cingulum is stronger on one of the isolated lower molars (UF 182869), than on the type. The anterior cingulum or mesial cingulum is well-developed with a small cuspid at its anterolingual termination. This cuspid is larger in the two isolated lower mo- lars. The postcingulum or distal cingulum is weak. The left c1 (UF 182803; Fig. 12 D-F) of Oli- gopteryx hamaxitos from the Brooksville 2 LF has a tall, slender main cusp with a narrow root. It has a less anteroposteriorly elongate and less transverse- ly narrowed crown base than those of O. florida- nus. It bears strong labial and lingual cingula with a prominent posterolingual secondary cusp and weaker posterolabial, anterolabial, and anterolin- gual cingular cusps. Between these cingular cusps, the cingulum shows an anterior indentation to ac- commodate the lower incisors and a posterior in- dentation to accommodate the p2. Dentary.—The holotype partial dentary (UF 157768) of Oligopteryx hamaxitos preserves the ventral margin of the horizontal ramus from below p4 to m2. The ramus has a straight ventral mar- gin below the two anterior molars. Ventral to the anterior alveolus of p4 there is a triangular-shaped Figure 14. Mandible of Oligopteryx hamaxitos from Brooksville 2 LF. UF 157768 (holotype), left dentary fragment with m1- m2 in occlusal (A) and lingual (B) views. 184 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) flange that is very similar to the flange extending posteriorly from the mandibular symphysis in O. floridanus. No specimens in the Brooksville sam- ple preserve the portion of the dentary that would reveal the presence (or absence) of a p3. The pres- ence of a p3, although tiny and single-rooted, is one of the most diagnostic features of O. floridanus. Humerus.—Both the proximal (UF 179936) and distal (UF 182792, paratype; 179909) ends of the humerus are represented in the sample of Oli- gopteryx hamaxitos from Brooksville 2. There is also a distal humerus from I-75 (UF 121714). Ex- cept for their much smaller size, these four speci- mens are very similar to proximal and distal humeri of O. floridanus from Brooksville 2. Measurements of the humerus and radius of O. hamaxitos are pre- sented in Table 3. The single proximal end of a humerus from Brooksville 2 referred to Oligopteryx hamaxitos (UF 179936) is somewhat damaged. In posterior view, the humeral head is elliptical in shape, some- what transversely flattened, rounded distally, and more pointed or triangular proximally. The head projects farther proximally than either the greater or lesser tuberosities. The head is not in the center of the shaft but is shifted noticeably laterally, locat- ed closer to the greater tuberosity than to the lesser tuberosity. The head is oriented at a slight angle to the shaft, canted distally toward the greater tu- berosity. The greater tuberosity is rather elongated, rounded at its proximal end, and oriented at about a 45° angle to the shaft. The lesser tuberosity is con- siderably larger than the greater tuberosity, forming a large, triangular-shaped process with a squared- off proximal end. The greater and lesser tuberosi- ties project about the same distance proximally. In proximal view, the pointed proximal portion of the head extends anteriorly to the base of the pectoral ridge. The larger size of the lesser tuberosity com- pared to the greater tuberosity is even more evident when viewed proximally. The lesser tuberosity is a prominent, triangular-shaped process extending medially, with a rounded anterior edge and straight posterior edge. It is about three times larger than the greater tuberosity. In this same view, the greater tuberosity is a narrow process with rounded edges, somewhat elongated in the anteroposterior dimen- sion, with a shallow indentation about halfway along its length. The supraglenoid fossa is very shallow. In lateral view, the pectoral ridge is rather short, broad, especially proximally, and triangular in shape. There is some variation in the shape of the pectoral ridge in the sample of proximal humeri from Brooksville 2; in most specimens this ridge is triangular but in several it is more rectangular. In medial view, the pectoral ridge bears a distinct, raised ridge that is oriented proximodistally, divid- ing the pectoral ridge into two approximately equal sections. There is a weak medial ridge on the shaft distal to the lesser tuberosity. The distal articular surface of the humerus of Oligopteryx hamaxitos (description based on UF 121714, 179909, 182792) is slightly offset from the shaft, with the lateral ridge of the capitulum lateral to the shaft (Fig. 15). The transversely narrow but proximodistally deep lateral ridge of the capitulum is separated by a very shallow groove from the bul- bous almost spherical medial ridge of the capitu- lum. The lateral ridge of the capitulum is the tallest portion of the distal articular surface, easily twice as high in the proximal-distal dimension as in the transverse dimension. The trochlea meets the me- dial ridge of the capitulum at about a 45° angle and the two facets are separated by a prominent groove. The trochlea is separated from the epitrochlea (me- dial epicondyle) by a deep rounded notch. The spinous process of the epitrochlea (distal spinous process) has a sharply triangular distal tip. The spi- nous process is somewhat less prominent than in O. floridanus. In medial view, the spinous process has a slightly more posterior orientation compared to O. floridanus in which this process is more vertical and parallel to the shaft. The radial fossa is shallow. Radius.—The proximal end of the radius of Oligopteryx hamaxitos is represented by three spec- imens from Brooksville 2, UF 179983 (paratype) and UF 179914, 179915. The proximalmost exten- sion of the radius has a gently rounded curvature, not triangular shaped as in most bats. The proximal articular surface consists of three slightly concave articular facets, corresponding to the three articular facets on the distal end of the humerus, the lateral MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 185 and medial ridges of the capitulum and the troch- lea. The facet for the medial ridge of the capitulum is a large, rounded concavity that occupies almost half of the proximal articular surface of the radius. The articular facet for the lateral ridge of the ca- pitulum is shallower and narrower transversely but is taller or deeper in the proximodistal direction. The articular facet for the trochlea is nearly flat and consists of a narrow, medially directed process that is blunt or squared-off on its medial surface. Distal to the articular surface for the trochlea is a deep pit, the flexor fossa. A single distal radius (UF 179977) of Oli- gopteryx hamaxitos is known from Brooksville 2. The distal end of the radius exhibits limited mor- phological characters. In anterior view, the distal edge of the articular surface is essentially straight, nearly horizontal to the shaft. There is a slight con- vexity representing the styloid process on the an- terolateral edge of the articular surface. In posterior view, the distal edge of the articular surface forms a 45° angle to the shaft, trending distally from lat- eral to medial. The distal articular surface is deeply concave where it articulates with the lunar bone of the carpus. Just proximal to the lateral edge of the distal articular surface on the posterior surface of the shaft is a prominent, triangular-shaped, later- ally oriented process that corresponds to the distal- most portion of the ulna comparIsons wITh oTher emBallonurIdae As noted above, Oligopteryx hamaxitos and O. floridanus are very similar, with the primary dif- ference between the two species being the much smaller size of O. hamaxitos (See comparative measurements of the teeth in Tables 1 and 2 and the humerus and radius in Table 3). Other than size, the most notable difference between these two spe- cies is the presence of a well-developed metaloph on M1 and M2 of O. hamaxitos. Upper M1s and M2s of O. floridanus generally lack the metaloph or have this character very weakly developed. In addi- tion to their small size, the upper canines (C1) of O. hamaxitos differ from those of O. floridanus in the Figure 15. Distal humerus of Oligopteryx hamaxitos from Brooksville 2 LF. UF 182792 in anterior (A), posterior (B), lateral (C), distal (D), and medial (E) views. 186 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) lesser angle between the root and crown, weaker lingual cingulum, and more distinct anterior sec- ondary cusp. The lower canine (c1) of O. hamaxi- tos is more anteroposteriorly elongate and has a less transversely narrowed crown base than lower ca- nines of O. floridanus. The spinous process on the distal end of the humerus in O. hamaxitos is some- what less prominent and has a slightly more poste- rior orientation than in O. floridanus. Differences between O. hamaxitos and. Karstopteryx gunnelli from the latest Oligocene Buda LF, are the same as the characters that distinguish K. gunnelli from O. floridanus, as discussed above in the account of the latter species. In addition, O. hamaxitos is smaller than K. gunnelli, and also differs in the presence of a paraloph and metaloph on M1, both of which are absent in K. gunnelli. Comparisons of morphologi- cal characters of O. floridanus with those of all oth- er extinct and extant genera of emballonurids (see above), including the genus Floridopteryx from the the early Miocene Thomas Farm LF, apply also to O. hamaxitos. Size comparisons of O. hamaxitos with other genera of emballonurids are also men- tioned above under the O. floridanus account. KARSTOPTERYX new genus Type Species.—Karstopteryx gunnelli. Included Species.—Only the type species is known. Diagnosis.—All characters pertain to the ho- lotype M1, the only tooth represented in the hypo- digm of Karstopteryx gunnelli. The M1 has a great- ly reduced anterolabial portion of the tooth, includ- ing the stylar shelf labial to the paracone, but has a well developed, rounded parastyle at the antero- labial terminus of the paracingulum. The preparac- rista is very short and oriented anteriorly. A weakly concave indentation is present in the metafossa labial to the metacone, not a sharply V-shaped in- dentation. The metacone is taller than the paracone. The paracingulum curves posterolingually from the parastyle to the protocone. The weak hypocone is not separated from the protocone by a distinct notch in the postprotocrista. A paraloph and meta- loph are absent. The trigon is triangular in shape not squarish. The talon has rounded lingual, labial, and posterior margins and lacks a deep, central pit. Etymology.—Karst (German), a type of to- pography formed by the dissolution of limestone, forming caves, sinkholes and other solution fea- tures, and pteryx (Greek), wing, a common ending for generic names in the family Emballonuridae. ‘Karst’ refers to the topography that character- izes much of northern peninsular Florida, where the limestone terrain has been extensively eroded forming numerous caves, sinkholes, fissures, and other karst-derived fossil deposits that have pro- duced bats, including not only the type locality, the Buda Quarry, but also the other three fossil local- ites described in this paper. KARSTOPTERYX GUNNELLI new species Fig. 4 G-H Holotype.—UF 97386, right M1 (Fig. 4 G-H) Buda Local Fauna, late Oligocene (early late Arikareean), Alachua County, Florida. This species is known only from the holotype. Type Locality and Age.—Buda Local Fauna, Buda Quarry, near Buda, about 7 km southwest of High Springs, Alachua County, Florida, early late Arikareean (Ar3), latest Oligocene. Occurrence.—Known only by the type spec- imen from the type locality. Etymology.—Named for the late Gregg F. Gunnell in honor of his many important contribu- tions to our knowledge of Tertiary Chiroptera. Diagnosis.—Same as for the genus. morphologIcal descrIpTIon A large emballonurid is represented in the Buda LF by a single, well-preserved M1 (UF 97386; Fig. 4 G-H), here described as the new ge- nus and species Karstopteryx gunnelli. This tooth is similar in size and morphology to the M1s of Oligopteryx floridanus, from Brooksville 2 and I-75 (Table 1), but there are several important dif- ferences. Although the anterolabial portion of the M1 of K. gunnelli is greatly reduced, in particular the stylar shelf labial to the paracone, preparacrista and postparacrista, there is a prominent parastyle with a rounded anterior projection at the anterola- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 187 bial terminus of the paracingulum. The parastyle is at the same level as the paracone and metacone. The preparacrista is very short and anteriorly ori- ented. The parafossa consists of a slight indenta- tion in the labial margin labial to the paracone and the metafossa forms a gently concave indentation labial to the metacone. The mesostyle is rounded, whereas the metastyle is narrow, compressed an- teroposteriorly, and extends somewhat farther labi- ally than the mesostyle. In lingual view, these two cusps are about the same height, with the mesostyle vertical and the metastyle oriented slightly poste- riorly. The paracone and metacone are similar in size and shape in occlusal view but the paracone is considerably lower in lingual view. The paracingu- lum (= precingulum) curves gently posteriorly and lingually, merging with the preprotocrista, and then meeting the protocone at the same level as the para- cone anteroposteriorly. The hypocone is weak and there is no V-shaped notch in the postprotocrista separating the hypocone from the protocone in lin- gual view. Both a paraloph and metaloph are ab- sent. The trigon basin is rather shallow and triangu- lar or V-shaped. The trigon and talon are separated by a low but distinct ridge that extends from the postprotocrista between the protocone and hypo- cone and is directed posterolabially to the poste- rior base of the metacone. The talon has rounded lingual, labial, and posterior margins and lacks a deep, central pit. The posterolabial margin of the talon extends farther posteriorly than the metacin- gulum (= postcingulum) and is separated from it by a distinct, V-shaped notch. comparIsons wITh oTher emBallonurIdae The Buda emballonurid M1 (UF 97386), described here as Karstopteryx gunnelli, shares characters with the M1s of Oligopteryx floridanus and O. hamaxitos from Brooksville 2, in particu- lar the overall reduction of the anterolabial portion of the tooth, including the stylar shelf labial to the paracone, and the greatly reduced preparacrista. However, the Buda M1 differs from the two spe- cies of Oligopteryx in several important features, the most obvious being the more prominent para- style, which is larger in the Buda tooth than in all but one M1 of O. floridanus (UF 182787). Another difference is the anterior orientation of the tiny pre- paracrista in K. gunnelli. The preparacrista is also greatly reduced in O. floridanus and O. hamaxitos but is more posteriorly oriented and parallel to the postparacrista. The ectoflexus on the labial mar- gin of the Buda M1 has a very slight indentation in the metafossa labial to the metacone in K. gun- nelli, whereas the two species of Oligopteryx have a stronger V-shaped emargination in the metafossa of M1. In lingual view, the paracone is lower than the metacone in K. gunnelli, whereas these two cusps are the same height in O. floridanus and O. hamaxitos. The paracingulum curves gently pos- terolingually to meet the protocone in K. gunnelli, with the protocone located directly lingual to the paracone. In the two species of Oligopteryx, the paracingulum on M1 does not curve posteriorly but forms the rather straight anterior margin horizontal to the long axis of the tooth, with the protocone in a more anterior position at the anterolingual cor- ner of the M1, slightly anterior to the paracone. The M1 of K. gunnelli has a very weak hypocone and lacks a distinct notch in the postprotocrista between the hypocone and protocone, whereas O. floridanus and O. hamaxitos have a more promi- nent hypocone and a deep V-shaped notch in the postprotocrista separating the hypocone and proto- cone. A paraloph and metaloph are both lacking on the M1 of K. gunnelli, whereas O. hamaxitos, has both the paraloph and metaloph well developed on this tooth, and O. floridanus has an intermediate condition, possessing a weak paraloph but lack- ing a metaloph. The trigon is somewhat smaller and triangular-shaped in K. gunnelli, compared to the more squarish trigon in O. floridanus and O. hamaxitos. The anterior margin of the trigon in K. gunnelli consists of the paracingulum that curves posterolingually from the prominent parastyle, merges with the preprotocrista, and then meets the protocone. The posterior margin of the trigon con- sists of a low, weak ridge that extends from the pos- terior base of the metacone anterolingually to meet the postprotocrista between the protocone and hy- pocone. The trigon is somewhat larger in O. flori- danus and O. hamaxitos and squarish not triangular 188 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) in shape, particularly the anterior and lingual mar- gins. Compared to the two species of Oligopteryx from Brooksville 2 and I-75, the talon of K. gun- nelli is smaller and has rounded posterior, lingual, and labial margins, with the anterolabial portion of the talon meeting the lingual end of the metac- ingulum posterior to the base of the metacone. In O. floridanus and O. hamaxitos, the talon of M1 is more angular and noticeably larger than in K. gun- nelli in both the anteroposterior and labiolingual dimensions. The talon of the two species of Oli- gopteryx from Brooksville 2 and I-75 has a well- developed, triangular posterolingual projection and the posterior margin trends slightly anterolingually from the posterolingual corner to the anterolabial corner located posterior and labial to the end of the metacingulum and slightly labial to the base of the metacone. The differences between the single M1 of K. gunnelli and the M1s of O. floridanus and O. hamaxitos are significant and indicate this tooth represents a distinct genus and species of embal- lonurid. The M1 of Karstopteryx gunnelli is con- siderably different from the M1 of Floridopteryx poyeri, the youngest emballonurid from the Ce- nozoic of Florida from the early Miocene (early Hemingfordian, He1) Thomas Farm LF (see next species account). The single M1 of Floridopteryx is damaged, with the posterolabial portion of the tooth missing. F. poyeri is a small bat, with the anteroposterior length of M1 (0.85 mm) less than half that of the much larger K. gunnelli (1.75 mm). Both Karstopteryx and Floridopteryx have a well- developed parastyle, although this feature is larger in the Buda M1 with a more prominent, rounded anterior projection. The labial margin and labial cusps of the M1 differ between the two species. K. gunnelli has a well-developed indentation in the parafossa labial to the paracone, a rather small me- sostyle, and a weak indentation in the metafossa labial to the metacone, whereas Floridopteryx has a weak indentation in the parafossa, a large, bul- bous mesostyle, and appears to have a deeper in- dentation in the metafossa, although the metacone, premetacrista, and labial margin of the tooth poste- rior to the mesostyle are damaged. The protocone is more anteriorly placed in Floridopteryx, at the anterolabial corner of the M1 somewhat anterior to the paracone, whereas the protocone is more pos- terior in Karstopteryx directly lingual to the para- cone. A weak paraloph and metaloph are present in Floridopteryx, whereas these structures are absent in Karstopteryx. The lingual portion of the M1 is more rounded in Karstopteryx, especially the an- terior and posterior margins, whereas the lingual half of this tooth is more squarish in Floridopteryx, with both the anterior and posterior margins essen- tially straight and tranverse to the long axis of the tooth. The talon is considerably longer in Karstop- teryx with a rounded posterior margin, whereas the shorter talon in Floridopteryx is truncated posteri- orly with a straight posterior margin. Karstopteryx gunnelli differs from other ex- tinct genera of emballonurids from the Old World and living genera of emballonurids from both the Old World and New World in almost all of the same characters as discussed above under the Oli- gopteryx floridanus account. FLORIDOPTERYX new genus Type Species.—Floridopteryx poyeri. Included Species.—Only the type species is known. Diagnosis.—The m1/m2 are nyctalodont, the talonid is much broader than the trigonid, the talonid is also longer than the trigonid because of the anteroposterior compression of the trigonid, the paraconid and metaconid are close together along the lingual margin, the metaconid is anterior to the protoconid, the protocristid trends anterolingually from the protoconid to the metaconid, the parac- ristid is gently curved, the paraconid, metaconid, and entoconid are conical with rounded apices and vertically oriented, the protoconid and hypoconid are larger than the lingual cusps with sharply point- ed apices, and the hypoconulid is rather large. The m3 has the trigonid broader than the talonid, the paraconid and metaconid are located close to one another along the lingual margin but both cusps are more posterior than on m1/m2, the paracristid and protocristid meet at a more acute angle than on MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 189 m1/m2, the protoconid and metaconid are located at the same level and the protocristid is at a right angle to the long axis of tooth. The dentary has a long mandibular symphysis extending posteriorly to the p4 and with a rounded projection ventral to the p2 and p4, a p3 is absent, the anterior alveolus of the m1 is flattened anteroposteriorly, a large el- liptical mental foramen is present ventral to the p2, a tiny mental foramen is present on the symphysis ventral to the incisor alveoli, the ascending ramus is upturned dorsally posterior to the toothrow, the coronoid process is short and slender dorsally, the angular process is at the same level as the alveo- lar margin of the toothrow and flares laterally, the articular process is dorsal to the alveolar margin, and the mandibular foramen is large, ventral to the toothrow, and below the tip of coronoid. The M1 has the parastylar region highly reduced anterior and labial to paracone, a well-developed parastyle is present at the anterolabial termination of the narrow paracingulum, the preparacrista very short and oriented labially, the mesostyle is bulbous and oriented anteriorly, the hypocone is small and not separated from the protocone by a notch, and the talon is squarish and somewhat truncated antero- posteriorly. Etymology.—Named for the state of Florida and pteryx (Greek), wing, a common ending for ge- neric names in the family Emballonuridae. FLORIDOPTERYX POYERI new species Fig. 16-19 Holotype.—UF 121132, right m1 Thomas Farm LF, early Miocene (early Hemingfordian), Gilchrist County, Florida. Paratypes.—UF 108661, complete eden- tulous right mandible with alveoli for i1-m3; UF 121134, partial right M1. Both paratypes are from Thomas Farm. Referred Specimens.—UF 121133, right m1 or m2; UF 108664, right m3; UF 121135, proximal end of left femur. All referred specimens are from Thomas Farm. MNI is 1. NISP is 6. Type Locality and Age.—Thomas Farm LF, 12 km NE of Bell, Gilchrist County, Florida, early Miocene, early Hemingfordian (He1). Occurrence.—Floridopteryx poyeri is known only from the Thomas Farm LF. Etymology.—Named for Arthur Poyer from the FLMNH in recognition of his important contri- butions to the study of Florida Cenozoic microver- tebrates, in particular screenwashing and sorting bats from Thomas Farm. Diagnosis.—Same as for the genus. morphologIcal descrIpTons Upper dentition.—Only one upper tooth of Floridopteryx poyeri was identified in the Thomas Farm sample, a shattered but nearly complete right M1 (UF 121134, paratype; Fig. 16), typically the most diagnostic tooth in emballonurids. The tooth is rather heavily worn, especially its lingual half. The posterolabial corner of the tooth is broken off and missing, including the tip of the metacone, metastyle, postmetacrista, part of the premetacris- ta, and the metacingulum (= postcingulum). As is typical of all other New World emballonurids, the anterolabial portion of the M1 is greatly reduced, in particular the stylar shelf labial to the paracone, preparacrista, and postparacrista (= parafossa or parastylar fovea). However, a fairly well-devel- oped, isolated parastyle is present at the antero- labial termination of the narrow paracingulum (= precingulum). The preparacrista is very short and oriented almost directly labially or slightly anteri- orly. The ectoloph on the labial margin of the M1 between the parastyle and mesostyle is gently con- cave with a slight indentation in the parafossa just anterior to the mesostyle. The mesostyle is a rather prominent bulbous cusp with a distinct anterior orientation. The lingual half of the M1 is squarish in shape, with both the anterior and posterolingual margins of the tooth relatively straight and trans- verse or horizontal to the anteroposterior axis of the tooth. Because of fairly heavy wear on the lin- gual portion of the M1, some of the following den- tal features may have been affected by wear. The narrow paracingulum extends lingually from the parastyle along the anterior margin of the M1, con- necting with the preprotocrista and protocone. The protocone is located at the anterolingual corner of 190 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 16. Upper molar of Floridopteryx poyeri from Thomas Farm LF. UF 121134, right M1 in occlusal view, photograph and line drawing. the tooth, slightly anterior to the paracone. The hy- pocone is barely distinguishable as a low rounded convexity along the posterolingual margin of the tooth. There is no notch in the postprotocrista sepa- rating the protocone from the weakly developed hypocone. Deep concavities or pockets are lacking in both the trigon and talon basins. A paraloph and metaloph are present but weak, although they may have been reduced by wear. The talon is squarish in shape, with a straight posterior margin and gen- tly convex lingual margin. The lingual margin of the tooth trends slightly posterolingually from the more labially positioned protocone to the postero- lingual corner of the talon. Lower dentition.—There are three isolated lower molars of Floridopteryx poyeri from Thom- as Farm, two m1 or m2 (UF 121132, holotype; UF 121133) and an m3 (UF 108664) (Fig. 17; Table 2). We identify the holotype (UF 121132) as an m1 because it was found in close association with a dentary of F. poyeri (UF 108661), described below, and the tooth is an exact fit for the m1 alveolus of this dentary. As discussed above under Oligopter- yx, it is difficult to separate isolated m1s and m2s of emballonurids, so we describe UF 121132 and 121133 together. The m3 is described in the fol- lowing paragraph. Using the length of the holotype m1, we predict an estimate of the body weight of F. poyeri of 12.6 g (by method of Gunnell et al., 2009). The holotype of Floridopteryx poyeri (UF 121132) is virtually unworn, UF 121133 is some- what more heavily worn. The talonid is much broader than the trigonid, suggesting that both UF 121132 and 121133 are probably m1s. In living New World emballonurids, on m1 the talonid is generally much broader than the trigonid, on m2 it is only slightly broader than the trigonid, and on m3 the trigonid is broader than the talonid. The talonid is also longer than the trigonid because of the anteroposterior compression of the trigonid, re- flected in the closely placed paraconid and meta- conid along the lingual margin. The metaconid is positioned anterior to the protoconid, such that the protocristid angles anterolingually from the proto- conid to the metaconid. The paracristid is gently curved, with a deep V-shaped notch about half- way between the paraconid and protoconid. There is also a deep V-shaped notch in the protocristid halfway between the protoconid and metaconid. The cristid obliqua meets the trigonid directly pos- terior to the tallest point on the protoconid in UF 121132 but is a bit more lingually placed on UF 121133, meeting the trigonid at the lingual base of the protoconid. In occlusal view, the entocristid is broadly U-shaped on UF 121132 but more sharply V-shaped and penetrates somewhat deeper into the talonid basin on UF 121133. The two m1/m2s of Floridopteryx are nyctalodont with the postcristid connecting the hypoconid to the hypoconulid. The hypoconulid is a rather prominent, posteriorly ori- ented cusp on the extreme posterolingual corner of the tooth immediately behind the entoconid. In lin- gual view, the paraconid, metaconid, and entoconid are conical with a rounded apex, vertically orient- ed, and similar in size, with the metaconid and en- toconid almost identical in size and the paraconid slightly smaller. In labial view, the protoconid and hypoconid are similar in height, with the protoco- nid slightly taller; both are easily twice as tall as the three main lingual cusps. The protoconid and hypoconid are also more sharply pointed than the lingual cusps. The anterior cingulum is well-devel- oped, the labial cingulum is not particularly strong, MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 191 and the posterior cingulum is weak. The roots on UF 121132 are intact, with the anterior root flat- tened anteroposteriorly and the posterior root rounded in cross-section. The shape of the roots in this tooth match the shape of the alveoli of m1 in an edentulous mandible (UF 108661) referred to Flo- ridopteryx poyeri (see dentary description below), in particular the noticeably flattened anterior root, strongly suggesting that the holotype (UF 121132) is an m1. A right lower molar (UF 108664) referred to Floridopteryx poyeri from Thomas Farm is identi- Figure 17. Lower molars of Floridopteryx poyeri from Thomas Farm LF. A-C, UF 121132 (holotype), right m1 or m2 in oc- clusal (and slightly labial, A), labial (B), and lingual (C) views; D-F, UF 121133, right m1 or m2 in occlusal (D), labial (slightly posterior, E), and lingual (slightly posteroventral, F) views; G-I, UF 108664, right m3 in occlusal (G), labial (H), and lingual (I) views. 192 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 18. Floridopteryx poyeri from Thomas Farm LF. UF 108661, edentulous right dentary, in alveolar (A), lingual (B), and labial (C) views. The dentary of Floridopterx lacks teeth but is otherwise almost perfectly intact, except for miss- ing the tips of the coronoid and angular processes (Fig. 18). It is similar to the dentary of extant em- ballonurids in having a comparatively elongated mandibular symphysis, reduced coronoid process, and laterally flaring angular process. Dentaries of vespertilionids, the most common Thomas Farm bats in the same size range, generally have a short- er symphysis, tall, rounded coronoid process, and smaller posteriorly oriented angular process. In dorsal aspect, the horizontal ramus of Floridopterx poyeri is straight from the m3 anterior to the m1 and then curves gently inward (lingually) toward the symphysis at about the level of p4. The mandibular symphysis is long, extending from the anterior tip of the dentary posteriorly to a point ventral to the anterior alveolus of p4. Ventral to the p2 and p4, a gently rounded ventral projection forms the poste- riormost extension of the mandibular symphysis. The symphyseal region and ventral margin of the horizontal ramus anterior to m1 form a gentle ob- tuse angle. Between p4 and m3 the ventral margin of the horizontal ramus is essentially straight, with only a slight ventral curvature below m3. There are fied as an m3 based on the broader trigonid com- pared to the talonid (Fig. 17 G-I). This tooth is identified as an emballonurid by the tall, narrow, V-shaped, labially directed entocristid. The paraco- nid and metaconid are located close to one another along the lingual margin, although both cusps are more posteriorly positioned than on m1/m2. Be- cause of the more posterior location of the paraco- nid and metaconid, the paracristid and protocristid meet at a more acute angle than on m1/m2. The paracristid is straight on m3, more curved on m1/ m2. The protoconid and metaconid are located at the same level, with the protocristid transverse to the long axis of the tooth. Although the entocristid is sharply V-shaped, it is slightly shallower than on m1/m2. The cristid obliqua meets the trigonid in a more lingual position than on m1/m2, about half- way between the protoconid and metaconid, form- ing a more acute angle. The hypoconulid is tiny. In lingual view, the entoconid and metaconid are similar in height, vertically oriented, and conical with a rounded apex, the slightly lower paraconid is more anteriorly oriented. In labial view, the pro- toconid is considerably taller than the hypoconid. The anterior cingulum and labial cingulum are strong; the posterior cingulum is rather weak. Dentary.—A nearly complete edentulous right dentary (UF 108661, paratype; Fig. 18) from Thomas Farm was collected on the same day and from the same sample of screenwashed concentrate as the holotype of Floridopteryx poyeri, an isolated m1 described above (UF 121132). The tooth is an exact fit in the empty alveolus for the m1 in the dentary. It seems unlikely this is a coincidence be- cause emballonurids are otherwise extremely rare from Thomas Farm (six specimens from a sample of more than 3,000 bat fossils). The dentary and tooth are almost certainly from the same individual but became separated during the screenwashing process. However, we have given these specimens separate catalog numbers because we cannot be ab- solutely certain they belong to the same individual, although without question they belong to the same species, Floridopteryx poyeri. We have arbitrarily chosen the m1 (UF 121132) as the holotype and the dentary (UF 108661) as a paratype. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 193 two mental foramina. The posterior mental foramen is large, deep, elliptical in shape, and located ven- tral to the p2, about one-third the distance between the alveolar and ventral margins of the horizontal ramus. A tiny mental foramen is located much far- ther anteriorly almost on the mandibular symphy- sis, immediately ventral to the alveoli for the in- cisors. The canine alveolus is slightly elliptical in shape, longer in the anteroposterior dimension. The single alveolus of p2 is round and about half the size of the canine alveolus. There is no alveolus for a p3, in agreement with modern New World em- ballonurids that also lack a p3, but differing from Oligopteryx that possesses a tiny, single-rooted p3. The p4 is double-rooted with the posterior alveolus rounded and anterior alveolus somewhat flattened along the anterior margin and set at a slight angle to the toothrow. There is no diastema between c1 and p2 and a very short diastema between p2 and p4. Five of the six molar alveoli are rounded in shape, but the anterior alveolus of the m1 is distinctly flattened anteroposteriorly. An examination of the well-preserved and complete roots of the m1 of Floridopteryx poyeri that we believe is associated with this dentary (UF 121132, holotype), confirms that the anterior root is flattened anteroposteriorly while the posterior root is round in cross-section. As is typical of emballonurids, the ascending ramus of the dentary is upturned dorsally posterior to the toothrow in Floridopteryx, although not to the degree observed in certain other members of this family such as Peropteryx and Saccopteryx. The tip of the coronoid process is missing but the base indicates that it was clearly rather short and slender dorsally as in emballonurids, and unlike the taller, broader, more rounded coronoid of most vespertilionids. The angular process is located at about the same level as the alveolar margin of the mandibular toothrow and, although its tip is bro- ken, it has a distinct lateral orientation. In dorsal view, the angular process flares noticeably lateral to the lateral edge of the horizontal ramus. The ar- ticular process or condyle is intact and is located dorsal to the alveolar margin of the toothrow. In posterior view, the articular process has a flat to slightly concave dorsal articular surface, a convex ventral margin, and is comparatively deep in the dorsoventral dimension. The mandibular foramen on the lingual surface of the dentary is large, deep, elliptical in shape, oriented anteroposteriorly, and is located just ventral to the toothrow below the tip of the coronoid. In Peropteryx and Saccopteryx, the mandibular foramen is much smaller, more round- ed, and is located dorsal to the alveolar margin of the toothrow, reflecting the more strongly dorsally upturned ascending ramus in these two genera. Femur.—A single proximal end of a left fe- mur (UF 121135) referred to Floridopteryx poyeri is the only emballonurid postcranial element iden- tified in the Thomas Farm sample (Fig. 19). This specimen is similar to, although somewhat smaller than, a proximal femur of Oligopteryx floridanus from Brooksville 2 described above. The femoral head is generally spheroidal but is slightly flat- tened proximally with a well-developed, rounded fovea capitis. The head is located in the center of the shaft but is slightly canted or angled medially and has a distinct neck separating it from the shaft. The lesser trochanter is slightly larger and extends farther proximally than the greater trochanter. The lesser trochanter is elongated with a rather pointed tip, while the greater trochanter is shorter with a blunt squarish tip. There is a well-developed me- dial ridge on the medial edge of the shaft distal to the lesser trochanter. comparIsons wITh oTher genera oF emBallonurIdae We compared Floridopteryx poyeri from Thomas Farm to all 14 living genera and the seven other extinct genera of Emballonuridae, Tachyp- teron, Vespertiliavus, Pseudovespertiliavus, Dho- farella, and Afrillonura from the Old World, and Oligopteryx and Karstopteryx from the Oligocene of Florida (see list of modern comparative material examined in Appendix 1). Dental characters of se- lected extinct and living species of Emballonuridae are presented in Table 4 and Appendix 2. Among other extinct genera of emballonurids, we consider the comparisons of Floridopteryx with Oligopter- yx and Karstopteryx to be the most critical based on their close geographic occurrence and possible 194 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Figure 19. Floridopteryx poyeri from Thomas Farm LF. UF 121135, left proximal femur, in ventral (A), anterior (B), dorsal (C), posterior (D), and proximal end (E) views. phylogenetic relationship. These are the only three described extinct genera of emballonurids from the New World. Features that distinguish the M1 of Floridop- teryx from the two species of Oligopteryx, O. flori- danus and O. hamaxitos, are the well-developed parastyle, weak paracingulum, bulbous anteriorly directed mesostyle, weak hypocone, shallow con- cavities in the trigon and talon basins (may be par- tially obscured by heavy wear), and the truncated and squarish talon, especially the posterior margin. Floridopteryx shares several of these features, in- cluding a well-developed parastyle and reduced hypocone not separated from the protocone by a notch in the postprotocrista, with Karstopteryx gunnelli from the latest Oligocene Buda LF. Com- pared to F. poyeri, the M1 of K. gunnelli is much larger and has a more prominent parastyle with a rounded anterior projection. The characters of the M1 shared by F. poyeri and K. gunnelli but not the two species of Oligopteryx, suggest the possibility that Floridopteryx may have evolved from a spe- cies similar to K. gunnelli in the early Miocene (lat- est Arikareean or earliest Hemingfordian). The lower molars of Floridopteryx and those of Oligopteryx floridanus and O hamaxitos are sim- ilar, but there are differences. Both genera have the talonid broader than the trigonid on m1 and m2, but the talonid is relatively broader in Floridopteryx. The trigonid is compressed anteroposteriorly with the paraconid and metaconid close together along the lingual margin in both genera, but the metaco- nid is more anterior in Floridopteryx. Because of the more anterior location of the metaconid in Flo- ridopteryx, the protocristid is even more strongly angled anterolingually from the protoconid to the metaconid. The entoconid is located more anteri- orly, the hypoconulid is smaller, and the entocristid is more labially inflected in Floridopteryx. On the m3, the metaconid is located farther posteriorly in Floridopteryx and the protocristid is at a right angle to the long axis of the tooth, not slightly angled as in the m3 of Oligopteryx. There are also differenc- es between these two genera in the structure of the dentary, the most important of which is the lack of a p3 alveolus in Floridopteryx and the presence of this alveolus (and tooth) in Oligopteryx. No lower teeth are known of Karstopteryx excluding com- parisons with the lower dentition of Floridopteryx. Floridopteryx is readily distinguished from the two European Eocene emballonurid genera Ta- chypteron and Vespertiliavus. Both of these genera possess a p3, which is lacking in Floridopteryx. Compared to Tachypteron and Vespertiliavus, the parastylar region of M1 is strongly reduced in Flo- ridopteryx with a small parastyle and shorter pre- paracrista. In addition, the M1 of Floridopteryx differs from Vespertiliavus in the near absence of a hypocone and the smaller talon with a straight (not posteriorly expanded) posterior margin. The lower molars are difficult to compare in the two specimens of Tachypteron franzeni from the mid- dle Eocene Messel site in Germany, both of which have the upper and lower dentitions tightly oc- cluded (Storch et al., 2002). Differences that can be observed include the weaker labial cingula on the lower molars and the more reduced coronoid process of the dentary in Floridopteryx. Compared to two species of Vespertiliavus recently described from the early to middle Eocene of Tunisia in northern Africa (Ravel et al., 2016), the m1 and m2 of Floridopteryx have a much broader talonid compared to the trigonid, a taller and more verti- cal paraconid, more closely placed paraconid and metaconid, more posteriorly directed protocristid owing to the more anterior location of the metaco- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 195 nid, larger entoconid, and more labially inflected entocristid. The genus Pseudovespertiliavus, re- cently described from the early to middle Eocene of Algeria in northern Africa (Ravel et al., 2016), is similar to Vespertiliavus and differs from Flo- ridopteryx in the same characters that the Florida Miocene genus differs from Vespertiliavus. There are two described species of the ex- tinct emballonurid genus Dhofarella, one from the late Eocene of Egypt and the second from the ear- ly Oligocene of Oman (Sigé et al., 1994; Gunnell et al, 2008). Only D. thaleri from Oman has the M1 preserved. Compared to D. thaleri, the M1 of Floridopteryx has a smaller parastyle and reduced parastylar region, shorter labially oriented pre- paracrista, shallower notch in the parafossa labial to the paracone, and a straight (rather than rounded) posterior margin of the talon basin. Neither species of Dhofarella preserves the dentary anterior to the m1, so the presence or absence of p3 in this genus cannot be determined. The m1 and m2 of Floridop- teryx differ from those of Dhofarella in the broader talonid, more closely placed paraconid and meta- conid, taller and more vertical paraconid, strongly angled protocristid from the more anteriorly placed metaconid to the protoconid, and the sharp labial inflection of the entocristid. Compared with the recently described middle Miocene emballonurine Afrillonura na- mibensis ffrom the Berg Aukas 1 site in the Otavi Mountain karst deposits of Namibia (Rosina and Pickford, 2021), the M1 of Floridopteryx differs in having a smaller parastyle, longer preparacrista, weaker paraloph and metaloph, poorly developed hypocone, and lack of a notch in the postprotocrista separating the protocone and hypocone. Compared to the m1 of Afrillonura, the m1 of Floridopteyx has the trigonid more compressed and shorter an- teroposteriorly with the paraconid and metaconid closer together, and the postcristid more strongly angled posterolabially from the more anteriorly lo- cated metaconid to the protoconid. The two genera in the Old World emballon- urid subfamily Taphozoinae, Taphozous and Sac- colaimus, are very similar dentally, and have in the past been considered congeneric. Our comparisons of Floridopteryx apply to both Taphozous and Sac- colaimus. The M1 of Floridopteryx differs from the M1 of the two taphozoines in its smaller size, overall shape of the tooth (more anteroposteriorly compressed in Floridopteryx, squarish in taphozo- ines), reduced parastylar area, much smaller para- style, presence of a paracingulum, more closely placed paracone and metacone, and a squarish tal- on basin. Compared to taphozoines, the lower m1 and m2 of Floridopteryx have an anteroposteriorly compressed trigonid, much broader talonid, more posterior position of the paraconid located close to the metaconid, anterolingually angled protocristid from the more posterior protoconid to the more an- teriorly placed metaconid and V-shaped entocristid oriented labially. Floridopteryx differs in several dental fea- tures from the Old World emballonurine genera Emballonura, Mosia, and Paremballonura, all of which were at one time included in Emballonura. The M1 of Floridopteryx is narrower anteroposte- riorly and wider in the transverse or labiolingual dimension (squarish in Emballonura except for talon), the ectoloph is not as shallow, the paracone and metacone are located farther apart and both in a more lingual position, the trigon and talon basins are not deeply pocketed, and the hypocone is very weak and is not separated from the protocone by a prominent notch in the postprotocrista. In several characters, the lower molars of Emballonura, Mo- sia, and Paremballonura are similar to taphozoines and differ from Floridopteryx. Compared to these three genera, the lower m1 and m2 of Floridopter- yx have a compressed trigonid with the paraconid and metaconid close to one another, the metaconid is anterior to the protoconid the talonid is broad- er, the talonid basin lacks a deep concave pit, the cristid obliqua meets the trigonid at a more acute angle, the entocristid is strongly curved/angled and is oriented labially, and the hypoconulid is more prominent. Dentally, the living African emballonurine genus Coleura is more similar to Floridopteryx and several genera of New World emballonurines than are Emballonura, Mosia, and Paremballonura. The M1 of Floridopteryx is similar in size to that 196 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) of Coleura but the parastyle is better developed and extends well anterior to the paracingulum, a short preparacrista is present, a paraloph is absent and the metaloph is weak, a deep concavity is lack- ing in the trigon basin, and the hypocone is highly reduced. Compared to Coleura, the m1 and m2 of Floridopteryx have the trigonid more compressed, the metaconid is more anterior, the protocristid is oriented anterolingually from the protoconid to the metaconid, the cristid obliqua meets the trigonid at a more acute angle and connects to the trigonid in a more lingual position, and the entocristid is more labially inflected. Our comparisons indicate that the Florida early Miocene genus Floridopteryx is the oldest member of the endemic New World emballonurid tribe Diclidurini (See discussion above of the sys- tematic relationships of Oligopteryx regarding the possibility this genus may also be an early diclidu- rine). We compared fossils of Floridopteryx to at least one species in each of the eight extant genera of New World diclidurines, that have been further subdivided into two subtribes (Lim, 2007; Lim et al., 2008): the Diclidurina (Balantiopteryx, Cor- mura, Cyttarops, Diclidurus, and Peropteryx) and the Saccopterygina (Centronycteris, Rhynchonyc- teris, and Saccopteryx). The comparisons are in al- phabetical order within each of the two diclidurine subtribes, beginning with the Diclidurina. Compared to Balantiopteryx, the M1 of Floridopteryx is larger and also differs in other features including the overall shape of the tooth, which is broader in the transverse dimension caus- ing the paracone and protocone to be more widely separated, more lingual location of the parastyle, longer preparacrista, better developed mesostyle, shallower concavity in the trigon basin, and very weak hypocone. The m1/m2 of Floridopteryx and Balantiopteryx are similar in the strong anteropos- terior compression of both the trigonid and talonid and by the much broader talonids compared to the trigonids, but the lower molars of Floridopteryx differ in the separation of the paraconid and meta- conid by a deeper notch, larger and more bulbous entoconid located in a more anterior position closer to the metaconid, and the location of the hypoconu- lid directly posterior to the entoconid. The M1 of Floridopteryx and Cormura are more similar than the Florida fossil is to most other extant genera of emballonurids, but there are some differences. The M1 of Floridopteryx is somewhat broader transversely, the parastyle is smaller and located more lingually, the protocone is in a more lingual position and more widely separated from the paracone, and the mesostyle is larger and more bulbous. Compared to Cormura, the m1 of Flori- dopteryx has the trigonid more compressed with the paraconid located more posteriorly and the metaconid more anteriorly, the paraconid is taller and on the lingual margin, the protocristid angles anterolingually from the protoconid to the meta- conid, and the cristid obliqua forms a more acute angle where it meets the trigonid. The talonid of m3 is broader in Floridopteryx. The M1 of Floridopteryx differs from that of the diclidurine Cyttarops in the lower, more round- ed cristae, larger parastyle, more prominent pre- paracrista, narrow paracingulum, more posterior position of the protocone, the near absence of a hy- pocone, and the less prominent concavities in the trigon and talon basins. The m1/m2 of Floridop- teryx differ from the lower molars of Cyttarops in the lower and more bulbous cusps, more posterior location and vertical orientation of the paraconid, closely placed paraconid and metaconid, anterolin- gual orientation of protocristid from protoconid to metaconid, much broader talonid, and larger hypo- conulid. The M1 of Floridopteryx differs from Di- clidurus in the presence of a larger parastyle lo- cated somewhat lingual to the anterolabial margin, broader paracingulum, more lingual position of the paracone, better developed mesostyle located more anteriorly, lack of deep concavities in the trigon and talon basins, and the near absence of a hypocone. The m1/m2 of Floridopteryx differs from Diclidu- rus in the much broader talonid compared to trigo- nid, strong anteroposterior compression of both the trigonid and talonid, closely placed paraconid and metaconid, larger paraconid located more posteri- orly, more anterior location of the metaconid, en- tocristid more sharply angled labially, and a larger MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 197 hypoconulid. Compared to Peropteryx, the M1 of Flori- dopteryx is broader transversely with the protocone positioned more lingually, lacks a deep concave pit in the trigon basin, and the hypocone is very weak. The m1/m2 of Floridopteryx differ from these low- er molars of Peropteryx in having a broader talonid compared to the trigonid, paraconid vertical, meta- conid more anterior and closer to the paraconid, protocristid strongly angled, cristid obliqua form- ing a more acute angle and meeting the trigonid farther lingually, and entocristid not as sharply V- shaped and less labially inflected. The M1 of Floridopteryx differs from Cent- ronycteris in its much larger size, lack of a deep V-shaped notch in the parafossa labial to the para- cone, more lingual location of the paracone and protocone, lack of deep pits in the trigon and talon basins, and the near absence of a hypocone. The m1/m2 of Floridopteryx and Centronycteris are similar in size but Floridopteryx has the paraconid located more posteriorly, the protocristid angled anteriorly from the protoconid to the metaconid, the cristid obliqua forming a more acute angle con- necting to the trigonid about halfway between the protoconid and metaconid, lacking a deep pit in the talonid basin, less sharply V-shaped entocristid that does not extend as far labially, and larger hypoco- nulid. Rhynchonycteris is the smallest New World emballonurid and differs from Floridopteryx in many dental features besides its much smaller size. Compared to Rhynchonycteris, the M1 of Flori- dopteryx has all cristae on the ectoloph somewhat lower and thicker and not sharp and blade-like, weaker parastyle, paracone and metacone located farther from the labial margin, lack of deep con- cave pits in the trigon and talon basins, and a very weak hypocone. The m1/m2 of Floridopteryx are shorter in the anteroposterior dimension, the proto- conid is in a more posterior position, the paraconid and metaconid are not as close together, the meta- conid is smaller, the lingual margin of the talonid is not as deeply concave because the entocristid is not a sharply deflected labially, and the entoconid is not laterally compressed. Among extant New World diclidurines, Flo- ridopteryx poyeri is most similar in size and dental morphology to Saccopteryx, specifically the spe- cies S. bilineata. However, there are notable dif- ferences between Floridopteryx and Saccopteryx. Compared to Saccopteryx, the M1 of Floridopteryx has a somewhat smaller parastyle that does not ex- tend as far anteriorly, the protocone is more lingual and farther from the paracone, and the trigon basin lacks a deep elliptically shaped pit or concavity. In the lower molars (m1/m2) of Floridopteryx, the talonid is relatively broader, the talonid basin lacks a deep pit, the paraconid is more vertical, the meta- conid is more anterior, the paraconid and metaco- nid are closer to one another, the protocristid is oriented at an angle to the long axis of the tooth, the cristid obliqua forms a more acute angle and attaches to the trigonid farther lingually, the ento- conid is larger and more inflated, the entocristid not as sharply V-shaped and is less labially inflected, and the hypoconulid is larger. The proximal end of the femur of Floridop- teryx differs from the femur of both Oligopteryx and the extant emballonurid Saccopteryx. The greater and lesser trochanters of Floridopteryx and Saccopteryx are similar in size and morphol- ogy, with the lesser trochanter more elongated and extending farther proximally than the greater tro- chanter. In Oligopteryx, the two trochanters extend proximally to the same level and the lesser tro- chanter is shorter, more robust, and triangular in shape. The femoral head is comparatively larger in Oligopteryx, approximately equal in width to the width of the shaft. The head is narrower than the shaft in Floridopteryx. The femoral head of Sac- copteryx is larger and is not canted medially com- pared to Floridopteryx. DISCUSSIONDISCUSSION sysTemaTIc relaTIonshIps Previous analyses of the systematic relation- ships within the family Emballonuridae include Barghoorn (1977) based on cranial and dental characters, Robbins and Sarich (1988) from pro- tein electrophoresis and immunology, Griffiths 198 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) and Smith (1991) on hyoid morphology, and more recent studies on molecular genetics (Lim, 2007; Lim et al., 2008; Goodman et al., 2012; Ruedi et al., 2012; Uvizl et al., 2019). Studies of both mor- phological and molecular characters demonstrate two major dichotomies among living species in the Emballonuridae. The first separation is between the strictly Old World subfamily Taphozoinae, in- cluding the genera Taphozous and Saccolaimus, and the subfamily Emballonurinae, including four genera from the Old World and eight Neotropical genera. The second major dichotomy occurs within the Emballonurinae, between the Old World tribe Emballonurini (Emballonura, Coleura, Mosia, and Paremballonura) and the New World tribe Dicli- durini. Finally, the Diclidurini have been separated into two subtribes (Lim, 2007; Lim et al., 2008), the Diclidurina (Balantiopteryx, Cormura, Cyt- tarops, Diclidurus, and Peropteryx) and the Sac- copterygina (Centronycteris, Rhynchonycteris, and Saccopteryx). The diclidurines are a monophyletic group that had a common origin, presumably from either Africa or North America via Eurasia as dis- cussed below. Not surprisingly, the earliest fossil represen- tatives of the Emballonuridae are also the most primitive. Four extinct genera of emballonurids have been described from Eocene and Oligocene faunas in the Old World: Vespertiliavus from the early to middle Eocene of Tunisia (Ravel et al., 2016) and the middle Eocene to Oligocene of Eu- rope (Barghoorn, 1977; Sigé, 1990; Smith et al., 2012; Maitre, 2014); Pseudovespertiliavus from the early to middle Eocene of Algeria (Ravel et al., 2016); Tachypteron from the middle Eocene of Germany (Storch et al., 2002; Smith et al., 2012); and Dhofarella from the late Eocene of Egypt (Gunnell et al., 2008) and the early Oligocene of Oman on the Arabian peninsula (Sigé et al., 1994). It is difficult to evaluate certain characters in two of these genera, Dhofarella and Pseudovespertilia- vus, represented almost entirely by isolated teeth, for example, the presence or absence of p3. Both Tachypteron and Vespertiliavus have a well-devel- oped, double-rooted p3, the primitive condition for the Emballonuridae. The only other emballonurid known to possess a p3 is the Oligocene Oligopteryx from Florida, in which this tooth is very small and single-rooted. All living members of the Emballon- uridae, including both Taphozoinae and Emballon- urinae, as well as the early Miocene Floridopteryx from Florida, lack a p3. All four Eocene/Oligocene genera of Old World emballonurids are represented by the M1, the most diagnostic tooth in the Em- ballonuridae. Three of these genera, Tachypteron, Vespertiliavus, and Pseudovespertiliavus, have a well-developed parastylar region on the antero- labial margin of the M1, including a prominent parastyle, long preparacrista, strong paracingulum (= precingulum), and wide shelf labial to the para- cone (= parafossa of Ravel et al., 2016). Compared to these three Eocene genera, as well as the living taphozoines Taphozous and Saccolaimus, Dho- farella thaleri from the early Oligocene of Oman (Sigé et al., 1994) has the parstylar area on the M1 more reduced, with a shorter preparacrista and re- duced shelf labial to the paracone. However, when compared to most living emballonurines, as well as Oligopteryx, Karstopteryx, and Floridopteryx from Florida, the parastylar region in Dhofarella is better developed, with a longer preparacrista, wider shelf labial to the paracone (parafossa) with a noticeable V-shaped notch, larger parastyle, and broader paracingulum. Rosina and Pickford (2021) considered Dhofarella to be the oldest member of the Emballonurinae. They also regarded their re- cently described extinct genus Afrillonura from the middle Miocene of Namibia in southwestern Africa to be an emballonurine, similar in dental characters to the extant African emballonurine genus Coleura (Rosina and Pickford, 2021). Ravel et al. (2016) presented a phylogenetic analysis of 65 dental and mandibular characters of four extinct Old World genera of Emballonuridae, including one species each of Dhofarella, Pseu- dovespertiliavus, and Tachypteron, and five species of Vespertiliavus, as well as two living species of Old World emballonurids, Emballonura (= Parem- ballonura) atrata and Taphozous melanopogon. No New World emballonurids (Diclidurini) were included in their analysis. Their phylogeny placed the three oldest genera at the base of the tree, all MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 199 known from either the early or middle Eocene, with Tachypteron in the basalmost position, fol- lowed next by Pseudovespertiliavus, and then the five species of Vespertiliavus. Ravel et al. (2016) placed their new genus and species, Chambinycte- ris pusilli, from the early to middle Eocene Chambi fauna in Tunisia, at the very base of the emballon- urid clade, although they considered Chambinycte- ris to be “family indeterminate” in their taxonomic analysis. In the Ravel et al. (2016) phylogeny, the extinct genus Dhofarella from the late Eocene and early Oligocene was placed closer to the modern genera Taphozous and Emballonura (= Parembal- lonura) than to Tachypteron, Pseudovespertiliavus, and Vespertiliavus. It is difficult to assess the re- lationship of Dhofarella to living emballonurids based on the Ravel et al. (2016) phylogeny because they included only two modern taxa, one taphozo- ine and one Old World emballonurine. Gunnell et al. (2008) noted that Dhofarella sigei from the late Eocene of Egypt was most similar to the African emballonurine genus Coleura among living embal- lonurids. As mentioned above, Rosina and Pickford (2021) also considered the extinct middle Miocene genus Afrillonura to be closely related to Coleura and placed both Afrillonura and Dhofarella in the Emballonurinae. Considering that all currently known Eocene emballonurids are from either Europe or northern Africa, it seems likely that the Emballonuridae had an Old World origin. New World emballon- urids were derived from Old World emballonurids sometime prior to the oldest Western Hemisphere occurrence of this family in the early Oligocene (~30 Ma) of Florida. With the possible exception of the incompletely known late Eocene and early Oligocene Dhofarella, the Oligocene fossils of Oligopteryx floridanus and O. hamaxitos from I-75 and Brooksville 2 in Florida are among the oldest records of the Emballonurinae and may also be the earliest members of the Diclidurini. However, based on the presence of a p3, Oligopteryx appears to be more primitive than the remainder of the New World emballonurid assemblage. The latest Oligo- cene Karstopteryx is closely related to Oligopteryx and may also be an early diclidurine. We hesitate to name a new, higher-level taxon based on fragmen- tary fossil material, but the Florida Oligocene em- ballonurids may represent an extinct subtribe of the Diclidurini. The Thomas Farm LF contains a third extinct genus of New World Emballonuridae, Flo- ridopteryx. In contrast to Oligopteryx, Floridop- teryx lacks the p3, and the M1 has several features that are more similar to the living or crown group of Neotropical emballonurids (Diclidurini). Owing to the incomplete and fragmentary condition of the fossil sample of Floridopteryx, we are not able to determine to which of the two diclidurine subtribes (Diclidurina or Saccopterygina) this extinct genus belongs. We did not conduct a phylogenetic analysis of Oligopteryx, Karstopteryx, and Floridopteryx, New World diclidurines, and living and extinct Old World emballonurids, although we did make morphological comparisons of the extinct Florida genera to all living and extinct genera of emballon- urids (See Comparisons sections in the taxonomic accounts of Oligopteryx floridanus and Floridop- teryx poyeri). A phylogenetic analysis in Ravel et al. (2016) included most extinct species of embal- lonurids, with the exception of the recently de- scribed Afrillonura namibensis (Rosina and Pick- ford, 2021), together with several archaic genera of Eocene bats belonging to extinct families (e.g., Ic- aronycteris, Archaeonycteris, Palaeochiropteryx), several enigmatic Eocene genera (Chambinycteris, Chibanycteris, Khoufechia), as well two living Old World species of emballonurids (Taphozous melanopogon and Emballonura = Paremballonura atrata) and two living species in the Nycteridae (Nycteris gambiensis, N. grandis). Their phyloge- netic analysis was based on dental and mandibular characters in the groups listed above. We produced a table (Table 4) that assesses dental and mandibular characters in all eight extinct genera of emballonurids, including the five extinct genera from the Old World (Afrillonura, Dhofarel- la, Pseudovespertiliavus, Tachypteron, and Vesper- tiliavus) and the three new extinct genera from the Oligocene and Miocene of Florida described here- in (Floridopteryx, Karstopteryx, and Oligopteryx), as well as species representing six living genera 200 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) of emballonurids, including a taphozoine (Tapho- zous), an emballonurine (Coleura), and four dicli- durines (Balantiopteryx, Diclidurus, Peropteryx, and Saccopteryx). Characters 1–65 in Table 4 were originally proposed by Ravel et al. (2016) in their phylogenetic analysis of the Emballonuridae, as well as several outgroups mentioned above. In our evaluation of characters from Ravel et al. (2016) that are present in the Florida fossil emballonurids, we noticed that many of these characters are use- ful in separating fossil and modern emballonurids from archaic Eocene bats and nycterids but are not as helpful in elucidating relationships within the Emballonuridae. Therefore, we added 20 dental characters to Table 4 (Characters numbered 66–85, under New Characters–this study) that our com- parisons indicated were more informative in sepa- rating genera within the Emballonuridae and are particularly important in evaluating the relation- ships of the Florida fossils. Our Table 4 is limited to Emballonuridae and includes 14 species, nine of which were not analyzed by Ravel et al. (2016): the newly described Miocene species from Namibia Afrillonura namibensis; the extant African embal- lonurine species Coleura afra; three extinct species of Florida emballonurids, Oligopteryx floridanus, Karstopteryx gunnelli, and Floridopteryx poyeri; and four living species of New World diclidurine emballonurids, Balantiopteryx plicata, Diclidurus albus, Peropteryx macrotis, and Saccopteryx bilin- eata. Because the six extant emballonurid genera in Table 4 are each represented by more than one species, we examined at least two species within a genus to make certain the species we analyzed were representative of the genera. Appendix 2 pro- vides a list of the 85 dental and mandibular char- acters evaluated in Table 4, providing detailed de- scriptions of both the charcters and character states within each character. We feel that the limited number of characters available in the Florida Oligocene and Miocene em- ballonurid fossils would be insufficient to generate a robust phylogenetic analysis of the New World Emballonurinae (Diclidurini) or of the Emballon- uridae in general. Recent work has emphasized the desirability of using an approach toward deriving phylogenetic hypotheses using multiple integrated lines of evidence and congruent datasets (Solari et al. 2019) that are not yet available in the fossil emballonurids and their extant relatives. Therefore, we elect to provide a suite of potentially phyloge- netically informative dental and mandibular char- acters that are available in the Florida emballonurid fossils (Table 4, Appendix 2) that could be added to larger databases of morphological characters for extant taxa (for example, from various systems of the soft anatomy). These characters could be incor- porated with genomic and other types of data to perform even more comprehensive and robust phy- logenetic analyses. Taphonomy and paleoecology All fossil specimens of Florida Oligocene and early Miocene Emballonuridae are derived from paleokarst deposits. Emballonurids are unknown in Florida mid Tertiary bat sites from non-karst depo- sitional environments, including the late Oligocene White Springs LF and several early Miocene sites (Morgan and Czaplewski, 2012). The two most productive fossil sites for emballonurids in Flori- da, the early Oligocene I-75 LF and late Oligocene Brooksville 2 LF, consist of terrestrially derived clay and sand deposits filling small pockets or fis- sures in Paleogene marine limestones that represent remnants of former caves (Patton, 1969a; Hayes, 2000; Morgan and Hulbert, 2008). The taphonomy of these two sites is consistent with cave deposits, containing substantial samples of small mammals, including abundant bats, as well as other small ver- tebrates (mostly amphibians and reptiles, birds are rare or absent), and isolated teeth and small post- cranial elements of larger mammals (Hayes, 2000; Holman and Harrison, 2001; Morgan and Hulbert, 2008). The large emballonurid Oligopteryx florida- nus is the most common bat in the Brooksville 2 fauna, with a sample consisting of an NISP (num- ber of identifiable specimens) of more than 100, representing seven individuals, comprising about half of the chiropteran fossils from this site. The second most abundant bat in the Brooksville 2 LF is the mormoopid Koopmanycteris palaeomormoops, represented by about 50 specimens (Morgan et al., MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 201 2019). These same two species, the emballonurid O. floridanus and the mormoopid K. palaeomor- moops, are also the most common bats in the I-75 fauna. A smaller emballonurid, O. hamaxitos, is also present in Brooksville 2 and I-75 but is rather uncommon with two individuals in each fauna. The samples of mormoopid fossils in the I-75 and Brooksville 2 karst deposits are consistent with our proposed taphonomic scenario for these two sites, since almost all species in the Mormoopidae roost in caves (Smith, 1972). A partial radius represent- ing an indeterminate genus in the Natalidae is also known from I-75, and all living natalids are obli- gate cavernicoles (Morgan and Czaplewski, 2003; Tejedor, 2011). The abundance of Oligopteryx in the two Florida Oligocene karst sites suggests these emballonurids were also cave dwellers. In the context of elucidating the evolutionary history of roosting ecology in the Emballonuridae, it is instructive to examine the roosting habits of living members of this family. Most extant New World species of emballonurids roost in trees, in- cluding recesses between buttresses on the outside of trees, underneath fallen trees, within hollow trees, and under leaves (Nowak, 1994; Simmons and Voss, 1998). Several species of Neotropical emballonurids are known to roost in caves, in- cluding Balantiopteryx io, B. plicata, Peropteryx macrotis, and Saccopteryx bilineata (Goodwin and Greenhall, 1961; Nowak, 1994; Simmons and Voss, 1998; Ceballos et al., 2014). Goodwin and Greenhall (1961, p. 216) noted tbat Peropteryx macrotis “…has been found on Tobago roosting in association with Glossophaga longirostris in well- lighted limestone and coral caves that overlook the ocean.” Old World species of emballonurids more commonly roost in caves than do New World mem- bers of this family (Hill and Smith, 1984; Nowak, 1994). Species in the taphozoine genus Taphozous are called tomb bats for their tendency to roost in tombs, caves, or rock crevices, and species in the emballonurine genera Emballonura and Coleura also often roost in caves (Nowak, 1994; Bambini et al. 2006; Nkrumah et al., 2021). Among extinct Old World genera of Embal- lonuridae, most species in the genus Vespertiliavus from the Eocene and Oligocene in France and else- where in western Europe are from karst deposits that probably represent former caves (Sigé and Legendre, 1983; Maitre, 2014). Fossils of the re- cently described extinct emballonurine genus Af- rillonura and associated taphozoines in the gen- era Taphozous and Saccolaimus, from Namibia in southwestern Africa, are from Miocene cave- derived karst deposits (Rosina and Pickford, 2020, 2021). King et al. (2020) reported Taphozous and Saccolaimus from an early Pleistocene site in the Riversleigh World Heritage Area in Australia, con- sisting of an extensive series of karst deposits. The evidence from the Brooksville 2 and I-75 sites, including the geology, taphonomy, and vertebrate fauna, in particular the abundance of bats, strongly indicates that both species of Oligop- teryx were cavernicolous. This suggests a different paleoecology for the Florida Oligocene emballon- urids compared to the majority of modern Neotrop- ical species in this family, as noted above. The early Oligocene I-75 LF and late Oligocene Brooksville 2 LF are the oldest karst deposits in North America known to contain bats (Czaplewski et al., 2008; Morgan and Czaplewski, 2012; this paper). Conse- quently, the two species of emballonurids (Oligop- teryx floridanus and O. hamaxitos) and one species of mormoopid (Koopmanycteris palaeomormoops) described from these two sites, as well as a nata- lid from I-75 (Morgan and Czaplewski, 2003), are the oldest known cave-dwelling bats in the West- ern Hemisphere, taking into consideration the lack of pre-Pleistocene karst deposits in South America containing fossil bats. The early Miocene Thomas Farm site is also a paleokarst deposit, but the taphonomy and verte- brate fauna are quite different from the Brooksville 2 and I-75 sites. Thomas Farm is a large, deep (>30 m), sediment-filled sinkhole that accumulated sig- nificant samples of both large mammals and small vertebrates, including abundant bats (Pratt, 1989, 1990). The proposed taphonomic setting for the large sample of bats in the Thomas Farm LF is the presence of vertical limestone walls in the sinkhole containing a cave or caves that supported colonies of cave-dwelling bats (Pratt, 1989). Although bat 202 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) fossils occur throughout the stratigraphic section in the Thomas Farm site, they are concentrated in certain layers, in particular a lime sand near the top of the section. This lime sand contains large sam- ples of bats and other small vertebrates (rodents, amphibians, lizards, snakes, and birds) but a rather limited sample of larger mammals, suggesting this layer may represent a former cave deposit. The majority of bat fossils from Thomas Farm (>75%, consisting of several thousand specimens) belong to Suaptenos whitei, an extinct genus and species in the family Vespertilionidae that was almost cer- tainly a colonial cavernicolous bat. The second most common bat at Thomas Farm, Primonatalus prattae, is one of the earliest known members of the Natalidae, a family composed entirely of cave- dwelling species (Morgan and Czaplewski, 2003, 2012; Tejedor, 2011). Emballonuridae are rare at Thomas Farm, with only six specimens of Flori- dopteryx poyeri from a sample of more than 3,000 bat fossils (<1% of the total chiropteran fauna from this site). The rarity of Floridopteryx at Thomas Farm suggests this bat may not have been a cave dweller. This would support our hypothesis that Floridopteryx is a member of the Diclidurini, the extant tribe of Neotropical emballonurids, most species of which do not roost in caves as discussed above. The fossil record of emballonurids in Flor- ida suggests an ecological shift in the roosting preferences of New World emballonurids through time, from cavernicolous species in the Oligocene to tree-roosting (non-cave dwelling) species in the Miocene and continuing to the present with the predominance of tree-roosting species in the mod- ern Neotropical fauna. The late Oligocene and Miocene emballon- urid fossils from South America, including samples from La Venta, Colombia and Contamana, Peru, were preserved in sediments derived from fluvi- al, lacustrine, or tropical forest environments, not karst deposits (Czaplewski, 1997, 2005; Antoine et al., 2016). Only one of the South American Tertiary emballonurids has been identified below the family level, the extant genus Diclidurus from the middle Miocene La Venta Fauna (Czaplewski, 1997). The living species Diclidurus albus roosts in tropical vegetation, especially where palm trees are abun- dant (Ceballos and Medellín, 1988). In this species, which has white fur, sunlight filtered through the palm leaves on white fur gives the fur a greenish cast and provides a form of camouflage in the roost or in flight (as it does for the white phyllostomid bat, Ectophylla alba and other species; Brooke, 1990; Rodríguez-Herrera et al., 2007; Rydell et al., 2019). The occurrence of four species of Embal- lonuridae in the Oligocene and early Miocene of Florida also has paleoclimatic implications. The Emballonuridae is a tropical family of bats. The present distribution of emballonurid species in the New World closely tracks the occurrence of tropi- cal forests, which also defines the northern limits of the Neotropical Region (Olson et al., 2001). Sev- eral species of living emballonurids, in particular Balantiopteryx plicata, occur at the northern edge of the Neotropics in Mexico, as far north as the Tropic of Cancer (~23° North) in the lowlands of eastern Mexico and somewhat farther north (~27° N) along the Pacific coast of western Mexico (Sim- mons, 2005). The approximate latitudes of the Florida Oligocene and early Miocene emballon- urid records are somewhat farther north than the northernmost current Mexican records of species in this family: Brooksville 2 (28° N), I-75 (29° N), and Thomas Farm and Buda (30° N). No emballon- urids are known from the modern fauna of Florida, or from Florida fossil sites younger than the early Miocene Thomas Farm LF. The disappearance of emballonurids from Florida after the early Miocene reflects the overall trend in Florida Tertiary chiropteran faunas. Oli- gocene bat faunas (~24–30 Ma) are dominated by families now primarily tropical in distribution, in particular the Emballonuridae and Mormoopidae, but also including the Natalidae and Molossidae (Czaplewski et al., 2003a; Morgan and Czaplewski, 2003, 2012). The Vespertilionidae, widely distrib- uted in temperate regions at present, is represented by a single tooth in the early Oligocene I-75 LF and is absent from the late Oligocene Brooksville 2 LF (Morgan and Czaplewski, 2012). Two inde- terminate species of vespertilionids were reported MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 203 from the late Oligocene White Springs LF in north- ern peninsular Florida (Morgan and Czaplewski, 2012). White Springs is a non-karst deposit con- sisting of shallow marine and fluvial sediments that, in addition to marine vertebrates (sharks, rays, bony fish, sirenians), also contains a rather diverse land mammal fauna, including two species of bats, seven species of rodents, several partial articulated skeletons of the oreodont Mesoreodon floridensis, and other mammals indicative of an early late Ari- kareean age (Ar3; MacFadden and Morgan, 2003). A latest Oligocene age for White Springs is sup- ported by a strontium isotope age estimate of 24.4 Ma on marine mollusk shells from the same unit that produced the land mammal fauna (Jones et al., 1993). The early Miocene (~18 Ma) Thomas Farm LF also contains three families of tropical bats, Em- ballonuridae, Natalidae, and Molossidae, but the overall fauna is dominated (>90% of fossil sample) by the more temperate Vespertilionidae, including at least five species (Lawrence, 1943; Czaplewski and Morgan, 2000; Czaplewski et al., 2008; Mor- gan and Czaplewski, 2012). The chiropteran record in Florida between the early Miocene and early Pleistocene (~2-18 Ma) is sparse; only a few fossils are known almost all of which are vespertilionids (Morgan and Hulbert, 2008; Morgan and Czaplews- ki, 2012). Changes in the distributional patterns of Florida bats during the mid Cenozoic almost cer- tainly reflect overall climatic changes in the Florida peninsula and southeastern North America, from a tropical or subtropical climate in the Oligocene and early Miocene to a warm temperate climate from the middle Miocene to the present. A diverse middle Miocene paleoflora from the Florida Panhandle provides information on the vegetation and climate of Florida during the time period shortly after the early Miocene Thomas Farm LF. Jarzen et al. (2010) reviewed the palynol- ogy and Lott et al. (2019) reviewed the macroplant remains from the Alum Bluff flora in the Fort Pres- ton Formation along the Apalachicola River in the central Florida Panhandle (Latitude: 30°28’N). The Alum Bluff flora has been dated as middle Miocene (early Barstovian NALMA; ~16 Ma) based on an associated land mammal fauna consisting of four taxa of ungulates, as well as molluscan biostra- tigraphy and strontium isotope chronology of un- derlying and overlying marine units (Bryant et al., 1992). The two paleobotanical studies documented that the affinities of many of the identified palyno- morphs and macroplants from the Alum Bluff flora indicate a warm temperate climate, much like that found in the northern Gulf Coast region of Florida today. The plant community consisted of an elm (Ulmus)–hickory (Carya)–palm forest (Sabalites- an extinct genus near the modern sabal or cabbage palm Sabal), occurring in close proximity to an oak and pine-dominated landscape. This interpretation of the Alum Bluff flora is much different from a previous study more than a century earlier that in- terpreted this same flora as being tropical to sub- tropical in its affinities (Berry, 1916). The Alum Bluff flora closely resembles the modern floras of the northern Gulf Coast through the north-central and northern Atlantic coast regions of peninsular Florida and extending northward along the Atlantic coasts of Georgia and South Carolina. The results of a climate analysis of the Alum Bluff flora, using leaf margin and leaf area (Lott et al., 2019), gave estimates of 19°C mean annual temperature and 116 cm mean annual precipitation, both of which are similar to modern values of mean annual tem- perature and precipitation for the Florida Panhan- dle and the north-central Florida peninsula. No Oligocene or early Miocene paleofloras are known from Florida. An attempt to recover pol- len from the early Miocene Thomas Farm site was unsuccessful (Pratt, 1989, 1990). The presence of tropical bats in Oligocene and early Miocene ver- tebrate faunas from peninsular Florida provides a proxy for the mid Cenozoic climate and vegetation in this region. The prevalence of tropical families of bats in Florida Oligocene faunas (~24–30 Ma) is indicative of a tropical to subtropical climate and vegetation during this time period. In the early Miocene Thomas Farm LF, the predominance of the more temperate family Vespertilionidae, together with the uncommon occurrence of three families of primarily tropical bats, suggests that the climate had moderated by the late early Miocene (~16–18 204 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Ma), with subtropical habitats and vegetation pre- dominating. Based on the modern fauna, embal- lonurids in the New World are a particularly sen- sitive indicator of tropical forest habitats. The di- clidurine emballonurid Floridopteryx would seem to indicate the presence of subtropical habitats, or perhaps even small pockets of tropical vegetation, in the vicinity of the Thomas Farm sinkhole in the early Miocene. With the transition to a more tem- perate climate in the middle Miocene, as indicated by the warm temperate Alum Bluff flora, tropical groups of bats disappeared from Florida presum- ably because of the disappearance of tropical and/ or subtropical vegetation and habitats. Similar mid Cenozoic changes are also ob- served in the European chiropteran fauna, tran- sitioning from a tropical or subtropical fauna in the Eocene, Oligocene, and early Miocene to a temperate fauna after the middle Miocene (Leg- endre, 1980; Sigé and Legendre, 1983; Horáček, 2001; Gunnell and Simmons, 2005). Two genera of emballonurids are known from the middle Eo- cene of Europe, Tachypteron and Vespertiliavus, and Vespertiliavus also occurs in late Eocene and early Oligocene European faunas. A record of the extant genus Taphozous from the early Miocene is the youngest emballonurid from Europe (Legen- dre, 1980; Gunnell and Simmons, 2005). Vesper- tilionids underwent an expansion and diversifica- tion in European faunas during the Miocene (Sigé and Legendre, 1983; Horáček, 2001; Gunnell and Simmons, 2005), much as they did in the Mio- cene of Florida and elsewhere in North America (Czaplewski et al., 2008; Morgan and Czaplewski, 2012), and presumably for the same reason, the cooling climate and expansion of temperate habi- tats in the Northern Hemisphere. Tropical species of bats are unknown from fossil deposits in the Florida peninsula after the early Miocene, except for a brief appearance of several species in the Pleistocene, including (Mor- gan et al., 1988; Morgan, 1991, 2002): two species of Phyllostomidae, the vampire bats Desmodus ar- chaeodaptes and D. stocki, both now extinct; two species of Mormoopidae, the extant Mormoops megalophylla and the extinct Pteronotus cf. pris- tinus; and a large living species of Molossidae, Eumops underwoodi. Among these Pleistocene bats with Neotropical affinities, three species are extinct and the extant species M. megalophylla and E. underwoodi are currently extralimital to Florida, occurring no closer than the southwestern U. S. and northern Mexico. Morgan and Emslie (2010) attrib- uted the occcurrence of tropical/subtropical bats in Florida Pleistocene faunas, as well as other species of mammals and birds with tropical affinities, to changes in climate, vegetation, and biogeographic patterns during glacial and interglacial periods, be- ginning at about 2.6 Ma and continuing to the end of the Pleistocene. The modern chiropteran fauna of Florida consists of 20 species, mainly composed of Vesper- tilionidae (13 species), together with seven species belonging to two other families with tropical affin- ities, the Molossidae and Phyllostomidae (Marks and Marks, 2006). Three species of molossids are recorded from the modern Florida fauna, the wide- spread Brazilian or Mexican free-tailed bat Tadari- da brasiliensis, the Florida bonneted bat Eumops floridanus from southern peninsular Florida, and the velvety free-tailed bat Molossus molossus from the Florida Keys (Frank, 1997b; Marks and Marks, 2006). Eumops floridanus is endemic to the south- ern half of the Florida peninsula (Koopman, 1971; Timm and Genoways, 2004; Vannatta et al., 2021), and also has been identified from three late Pleis- tocene fossil deposits in southern Florida (Morgan, 1991, 2002). In fact, E. floridanus was originally described as the extinct species Molossides florida- nus based on a mandible from the late Pleistocene Melbourne LF in Brevard County about midway along the Atlantic Coast of Florida (Allen, 1932), before it was discovered as a living animal in Miami at the southern tip of the peninsula (Barbour, 1936; Koopman, 1971). The Melbourne LF is slightly north of the currently known range of E. floridanus (Vannatta et al., 2021). The biogeographic origin of E. floridanus appears to be from the West Indies, as genomic data indicate this species is very similar to E. ferox from Cuba (Timm and Genoways, 2004; McDonough et al., 2008; Bartlett et al., 2013). Four species in the Phyllostomidae, the West Indian fruit MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 205 bat Artibeus jamaicensis, the Cuban fig-eating bat Phyllops falcatus, the buffy flower bat Erophylla sezekorni, and the Cuban flower bat Phyllonycte- ris poeyi, are West Indian bats known in Florida only by rare records from the Florida Keys (Frank, 1997a; Marks and Marks, 2006). The records of the four phyllostomids from the Florida Keys almost certainly represent accidental occurrences from Cuba, probably during hurricanes. BIogeography and evoluTIonary hIsTory oF The emBallonurIdae The Emballonuridae is a pantropical fam- ily of bats found in Mesoamerica, South America, Africa, Southeast Asia, Australia, and some Pacific Islands (Simmons, 2005; Simmons and Cirranello, 2019). There are 14 genera and over 50 species of extant Emballonuridae (Simmons, 2005; Goodman et al., 2012; Simmons and Cirranello, 2019): six genera in the Old World (Coleura, Emballonura, Mosia, Paremballonura, Saccolaimus, and Tapho- zous) and eight genera in the New World (Balan- tiopteryx, Centronycteris, Cormura, Cyttarops, Diclidurus, Peropteryx Rhynchonycteris, and Sac- copteryx). The geographic occurrence of the six genera of Old World emballonurids is as follows: Saccolaimus and Taphozous have similar distribu- tions, occurring in Africa, southerm Asia, and Aus- tralia; Coleura is restricted to Africa, Madagascar off the east coast of Africa, and the Seychelles in the Indian Ocean; Emballonura is found in south- east Asia and the Pacific Islands; Paremballonura is known from two species in Madagascar; and the single species of Mosia occurs in eastern Indone- sia, New Guinea, the Bismarck Archipelago, and the Solomon Islands. Most genera of New World emballonurids have a widespread distribution in the Neotropical region, occurring from southern Mexico or northern Central America south throughout Central America and the tropical portion of South America, with at least one species found as far south as Paraguay (Simmons, 2005; Lopez-Aguirre et al., 2019; Sim- mons and Cirranello, 2019). Their occurrence mir- rors the distribution of tropical forests. The north- ernmost New World emballonurid, Balantiopteryx plicata, occurs as far north as southern Sonora and southern Chihuahua in northwestern Mexico (~27° N; Alvarez-Castañeda and Patton, 1999) and eastern San Luis Potosí in eastern Mexico (~22° N; López-Forment and Tellez-Giron, 2014). Five other species of emballonurids, Balantiopteryx io, Diclidurus albus, Peropteryx macrotis, Rhyn- chonycteris naso, and Saccopteryx bilineata, oc- cur in the tropical lowlands of eastern Mexico as far north as Veracruz, with several of these species found somewhat farther north in the Mexican Pa- cific coastal states of Guerrero, Jalisco, and Nayarit (Medellín et al., 2008; Ceballos et al., 2014). Em- ballonurids are absent from the West Indies, ex- cept for several continental islands in the southern Caribbean Sea that would have been connected to northern South America during the late Pleistocene low sea level stand, including Trinidad with five species of emballonurids, Tobago with three spe- cies, Margarita with two, and Aruba with a single species (Simmons, 2005). Peropteryx trinitatis oc- curs on these islands and on Grenada, an oceanic island at the southernmost end of the Lesser An- tillean archipelago. All of these islands, including Grenada, have a South American chiropteran fauna lacking Antillean endemic species (Eshelman and Morgan, 1985; Koopman, 1989; Genoways et al., 1998). The previous published fossil record of the Emballonuridae in the Western Hemisphere is sparse, even if records from late Pleistocene cave deposits are included. This is probably a result of the roosting habits of New World emballonurids, most of which are not typically found in caves (see Taphonomy and Paleoecology discussion above). Besides the Oligocene and early Miocene records from Florida described here, all other Tertiary re- cords of New World Emballonuridae are from South America, consisting of seven isolated teeth from four different localities. Two associated upper molars referred to the living genus Diclidurus and a canine of a smaller indeterminate emballonurid were identified from the middle Miocene La Venta Fauna in Colombia (Czaplewski, 1997; Czaplews- ki et al., 2003b). Four isolated emballonurid teeth were reported from Oligocene and Miocene faunas 206 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) from Contamana, Peru (Antoine et al., 2016 and supplementary data): two lower teeth from a late Oligocene (Deseadan SALMA) fauna in the Cham- bira Formation; and two records from the Miocene Pebas Formation, a large upper molar from an early Miocene (Colhuehuapian/Santacrucian SALMAs) fauna and a lower canine from a late Miocene (Mayoan/Chasicoan SALMAs) fauna. The embal- lonurid teeth from the Contamana faunas were not described and only identified to the family level. The Quaternary record of the Neotropical Emballonuridae consists entirely of living spe- cies from cave deposits. Dalquest and Roth (1970) identified Balantiopteryx io from a late Pleistocene deposit in Cueva de Abra in southern Tamaulipas, northeastern Mexico (just south of the Tropic of Cancer at about 23° N). This cave is near the north- ern limit of the Neotropics but is somewhat north of the current northernmost occurrence of B. io in southern Veracruz (Hall, 1981; Alvarez-Castañeda and Patton, 1999; Medellín et al. 2008). Peropteryx macrotis was identified from Late Quaternary cave deposits in Loltún Cave in the Yucatán peninsula of Mexico (Arroyo-Cabrales, 1992; Arroyo-Cabrales and Polaco, 2003), within the modern range of the species. Eshelman and Morgan (1985) identified P. macrotis from Late Quaternary deposits in Robin- son Crusoe Cave on Tobago in the southeastern Ca- ribbean Sea. The extant Peropteryx from Tobago has since been referred to P. trinitatis (Simmons and Voss, 1998; Simmons, 2005), suggesting that the Quaternary fossils probably belong to P. trini- tatis as well. Emballonurids are absent from the ex- tensive fossil record of bats from Late Quaternary deposits in the West Indies, most of which are from caves (Morgan, 2001). The Quaternary record of emballonurids in South America consists of the ex- tant P. macrotis from cave deposits in the states of Bahia and Minas Gerais, Brazil (Czaplewski and Cartelle, 1998; Lessa et al., 2005). Two new species of Emballonuridae belong- ing to the new genus Oligopteryx are described here from the Oligocene of Florida, O. floridanus and O. hamaxitos from the Whitneyan I-75 LF and late early Arikareean Brooksville 2 LF, and a third new species, Karstopteryx gunnelli, is described from the latest Oligocene (early late Arikareean) Buda LF. These three Oligocene species represent the oldest New World records of the Emballonuri- dae, ranging in age from ~24-30 Ma. There are nu- merous earlier records of emballonurids from the Old World, including: early to middle Eocene of Algeria and Tunisia in northern Africa (Vespertil- iavus, Pseudovespertiliavus); middle to late Eo- cene of Europe, primarily France and Germany (Tachypteron, Vespertiliavus); and late Eocene of Egypt (Dhofarella). The Emballonuridae apparent- ly originated in the Old World (Europe or Africa) in the early to middle Eocene (Storch et al., 2002; Ravel et al., 2016), and dispersed to North America by the early Oligocene from either Europe or Asia (this paper), or from South America by way of Af- rica (Teeling et al., 2005; Lim 2007, 2008, 2009, 2010). Molecular, morphological, biogeographic, and behavioral studies of modern Eastern Hemi- sphere and Western Hemisphere Emballonuridae (Teeling et al., 2005; Lim 2007, 2008, 2010; Lim and Dunlop 2008; Lim et al. 2008; Ruedi et al. 2012) have suggested that the New World Embal- lonurinae (Diclidurini) had an African origin. Teel- ing et al. (2005) and Lim (2007, 2008) proposed that emballonurids arrived in South America from Africa by overwater dispersal in the Oligocene be- tween 25 and 31 Ma, at about the same time as the better-known dispersal of platyrrhine monkeys and caviomorph rodents from Africa to South America (e.g., Flynn and Wyss, 1998). More recent fossil discoveries in South America have extended the record of caviomorphs back to the early Oligo- cene (Arnal et al., 2019) and middle Eocene (An- toine et al., 2012; Assemat et al., 2019) and that of platyrrhines and parapithecid primates to the late Eocene and early Oligocene, respectively (Bond et al., 2015; Seiffert et al., 2020). Prior to 2010, the earliest published record of the Emballonuri- dae in the New World consisted of several teeth from the middle Miocene (~12 Ma) of Colombia (Czaplewski, 1997; Czaplewski et a1., 2003b). More recently, the New World record of emballon- urids has been extended back into the Oligocene on both continents, with both early and late Oligo- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 207 cene (~24-30 Ma) samples from Florida in south- eastern North America (Morgan and Czaplewski, 2012; this paper) and isolated teeth from the late Oligocene (~25 Ma) of Contamana, Peru in South America (Antoine et al., 2016). The molecular di- vergence dates for the basal split of the New World emballonurines of 32.5 Ma (Lim, 2007) and 30 Ma (Teeling et al., 2005) are close in age to the oldest fossils of this group from the early Oligocene of Florida and are older than the earliest emballonurid in South America from the late Oligocene of Peru. The emballonurid Dhofarella, from the late Eocene Fayum deposits in Egypt in northern Afri- ca, has been allied with emballonurines (Gunnell et al., 2008; Ravel et al., 2016). The M1 of Dhofarella is more similar to that of emballonurines than to other Eocene emballonurids or extant taphozoines, and Gunnell et al. (2008) noted that Dhofarella is similar to the extant African emballonurine ge- nus Coleura. Dhofarella is also known from an early Oligocene fauna from Oman on the Arabian Peninsula, which is a part of the African-Arabian continental tectonic plate, although colliding with Eurasia beginning in the late Eocene (Sigé et al., 1994). There are two other Tertiary occurences of emballonurines in Africa after the late Eocene and early Oligocene records of Dhofarella, includ- ing the recently described middle Miocene embal- lonurine Afrillonura namibensis from Namibia in southwestern Africa (Rosina and Pickford, 2021) and an extinct species in the extant emballonurine genus Coleura, C. muthokai, from the Pliocene of Ethiopia (Wesselman, 1984). There are also sev- eral records of taphozoine emballonurids from the Miocene and Pliocene of Africa (Gunnell and Sim- mons, 2005; Gunnell, 2010; Rosina and Pickford, 2019, 2020; Gunnell and Manthi, 2020). Our mor- phological analysis suggests that Oligopteryx from the Oligocene of Florida is more similar dentally to living New World emballonurines than is Dho- farella of equivalent age. South America and Africa are currently separated by a minimum distance of about 2,500 km, although taking into account seafloor spread- ing from the Eocene to the present, this distance would have been considerably less when monkeys and rodents (and possibly bats) dispersed westward across the Atlantic. Emballonurids are certainly capable of long-distance dispersals over water, as their colonization of many Indian Ocean and Pa- cific Ocean islands attests. The failure of emballon- urids to similarly colonize the West Indian islands is an enigma. An African origin for the New World Emballonuridae would require two oceanic disper- sal events for this group, first from Africa to South America and then from South America to North America, both of which must have occurred prior to the early Oligocene (~30 Ma) record of Oligop- teryx in Florida. Lim (2009) hypothesized that em- ballonurids reached Florida by overwater dispersal in the Oligocene from South America across the West Indies. The absence of emballonurids in both the modern and fossil record of the West Indies suggests the more likely overwater dispersal route from South America, if indeed the New World em- ballonurids reached South America first (see alter- nate hypothesis below), was northward across the Central American Seaway (CAS), and then over- land through Mesoamerica and around the western margin of the Gulf of Mexico to Florida. Lim (2010) further elaborated upon the colo- nization of South America by emballonurids with a model for the diversification of the monophyletic Neotropical diclidurines, proposing an early split of the two subtribes about 27 Ma with Saccopter- ygina as primarily forest-adapted and Diclidurina as primarily savanna-adapted. He suggested that after a period of stasis, a rapid diversification of genera began in the early Miocene, spurred by a forest-savanna mosaic resulting from a marine in- cursion into the western Amazon (Lim, 2010), with the appearance of the eight modern New World genera in the early to middle Miocene between 14 and 19 Ma. The extinct diclidurine genus Floridop- teryx is known from the early Miocene (~18 Ma) during the time interval when Lim (2010) proposed the modern genera of diclidurines evolved. The earliest record of an extant genus of diclidurine is Diclidurus, which first appeared at the end of this time interval in the middle Miocene (~12 Ma) of Colombia. Generic differentiation within the Dicli- durini is presumed to have occurred primarily in 208 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) South America (Lim, 2008), or a New World fa- milial differentiation in Mesoamerica and northern South America (López-Aguirre et al., 2019). The small sample of isolated teeth of emballonurids from the late Oligocene and Miocene of Contama- na, Peru and middle Miocene of La Venta, Colom- bia (Czaplewski, 1997; Czaplewski et al., 2003b; Antoine et al., 2016) are not complete enough to evaluate Lim’s hypothesis of emballonurid diver- sification, which was based on inferences from data on modern specimens. It is also possible with a North American (instead of African) orgin, that some of the modern genera of emballonurids had already appeared in tropical Mesoamerica in the Miocene prior to their dispersal to South America. Only a Tertiary fossil record of emballonurids from Mexico or Central America will answer this ques- tion. López-Aguirre et al. (2019) used evidence from geographic patterns of phylogenetic diver- sity and phylogenetic endemism of the extant gen- era and species of Emballonuridae, in comparison with those of other families of bats, to confirm a hypothesis that niche conservatism limited the spread of emballonurids beyond a proposed tropi- cal to intertropical center of origin. They also found an unusual, spatially discontinuous cluster- ing of some emballonurid clades in certain parts of Central America with others in distant parts of South America that might indicate “in situ diver- sification of vicariant species, phylogenetically distant from other co-occurring species” (López- Aguirre et al., 2019, p. 1196). The Florida fossils described here fall completely outside of the fam- ily’s modern Western Hemisphere distribution (as mapped by López-Aguirre et al., 2019), and thus they shed new light on the family’s evolutionary biogeographic radiation in southeasternmost North America in the late Oligocene-early Miocene. The mid Cenozoic occurrence of emballonurids in the Florida peninsula probably affected the family’s later patterns of diversification and endemism to an unknown degree, unless the Florida fossils were members of a clade that became extinct without giving rise to any still-living descendants. We propose an alternative hypothesis for the origin of the New World Emballonuridae based on the available fossil evidence, which includes the presence of three extinct genera and four extinct species in this family in Florida from the early Oli- gocene through the early Miocene, between about 30 and 18 million years ago. We suggest that Oli- gopteryx from the early Oligocene of Florida was derived from a basal Old World emballonurid, a bat similar to Vespertiliavus from the Eocene of Europe, that gave rise to both the Old World (Emballonuri- ni) and New World (Diclidurini) emballonurines. Dhofarella appears to be too different dentally to be closely related to Oligopteryx. An ancestral em- ballonurid dispersed to North America using either a western route from Europe across the Canadian Arctic, or possibly from eastern Eurasia across Beringia, sometime in the middle to late Eocene, giving rise to Oligopteryx from the early Oligocene of Florida. The western European route is perhaps more plausible, because primitive emballonurids are well known from the Eocene and Oligocene of Europe but are unknown from the Tertiary of Asia (Gunnell and Simmons, 2005). Moreover, a diverse fauna of early Eocene (Wasatchian NALMA) ver- tebrates, including many tropical species, is known from Ellesmere Island in the eastern Canadian High Arctic region near Greenland (McKenna, 1975; Dawson, 1976; 1991; Estes and Hutchison, 1980; Eberle and McKenna, 2002). Although the Elles- mere Island fauna is older than the earliest known emballonurid from Europe (middle Eocene), this fauna confirms that tropical vertebrates occurred in the eastern Arctic during the Eocene and that this region was on a dispersal route for Eocene verte- brates between Europe and North America. We hypothesize that sometime prior to the late Oligocene appearance of emballonurids in Amazonian Peru, an ancestral diclidurine similar to Oligopteryx (but having lost the p3) dispersed southward from Central America overwater across the Central American Seaway (CAS) to the then- island continent of South America. The CAS sepa- rating North America and South America may have been as narrow as 200 km in the early Miocene at about 20 Ma (Montes et al., 2012), which was far less than the distance across the Atlantic Ocean MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 209 separating South America from Africa at this same time. Floridopteryx from the early Miocene of Florida was derived from Oligopteryx or a similar genus such as Karstopteryx, and probably evolved in southeastern North America. Emballonurids disappeared from temperate North America (e.g., Florida) after the early Miocene but probably con- tinued to inhabit tropical Mesoamerica, although the family has no fossil record in Mexico or Cen- tral America prior to the late Pleistocene. Embal- lonurids are known from Colombia and Peru from the late Oligocene to the late Miocene, between about 25 and 10 million years ago (Czaplewski, 1997; Czaplewski et al., 2003b; Antoine et al., 2016), but are unrecorded in South America be- tween the late Miocene and late Pleistocene. Sure- ly, the absence of emballonurids from tropical re- gions of both North America and South America from the late Miocene to the late Pleistocene is a collecting bias, considering the overall rarity of pre-late Pleistocene fossil deposits containing bats in the New World tropics. Bats are known from two early Miocene faunas in Panama, but no em- ballonurids were reported (Morgan et al., 2013). Based on the information in the previous paragraphs, there are two conflicting hypotheses for the origin of the Neotropical Emballonuridae. The fossil evidence presented here suggests the possibility that an ancestral emballonurid from Eurasia dispersed overland to North America in the Eocene, with subsequent overwater dispersal of an ancestral emballonurine southward from tropical North America across the CAS to South America prior to the late Oligocene. A second hypothesis based primarily on molecular data from modern emballonurids proposed an Oligocene (more like- ly Eocene) overwater dispersal of emballonurids from Africa to South America across the Atlantic Ocean (Teeling et al., 2005; Lim, 2007, 2009), which would also require a subsequent overwater dispersal across the CAS from South America to North America prior to the early Oligocene. A phy- logenetic analysis of the New World Emballonuri- dae indicates that all living Neotropical members of this family form a monophyletic group, the tribe Diclidurini, within the subfamily Emballonurinae (Lim, 2007; Lim et al., 2008). The two reported genera of Miocene emballonurids from the New World belong to the Diclidurini, the extinct genus Floridopteryx from the early Miocene of Florida and the somewhat younger record of the extant genus Diclidurus from the middle Miocene of Colombia (Czaplewski, 1997; Czaplewski et al., 2003b; this paper). Dental morphology indicates that Oligopteryx from the Oligocene of Florida may be a basal diclidurine, although we do not formally refer this extinct genus to a subfamily or tribe within the Emballonuridae. It is also possible that the Emballonuridae from North America and South America had a dual origin, from Eurasia and Africa, respective- ly, and that only the South American diclidurines survived and the North American members of the family became extinct after the early Miocene. Based on this hypothesis, the Oligocene and Mio- cene emballonurids from Florida and the Oligo- cene, Miocene, and modern emballonurids from South America would not be monophyletic. How- ever, our morphological analysis suggests the New World emballonurines are monophyletic (Tribe Diclidurini), with Floridopteryx being a member of the Diclidurini and Oligopteryx basal to the Di- clidurini if not a member of this tribe. The known South American Tertiary emballonurid fossils are very fragmentary, consisting entirely of isolated teeth, and as such provide little information per- taining to the phylogeny or monophyly of New World emballonurids. Additional information from the fossil re- cord is required before we can fully understand the evolutionary history of the Emballonuridae in the New World, specifically if the Neotropi- cal members of this family (tribe Diclidurini) had an African or Eurasian origin or possibly a more complicated dual origin. Over the past several de- cades, discoveries of Oligocene and Miocene em- ballonurids in both North America (Florida) and South America (Colombia and Peru) have greatly improved our knowledge of the fossil history of this family in the New World. Future discoveries of emballonurid fossils in Oligocene or older sites in North America and/or South America should 210 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) provide additional data on the antiquity of this family in the Western Hemisphere and its possible origin(s). BrIeF evoluTIonary hIsTory oF new world noc- TIlIonoIdea, molossIdae, and vesperTIlIonIdae The fossil record and evolutionary history of several other families of New World bats are per- tinent to a better understanding of the origin and biogeography of the Neotropical Emballonuridae. In addition to the Emballonuridae, three other families of bats, Phyllostomidae, Molossidae, and Vespertilionidae, occurred in both North America and South America prior to the onset of the Great American Biotic Interchange in the late Miocene (~9 Ma). An undescribed new genus of phyllosto- mine phyllostomid occurs in two early Miocene faunas (~18-21 Ma) in Panama in southernmost North America (Morgan et al., 2013) and an in- determinate phyllostomine was identified from a fauna of similar early Miocene age (~21 Ma) from Argentina in southern South America (Czaplews- ki, 2010). Four phyllostomids are known from the middle Miocene La Venta Fauna in Colombia (Czaplewski, 1997; Czaplewski et al., 2003b): an extinct genus and two extinct species of large phyl- lostomines, Notonycteris magdalenensis and N. sucharadeus; a smaller phyllostomine similar to the genera Tonatia and Lophostoma; and the lon- chophylline nectar bat Palynephyllum antimaster. Simmons et al. (2020) noted a p3 of late Miocene age (Mayoan SALMA; ~11 Ma) from Contama- na, Peru that appears to belong to a species in the phyllostomid subfamily Stenodermatinae but was identified as the primate Cebuella sp. by Marivaux et al. (2016). This tooth needs to be studied and confirmed but would constitute the oldest fossil re- cord of a fruit-eating stenodermatine bat. The only other pre-late Pleistocene records of phyllostomids from South America are two recently reported hu- meri of the vampire bat genus Desmodus, one each from Uruguay and Venezuela (Ubilla et al., 2019; Czaplewski and Rincon, 2020). The precise age of these two faunas is unclear but they are either late Pliocene or early Pleistocene. Besides the two ear- ly Miocene phyllostomine records from Panama, the only other pre-late Pleistocene phyllostomids from North America are post-Interchange records of Desmodus from the early Pleistocene of Florida, including the latest Blancan Inglis 1A LF and early Irvingtonian Haile 16A and Haile 21A LFs (Mor- gan et al., 1988). A similar evolutionary history to that of the New World Emballonuridae has been proposed for the chiropteran superfamily Noctilionoidea that includes five extant endemic Neotropical families, Phyllostomidae, Mormoopidae, Noctilionidae, Fu- ripteridae, and Thyropteridae, as well as the ex- tinct family Speonycteridae from the Oligocene of Florida (Czaplewski and Morgan, 2012) and the Mystacinidae from the Miocene to modern fauna of New Zealand and the Oligocene and Miocene of Australia (Hand et al., 2005). Most previous hy- potheses have proposed a Gondwanan origin for the Noctilionoidea, when Australia, Africa, and South America were connected through Antarctica, as re- cently as the late Eocene (Hand et al., 2005; Teel- ing et al., 2005; Gunnell et al., 2014). Gunnell et al. (2014) suggested that noctilionoids reached South America through a series of dispersal events from Australia across Antarctica. The main difference between the proposed evolutionary histories for the New World Emballonuridae and Noctilionoidea is that the emballonurids are thought to have been de- rived from Africa (Teeling et al., 2005; Lim, 2007, 2010), whereas noctilionoids had an Australian origin (Hand et al., 2005; Gunnell et al., 2014). As with the Emballonuridae, a North American/Eur- asian origin has also been proposed for the Noc- tilionidea, based on the occurrence of the oldest known members of this superfamily in the Oligo- cene of Florida, consisting of an extinct genus and species in the Mormoopidae and the extinct family Speonycteridae (Czaplewski and Morgan, 2012; Morgan et al., 2019). The oldest member of the extant family Mormoopidae, the extinct genus and species Koopmanycteris palaeomormoops (Mor- gan et al., 2019), and two species in the extinct ge- nus Speonycteris, S. aurantiadens and S. naturalis, in the extinct basal noctilionoid family Speonyc- teridae, have been described from the same two faunas in Florida that produced Oligopteryx flori- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 211 danus and O. hamaxitos, the early Oligocene I-75 LF and the late Oligocene Brooksville 2 LF. If the Noctilionoidea and Emballonuridae first dispersed to South America from either Africa or Australia via Antarctica, then their occurrence in the early Oligocene of Florida would require pre-Oligocene fossils of both groups from South America (neither of which are known that early from the continent) and two separate pre-Oligocene overwater disper- sal events, first from Africa or Australia/Antarctica to South America and then from South America to North America across the CAS. Species in the Molossidae occur in Oligo- cene and Miocene faunas in both North America and South America. Molossids have been identi- fied in eight Tertiary faunas in South America, including four described species in two genera: three extinct species in the extant genus Mormop- terus, M. faustoi from the late Oligocene Tremem- bé Fauna in Brazil, M. barrancae from the early Miocene Gran Barranca Fauna in Argentina, and M. colombiensis from the middle Miocene of La Venta, Colombia; and the extinct genus and species Potamops mascahehenes from La Venta (Paula Couto, 1956; Czaplewski, 1997, 2010; Czaplews- ki et al., 2003b). An indeterminate species of the extant molossid genus Eumops is known from La Venta, and there are also several South American records of Eocene, Oligocene, and Miocene molos- sids that were not identified below the family level (Czaplewski, 1996a, 1997, 2010; Czaplewski et al., 2003b; Antoine et al., 2016; Czaplewski and Campbell, 2017). North American Tertiary Molossidae are known from seven faunas, only one of which has been formally described. The oldest known molos- sid is from North America, Wallia scalopidens from the middle Eocene (Uintan NALMA) of Swift Cur- rent Creek, Saskatchewan, Canada (Storer, 1984; Legendre, 1985), although Smith et al. (2012) considered the familial status of Wallia uncertain pending the discovery of more complete speci- mens. The Swift Current Creek Fauna is 42-44 Ma in age (Storer, 1984), whereas the earliest molos- sid from the Old World, Cuvierimops from the late Eocene of France, is slightly younger at about 39 Ma (Legendre, 1985; Maitre, 2014). Antoine et al. (2016) listed lower teeth of a molossid from an early middle Eocene (Barrancan SALMA; 41-42 Ma) fauna in Contamana, Peru, but did not illus- trate or describe these specimens. Czaplewski et al. (2003a) described and illustrated an upper molar of an indeterminate genus of molossid from the late Oligocene Brooksville 2 LF and upper molars of two distinct species similar to Tadarida or Mor- mopterus from the early Miocene Thomas Farm, none of which were named. Several lower jaws and partial humeri of an unidentified molossid were re- ported from the early Miocene (early Hemingford- ian) Miller site in Florida (Morgan and Czaplewski, 2012). There are three records of molossids from post-Interchange late Pliocene (Blancan) faunas in North America: a lower premolar from the Deer Park LF in Kansas (Czaplewski et al., 2018) and a distal humerus from the Macaphalt Shell Pit LF in southern peninsular Florida (Czaplewski et al., 2003a), both referred to the extant genus Tadarida; and an M3 tentatively referred to the extant spe- cies Eumops perotis from the McRae Wash LF in Arizona (Czaplewski (1993). Eumops also occurs in the middle Miocene of Colombia suggesting a South American origin for this genus. Bats similar to or congeneric with Tadarida are known from the early Miocene of Florida but are unknown from pre-late Pleistocene faunas in South America, indicating a northern (North Amer- ican or Eurasian) origin for this genus. Tadarida is also known from Oligocene through the present in Europe (Gunnell and Simmons, 2005; Simmons, 2005). Because of the widespread occurrence of molossids in Tertiary faunas in North America, South America, and the Old World (Gunnell and Simmons, 2005), as well as the lack of a well-con- strained phylogeny that includes both living and extinct members of this family, it is premature to propose an evolutionary history for the New World members of this group. An African, Eurasian, or North American origin are all possible, and a dual origin for certain groups within the family is also a possibility. Further studies of modern and fos- sil molossids in both North and South America, as well as the recovery of additional fossils, are neces- 212 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) sary before we can develop a better understanding of the fossil history of this family in the Western Hemisphere. Czaplewski et al. (2003b) tentatively identi- fied the first member of the Vespertilionidae from the Tertiary of South America, based on a lower molar from the middle Miocene La Venta Fauna in Colombia. The Vespertilionidae is the most com- mon family of bats in Tertiary faunas in North America (Czaplewski et al., 2008), and species of vespertilionids also dominate the modern North American chiropteran fauna (Simmons, 2005). The fossil record suggests the Vespertilionidae had a northern origin in Eurasia or North America (Gunnell and Simmons, 2005), and dispersed from North America to South America across the CAS sometime before the middle Miocene. Four families of bats (Emballonuridae, Phyl- lostomidae, Molossidae, Vespertilionidae) have been recorded in both North America and South America in the middle Miocene or earlier (>12 Ma), well before the Great American Biotic Inter- change began at about 9 Ma (Morgan, 2008; Wood- burne, 2010). Clearly, the overwater dispersal capabilities of bats are greater than those of non- volant mammals, and as such bats should be evalu- ated separately when discussing mammals and the GABI. North America and South America were not known to share a single family of non-volant mammals prior to late Miocene onset of the GABI, until recently when Bloch et al. (2016) described the cebid monkey Panamacebus transitus from the early Miocene (late Arikareean) Lirio Norte fauna in Panama, the southernmost extension of North America in the Miocene. Tertiary fossils of platyr- rhine primates, and specifically the family Cebidae, were previously known only from South America. The Lirio Norte fauna and the somewhat younger early Miocene (early Hemingfordian) Centenario Fauna, both from the Panama Canal area, also pro- duced an as-yet-undescribed new genus of phyl- lostomid bat, another group previously thought to have been confined to South America before the Interchange (Morgan et al., 2013). Both the Lirio Norte and Centenario faunas also contain vertebrae of the boid snake Boa of South American origin (Head et al., 2012). Early Miocene deposits from Panama have produced a diverse paleoflora with strong South American or Gondwanan affinities (Jaramillo et al., 2014). The Panama fossils con- firm that an interchange of vertebrates and plants occurred between the tropical regions of North America and South America in the early Miocene (~18–21 Ma), when the two continents were sepa- rated by a seaway that may have been as narrow as 200 km (Montes et al., 2010), long before the beginning of the better-known GABI. Other than the cebid monkey, phyllostomine bat, boid snake, and a few species of freshwater reptiles, including several turtles and a caiman, almost all other verte- brates from the Lirio Norte and Centenario faunas are North American in origin. These two Panama- nian early Miocene faunas contain a diverse assem- blage of large mammals with North American af- finities including amphicyonid ( Amphicyonidae) and procyonid (Procyonidae) carnivorans, horses (Equidae), rhinos (Rhinocerotidae), chalicotheres (Chalicotheriidae), peccaries (Tayassuidae), an- thracotheres (Anthracotheriidae), oreodonts (Oreo- dontidae), camels (Camelidae), and protoceratids (Protoceratidae), as well as three North American families of rodents, the extinct Jimomyidae and the extant Sciuridae and Heteromyidae (MacFadden et al., 2014; Bloch et al., 2016). TERTIARY FOSSIL RECORD OF THE NEO- TROPICAL CHIROPTERA: AN UPDATE This contribution is the last paper in a series de- scribing six new genera and eight new species of Oligocene and Miocene bats from Florida belong- ing to four families with tropical affinities. Two of these are extant families now endemic to the New World tropics (Mormoopidae, Natalidae), one is an extant family with a pantropical distribution (Em- ballonuridae), and one is an extinct family (Spe- onycteridae) belonging to the superfamily Noctil- ionoidea that also includes five living families en- demic to the Neotropics (Morgan and Czaplewski, 2003; Czaplewski and Morgan, 2012; Morgan et al., 2019; this paper). We present an update of our review from a MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 213 decade ago of the Tertiary fossil record of New World bats with tropical affinities (Morgan and Czaplewski, 2012). Although we realize this dis- cussion includes some repetition from previous sections of this paper, our intention is to provide a comprehensive, stand-alone status report of the Tertiary chiropteran fauna with Neotropical af- finities from the Western Hemisphere. These re- cords include not only localities within the modern Neotropical biogeographic region, primarily from South America, but also the diverse fauna of bats with Neotropical affinities mentioned above from mid Cenozoic localities in Florida, which is located in the modern Nearctic biogeographic region. It is important to clarify here that the current biogeo- graphic regions are based on the historic flora and fauna (Olson et al., 2001), whose ecosystems have been radically altered by the 21st century (Keith et al., 2022), and that those regions may have had quite different geographic boundaries in the geo- logic past. The current Neotropical region includes South America, the West Indies, Central America, and Mexico north to the Tropic of Cancer at about 23°N. The Nearctic region includes the United States, Canada, and the temperate region of north- ern Mexico south to the Tropic of Cancer. Prior to the Great American Biotic Interchange beginning in the late Miocene (~9 Ma), fossil evidence suggests the Neotropical region would have had very differ- ent boundaries, including the then-island continent of South America and probably the West Indies but not Mesoamerica (Mexico south to Panama) which was a tropical province of the Nearctic region at that time. As discussed above, early Miocene fau- nas from Panama consist primarily of North Amer- ican species of mammals (MacFadden et al., 2014; Bloch et al., 2016), even though Panama’s current mammalian fauna has a strong Neotropical compo- nent (Eisenberg, 1989). Over the past three decades, our knowledge of the fossil history of the Neotropical Chiroptera has improved dramatically. Before 1990, only three named Tertiary taxa with Neotropical affini- ties belonging to two families (Phyllostomidae, Molossidae) had been described from the Western Hemisphere: an extinct genus and species of phyl- lostomine phyllostomid, Notonycteris magdale- nensis, from the middle Miocene La Venta Fauna in Colombia (Savage, 1951); an extinct species be- longing to an extant genus of molossid, Mormop- terus faustoi, from the late Oligocene Tremembé Fauna in Brazil (Paula Couto, 1956); and the earli- est known molossid, Wallia scalopidens, from the middle Eocene of Saskatchewan, Canada, original- ly named as a proscalopid mole (Storer, 1984) but later referred to the Molossidae (Legendre, 1985; also see Smith et al. 2012). Since 1990, Tertiary fossils representing eight of the nine families of bats now inhabiting the Neotropical region have been reported, includ- ing fossils from both South America and North America. The only Neotropical bat family current- ly lacking a Tertiary fossil record is the Furipteri- dae. Most of these new paleontological discover- ies are from four widely separated regions: five early Oligocene through early Miocene faunas in peninsular Florida in southeastern North America (Czaplewski et al., 2003a; Morgan and Czaplews- ki, 2003, 2012; Czaplewski and Morgan, 2012; Morgan et al., 2019; this paper); and three areas in South America: the middle Miocene of Colombia (Czaplewski, 1997; Czaplewski et al., 2003b); the middle Eocene through late Miocene in the west- ern Amazon Basin of Peru and Brazil (Czaplewski, 1996a; Czaplewski and Campbell, 2004, 2017; An- toine et al., 2016); and the early Eocene and early Miocene of Patagonia in Argentina (Tejedor et al., 2005; Czaplewski, 2010). Table 5 is a current list of all described taxa of Tertiary bats from South America and North America with Neotropical affinities, also includ- ing records identified only to the family or genus level. The location of these sites is indicated on maps of South America (Fig. 20) and North Amer- ica (Fig. 21). Two Eocene records of Molossidae are included in Table 5 but otherwise Eocene bats are excluded because their systematic relationships and biogeographic affinities with modern families are not well understood. Almost all Tertiary re- cords of bats from South America are included on this list, whereas the records from North America exclude the Vespertilionidae. The majority of post- 214 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Eocene Tertiary bats from North America have been referred to the Vespertilionidae (Czaplewski et al., 2008), and are not included here because they mostly represent temperate bats lacking ob- vious Neotropical affinities. Although some ex- tinct genera of Vespertilionidae known from North American Tertiary fossil deposits (Czaplewski et al., 2008) may be related to Neotropical vespertil- ionids, the phylogenetic relationships of these ex- tinct genera are not well enough understood to de- termine their biogeographic origins and affinities. Several extant genera of vespertilionids identified from North American Tertiary faunas (e.g., Eptesi- cus, Lasiurus, Myotis) occur in the modern fauna of the Neotropical region in Mesoamerica and South America (Czaplewski et al., 2008). The middle Miocene (~12–13 Ma; Laventan SALMA) La Venta Fauna in Colombia has pro- duced the most diverse Tertiary chiropteran fauna from the Western Hemisphere, numbering 14 spe- cies, including 12 genera (nine identified genera and three indeterminate but distinct genera) and six families (Czaplewski, 1997; Czaplewski et al., 2003b; Table 4). Among the nine families of bats known from the modern fauna of South America, only three are missing from the La Venta Fauna, Furipteridae, Mormoopidae, and Natalidae. The Furipteridae lack a Tertiary fossil record as noted above, whereas the Mormoopidae and Natalidae occur in the Oligocene and/or early Miocene of North America (Florida) but are unknown in South America before the late Pleistocene (Morgan and Czaplewski, 2003; Morgan et al., 2019). More than half of the La Venta chiropteran fauna belongs to two families: four species in the Phyllostomidae, the phyllostomines Notonycteris magdalenensis and N. sucharadeus, a phyllostomine near Tonatia or Lophostoma, and the lonchophylline Palyne- phyllum antimaster; and four species in the Molos- sidae, Eumops sp., Mormopterus colombiensis, Potamops mascahehenes, and an indeterminate species distinct from the three other molossids. The other six species of bats from La Venta include: two species of Emballonuridae, Diclidurus sp. and a smaller indeterminate genus; one member of Noctilionidae referred to the living species Noctilio Figure 20. Tertiary sites containing bats from South America (Chiroptera: Emballonuridae, Phyllostomidae, Noctilioni- dae, Thyropteridae, Molossidae, Vespertilionidae). Table 4 lists the taxa of bats known from most of these sites. The sites are numbered in order from oldest to youngest. Symbols designate different epochs: Eocene (filled circle); Oligocene (asterisk); Miocene (triangle), Pliocene (plus sign). Eocene: 1. Laguna Fría, Argentina (early Eocene), site not listed in Table 4 because the bat is not identified below the level of Chiroptera; 2. Pozo Formation, Contamana, Peru (middle Eo- cene, Barrancan); Oligocene: 3. Santa Rosa, Peru (early Oli- gocene), site not listed in Table 4 because the bat is not identi- fied below the level of Chiroptera. 4. Tremembé Formation, Brazil, (late Oligocene, Deseadan); 5. Chambira Formation, Contamana, Peru (late Oligocene, Deseadan). Miocene: 6. Gran Barranca, Argentina (early Miocene, Colhuehuapian); 7. Pebas Formation, Contamana, Peru (early Miocene, Col- huehuapian/Santacrucian); 8. La Venta, Colombia (middle Miocene, Laventan); 9. Pebas Formation, Contamana, Peru (late Miocene, Mayoan/Chasicoan); 10. Juruá River, Brazil (late Miocene, Huayquerian); 11. Río Acre, Peru (late Mio- cene, Huayquerian). Pliocene: 12. Kiyú, Uruguay (late Plio- cene-middle Pleistocene); 13. El Breal de Orocual, Venezuela (late Pliocene/early Pleistocene). MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 215 Figure 21. Tertiary sites containing bats with Neotropical affinities from North America (Chiroptera: Emballonuridae, †Speonycteridae, Phyllostomidae, Mormoopidae, Natalidae, Molossidae). Table 4 lists the taxa of bats known from each of these sites. The sites are numbered in order from oldest to youngest. Symbols designate different epochs: Eocene (filled circle); Oligocene (asterisk); Miocene (triangle), Pliocene (plus sign). Eocene: 1. Swift Current Creek, Saskatchewan, Canada (middle Eocene, Uintan). Oligocene: 2. I-75, Florida (early Oligocene, Whitneyan); 3. Brooksville 2, Florida (late Oligocene, early Arikareean); 4. Buda, Florida (latest Oligocene, late Arikareean). Miocene: 5. Lirio Norte, Panama (earliest Miocene, latest Arikareean); 6. Centenario, Panama (early Miocene, early Hemingfordian); 7. Miller, Florida (early Miocene, early Hemingfordian); 8. Thomas Farm, Florida (early Miocene, early Hemingfordian). Pliocene: 9. McRae Wash, Arizona, (late Pliocene, Blancan); 10. Deer Park, Kansas (late Pliocene, Blancan); 11. Macasphalt Shell Pit, Florida (late Pliocene, Blancan). albiventris; two species of Thyropteridae, both re- ferred to living species, Thyroptera lavali and T. cf. tricolor; and an indeterminate genus and species of Vespertilionidae. La Venta is one of the most di- verse Tertiary chiropteran faunas known (Gunnell and Simmons, 2005), and among the richest from a non-karst depositional environment. Two main units within the La Venta stratigraphic section have produced most of the bat fossils, the “Fish Bed” representing a lacustrine depositional environment and the “Monkey Beds” that sample a tropical for- est habitat (Czaplewski, 1997). In comparison, the most diverse Tertiary bat fauna from North America, Thomas Farm, Florida, is an early Miocene karst deposit with nine spe- cies. Sample size is not the reason for the differ- ence in the number bat species in the two faunas, because Thomas Farm has an order of magnitude more individual bat fossils than does La Venta (Pratt, 1989; Czaplewski, 1997; Czaplewski et al, 2003b; Morgan and Czaplewski, 2003; this paper). At present, Colombia has well over 100 species of bats, whereas Florida has only 20 species (Sim- mons, 2005; Marks and Marks, 2006). The fossil record suggests the much higher diversity of bats in Colombia compared to Florida was also typical of Miocene faunas. This pattern reflects the well- known observation that chiropteran diversity or species richness in the tropics is far greater than in temperate regions (Hill and Smith, 1984; Nowak, 1994). La Venta is about 3° North of the Equator whereas Thomas Farm is 30° North. The remainder of the Tertiary chiropteran re- cord from South America is widely scattered, with records from Argentina, Uruguay, southern Brazil, the western Amazon basin in Brazil and Peru, and Venezuela, and mostly consists of small samples of isolated teeth and other fragmentary speci- mens (Table 5). No more than three taxa of bats are known from any of these localities. Until re- cently, there was only one record of an Eocene bat from South America, consisting of two teeth of an indeterminate family from the Laguna Fría site in Patagonia, Argentina of early Eocene age (M. Teje- dor et al., 2005). A lower molar of a bat of indeter- minate affinity from the Santa Rosa LF in Ama- zonian Peru was originally considered late Eocene in age (Czaplewski and Campbell, 2004), but is now thought to be early Oligocene (Seiffert et al. 2020; K. Campbell, pers. communication). A small sample of isolated bat teeth was recently reported from sediments representing a tropical rainforest habitat in the middle Eocene (Barrancan SALMA) Pozo Formation in Contamana, Amazonian Peru (Antoine et al., 2012, 2016 and supplementary data). These fossils were identified in a table in a 216 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Table 5. Tertiary bats from South America and North America with Neotropical affinities. Pleistocene records are not included in this table. The order of families follows Simmons (2005). Within a family (subfamily for the Phyllostomidae), the genera and species are listed in alphabetical order, with undescribed or indeterminate genera at the end of the family. Extinct taxa are indicated by a dagger (†). The type locality for a species is indicated by an asterisk (*). Abbreviations: indet. (indeterminate, refers to fossils that are too incomplete for a positive identification to a higher taxonomic level than indicated below); NALMA (North American land mammal age), SALMA (South American land mammal age). Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 217 Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Table 5. Cont. Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. 218 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Element, species, site, proximal proximal width of distal distal distal and catalog number width depth shaft width width depth maximum articular surface Humerus Oligopteryx floridanus Brooksville 2 UF 179904 (paratype) 4.3 4.8 1.6 – – – UF 179935 4.1 4.8 1.7 – – – UF 179936 4.3 5.0 – – – – UF 179959 4.2 4.9 1.7 – – – UF 179963 4.1 – – – – – UF 179909 – – 1.4 3.7 2.8 1.8 UF 179910 – – 1.7 3.8 2.9 2.1 UF 179939 – – – 3.9 – 1.9 UF 179964 (paratype) – – 1.6 3.8 3.0 1.9 UF 179965 – – 1.5 3.6 2.9 1.9 UF 179966 – – 1.6 3.6 2.8 1.9 UF 179967 – – 1.5 3.6 2.9 2.0 UF 179968 – – – 3.9 – 1.7 N 5 4 9 8 6 8 M 4.2 4.9 1.6 3.7 2.9 1.9 OR 4.1– 4.8– 1.4– 3.6– 2.8– 1.7– 4.3 5.0 1.7 3.9 3.0 2.1 I-75 UF 121710 – – – 3.9 2.8 2.0 Oligopteryx hamaxitos Brooksville 2 UF 182792 (paratype) – – 1.2 3.1 2.1 1.4 I-75 UF 121714 – – 1.3 2.8 2.3 1.5 Radius Oligopteryx floridanus Brooksville 2 UF 179911 (paratype) 2.8 3.2 1.5 – – – UF 179912 – 3.3 1.6 – – – UF 179913 2.9 3.2 – – – – UF 179940 2.9 3.3 1.6 – – – UF 179942 2.8 3.2 1.6 – – – UF 179969 2.8 3.1 1.5 – – – UF 179971 2.9 3.1 1.6 – – – UF 179972 2.9 3.3 1.6 – – – UF 179981 2.9 3.2 1.6 – – – UF 179982 2.8 3.1 1.5 – – – UF 179944 – – 1.4 2.5 – 2.0 UF 179974 – – 1.3 2.5 – 1.9 UF 179975 – – 1.5 2.5 – 1.9 UF 209956 – – 1.3 2.6 – 1.9 UF 209957 – – 1.4 2.5 – 1.8 N 9 10 14 5 – 5 M 2.9 3.2 1.5 2.5 – 1.9 OR 2.8– 3.1– 1.3– 2.5– – 1.8– 2.9 3.3 1.6 2.6 – 2.0 I-75 UF 121711 3.0 3.2 – – – – Radius (cont.) Oligopteryx hamaxitos Brooksville 2 UF 179914 2.0 2.1 1.2 – – – UF 179915 2.1 2.2 1.1 – – – UF 179983 (paratype) 2.1 2.2 1.1 – – – UF 179977 – – 1.1 2.1 – 1.4 N 3 3 4 – – – M 2.1 2.2 1.1 OR 2.0– 2.1 1.1– 2.1 2.2 1.2 Table 5. Cont. Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. 1Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. 6This record of cf. Desmodus sp. from Venezuela was considered late Pliocene or early Pleistocene in age (Czaplewski and Rincon, 2020). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainty in the age, the SALMA is not provided. 7First reported as the primate Cebuella (Marivaux et al., 2016) but reidentified as the lower premolar of a stenodermatine phyllostomid (Simmons et al., (2020). 8Originally described as the extinct species †Thyroptera robusta but synonymized with the living species T. lavali by Czaplewski (1996b). 9These two late Pliocene records of an indeterminate species of Tadarida may represent the same species but consist of elements that are not comparable. 10These two early Miocene records of Tadarida/Mormopterus from Thomas Farm are distinct species based on a significant difference in size (Czaplewski et al., 2003a). 11The two records of molossids from Contamana, Peru listed here as genus and species indet. almost certainly represent two distinct taxa based on their difference in age. 12This is the only Tertiary record of the Vespertilionidae in South America. Numerous genera and species of vespertilionids are known from Tertiary faunas in North America (Czpalewski et al., 2008), but are not listed here because they do not have obvious Neotropical affinities. Family, genus, and species Locality/localities Age and NALMA/SALMA References and country/US state Emballonuridae Diclidurus species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Floridopteryx poyeri Thomas Farm, Florida* early Miocene, early Hemingfordian This paper †Karstopteryx gunnelli Buda, Florida* latest Oligocene, late Arikareean This paper †Oligopteryx floridanus I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean †Oligopteryx hamaxitos I-75, Florida early Oligocene, Whitneyan This paper Brooksville 2, Florida* late Oligocene, early Arikareean genus and species indet.1,2 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) genus and species indet.1,3 Contamana, Peru late Oligocene, Deseadan Antoine et al. (2016) Chambira Formation Contamana, Peru early Miocene, Colhuehuapian- Antoine et al. (2016) Pebas Formation Santacrucian Contamana, Peru late Miocene, Mayoan-Chasicoan Antoine et al. (2016) Pebas Formation †Speonycteridae4 †Speonycteris aurantiadens I-75, Florida early Oligocene, Whitneyan Czaplewski and Morgan (2012) Brooksville 2, Florida* late Oligocene, early Arikareean †Speonycteris naturalis I-75, Florida* early Oligocene, Whitneyan Czaplewski and Morgan (2012) Phyllostomidae4: Phyllostominae †Notonycteris magdalenensis La Venta, Colombia* middle Miocene, Laventan Savage (1951) Czaplewski (1997) Czaplewski et al. (2003b) †Notonycteris sucharadeus La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Tonatia or Lophostoma La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) species indet. Czaplewski et al. (2003b) undescribed genus and species Lirio Norte, Panama early Miocene, late Arikareean Morgan et al. (2013) Centenario, Panama early Miocene, early Hemingfordian genus and species indet.1 Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) Phyllostomidae: Desmodontinae Desmodus sp. Kiyú, Uruguay late Pliocene/middle Pleistocene5 Ubilla et al. (2019) Raigón Formation cf. Desmodus sp. El Breal de Orocual late Pliocene/early Pleistocene6 Czaplewski and Rincon (2020) Venezuela Phyllostomidae: Lonchophyllinae †Palynephyllum antimaster La Venta, Colombia* middle Miocene, Laventan Czaplewski et al. (2003b) Phyllostomidae: Stenodermatinae Contamana, Peru late Miocene, Mayoan Simmons et al. (2020) genus and species indet.7 Pebas Formation Mormoopidae4 †Koopmanycteris palaeomormoops I-75, Florida early Oligocene, Whitneyan Morgan et al. (2019) Brooksville 2, Florida* late Oligocene, early Arikareean Noctilionidae4 Noctilio albiventris La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Noctilio lacrimaelunaris Rio Acre, Peru* late Miocene, Huayquerian Czaplewski (1996a) Thyropteridae4 †Amazonycteris divisus Rio Juruá, Brazil* late Miocene, Huayquerian Czaplewski and Campbell (2017) Thyroptera lavali8 La Venta, Colombia middle Miocene, Laventan Czaplewski (1996b, 1997) Czaplewski et al. (2003b) Thyroptera cf. tricolor La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Natalidae †Primonatalus prattae Thomas Farm, Florida* early Miocene, early Hemingfordian Morgan and Czaplewski (2003) genus and species indet.1 I-75, Florida early Oligocene, Whitneyan Morgan and Czaplewski (2003) Molossidae Eumops cf. perotis McRae Wash, Arizona late Pliocene, Blancan Czaplewski (1993) Eumops species indet. La Venta, Colombia middle Miocene, Laventan Czaplewski (1997) Czaplewski et al. (2003b) †Mormopterus barrancae Gran Barranca, Argentina* early Miocene, Colhuehuapian Czaplewski (2010) †Mormopterus colombiensis La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) †Mormopterus faustoi Tremembé, Brazil* late Oligocene, Deseadan Paula Couto (1956) Mormopterus species indet. Gran Barranca, Argentina early Miocene, Colhuehuapian Czaplewski (2010) †Potamops mascahehenes La Venta, Colombia* middle Miocene, Laventan Czaplewski (1997) Tadarida species indet.9 Macasphalt Shell Pit, Florida late Pliocene, Blancan Czaplewski et al. (2003a) Deer Park, Kansas late Pliocene, Blancan Czaplewski et al. (2018) Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 Tadarida or Mormopterus Thomas Farm, Florida early Miocene, early Hemingfordian Czaplewski et al. (2003a) species indet. 10 †Wallia scalopidens Swift Current Creek* middle Eocene (Uintan) Storer (1984) Saskatchewan, Canada Legendre (1985) genus and species indet.1,11 Contamana, Peru middle Eocene, Barrancan Antoine et al. (2016) Pozo Formation genus and species indet.1,11 Contamana, Peru late Miocene, Mayoan Antoine et al. (2016) Pebas Formation genus and species indet.1 Rio Acre, Peru late Miocene, Huayquerian Czaplewski (1996) Molossidae (cont.) genus and species indet.1 Rio Juruá, Brazil late Miocene, Huayquerian Czaplewski and Campbell (2017) Vespertilionidae genus and species indet.12 La Venta, Colombia middle Miocene, Laventan Czaplewski et al. (2003b) 1 Taxa listed as genus and species indet. (indeterminate) are included in this table only if they are clearly distinct from other described Tertiary members of the family. Taxa identified in the literature as genus and species indet. because of their incomplete or fragmentary condition are not included here. 2 This taxon is much smaller than Diclidurus, the other emballonurid identified from La Venta, Colombia. 3 The three records of emballonurids from Contamana, Peru listed here as genus and species indet. almost certainly represent three distinct taxa based on their difference in age. 4 The extinct family Speonycteridae and the extant families Phyllostomidae, Mormoopidae, Noctilionidae, and Thyropteridae, as well as the Furipteridae (lacks a Tertiary fossil record), are placed in the superfamily Noctilionoidea. 5 This record of Desmodus sp. from Uruguay was considered late Pliocene or early to middle Pleistocene in age (Ubilla et al., 2019). This specimen may represent one of only two Pliocene records of bats from South America. Because of the uncertainly in the age, the SALMA is not provided. supplementary data file and were not described or illustrated. We consider these and other identifica- tions of bats from Contamana (see Oligocene and Miocene bats below and in Table 5) to be tentative pending further comparisons. Lower teeth identi- fied as Molossidae from a middle Eocene fauna in Contamana would be among the earliest records of this family. A fragmentary lower tooth identified as Phyllostomidae would be by far the oldest re- cord of this family, otherwise unknown before the early Miocene (see below), although we question whether a partial lower tooth is identifiable to the family level. The Oligocene record of Chiroptera from South America is somewhat better than the Eocene record. Two partial skeletons of bats are known from lacustrine deposits of the Tremembé Forma- tion of late Oligocene age (Deseadan SALMA) in the state of São Paulo in southern Brazil, including Mormopterus faustoi, an extinct species of an ex- tant genus of Molossidae, and a second more com- plete but crushed skeleton of unknown familial af- finity (Paula Couto, 1956; Mezzalira, 1966; Paula Couto and Mezzalira, 1971). Lacustrine sediments of the Chambira Formation of late Oligocene age (Deseadan) from Contamana, Peru, have produced two lower teeth of an emballonurid and an upper molar and upper and lower canines of a supposed “rhinolophoid” (Antoine et al., 2016, and supple- mentary data), a group otherwise unknown from the New World, unless emballonurids are consid- ered rhinolophoids. We consider the identification MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 219 of rhinolophoids from Contamana to be question- able pending further comparisons. Compared to the Eocene or Oligocene, bats are much better known from the South American Miocene (~7–20 Ma), including 14 species from the middle Miocene of La Venta, Colombia listed above (Czaplewski, 1997; Czaplewski et al., 2003b; Table 4). Five other Miocene faunas from South America have produced bats. The early Miocene Gran Barranca Fauna (Colhuehuapian SALMA) from Patagonia in southern Argentina consists of three species (Czaplewski, 2010): a lower mo- lar of an indeterminate genus of phyllostomine phyllostomid; a mandible with p3-m2 named as the new species Mormopterus barrancae; and a smaller species of Mormopterus. Two faunas from the Miocene Pebas Formation in Contamana, Peru have produced bats (Antoine et al., 2016 and sup- plementary data; Table 5): an emballonurid from an early Miocene fauna (Colhuehuapian or Santa- crucian SALMA) and a different taxon of embal- lonurid, a molossid, and a possible stenodermatine phyllostomid (Simmons et al., 2020) from a late Miocene fauna (Mayoan or Chasicoan SALMAs). Late Miocene (Huayquerian SALMA) faunas from exposures along three rivers in the western Ama- zon basin in Brazil and Peru, Rio Acre, Rio Purus, and Rio Juruá, have produced bats. The only ex- tinct species in the Noctilionidae, Noctilio lacri- maelunaris, was described from the Rio Acre and a small molossid of an indeterminate genus was identified from the Rio Purus, both referred to the Rio Acre Fauna from Peru (Czaplewski, 1996a). An extinct genus and species in the Thyropteridae, Amazonycteris divisus, was described from the Rio Juruá in Brazil, together with an isolated tooth of an indeterminate genus of molossid (Czaplewski and Campbell, 2017). Until recently, Pliocene bats were unknown from South America. Ubilla et al. (2019) reported a possible Pliocene bat from the Kiyú site in Uru- guay, a humerus of a large species of the vampire bat Desmodus (Phyllostomidae: Desmodontinae) The age of the fossil vampire from Uruguay is somewhat problematic, with the authors suggest- ing an age range between late Pliocene and middle Pleistocene (Ubilla et al., 2019). Shortly thereafter, Czaplewski and Rincon (2020) reported another humerus of a large species of Desmodus from the El Breal de Orocual asphaltic deposit (tar pit) in Venezuela of either late Pliocene or early Pleisto- cene age. Although these two Desmodus records from Uruguay and Venezuela may be early Pleisto- cene rather than Pliocene, and thus not technically Tertiary, they represent the only South American fossil records of the Chiroptera in the time interval spanning the late Miocene to the late Pleistocene. The Venezuelan record of Desmodus is intriguing because it is from northern South America and is similar in age to the oldest record of Desmodus in North America, from the early Pleistocene (latest Blancan, ~2 Ma) Inglis LF in Florida (Morgan et al., 1988). The rarity of Pliocene bats from South America hinders our ability to adequately evaluate the biogeographic history of the Neotropical chi- ropteran fauna. This is especially relevant because one of the most important biogeographic events in the history of the Western Hemisphere, the Plio- cene to early Pleistocene (~1–5 Ma) phase of the Great American Biotic Interchange, began in the early Pliocene with the initial connection of North America and South America at the Panamanian isthmus (O’Dea et al., 2016). The discovery of a diverse fauna of bats with tropical affinities from the Oligocene and early Miocene of Florida (Czaplewski et al., 2003a; Morgan and Czaplewski, 2003, 2012; Czaplewski and Morgan, 2012; Morgan et al., 2019; this pa- per), including the Emballonuridae, Mormoopidae, extinct noctilionoid family Speonycteridae, Natali- dae, and Molossidae, has significantly improved the chiropteran fossil record but has also complicated the narrative regarding the New World origins and evolutionary histories of these groups. Two Oligo- cene bat faunas from peninsular Florida are almost entirely composed of species with Neotropical af- finities, the early Oligocene (~30 Ma) I-75 LF and the late Oligocene (~26–28 Ma) Brooksville 2 LF. These two sites have been discussed in detail above as they have produced samples of a new genus and two new species of emballonurids, Oligopteryx floridanus and O. hamaxitos, described here. These 220 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) two sites have similar chiropteran faunas, including (Czaplewski et al., 2003a; Morgan and Czaplews- ki, 2003; Czaplewski and Morgan, 2012; Morgan et al., 2019; this paper; Table 5): the two embal- lonurids; an extinct genus and species of Mor- moopidae, Koopmanycteris palaeomormoops; and an extinct genus and species of basal noctilionoid in the extinct family Speonycteridae, Speonycteris aurantiadens. I-75 has two additional taxa with Neotropical affinities not found in Brooksville 2, Speonycteris naturalis and an indeterminate genus of Natalidae, and Brooksville 2 has an indetermi- nate genus of Molossidae not found in I-75. The mormoopid and natalid from I-75 are the oldest known members of those two families and the two species of Oligopteryx from this site are the oldest emballonurids from the New World. A single upper molar of a vespertilionid from I-75 represents the only bat from this site or Brooksville 2 that does not have clear Neotropical affinities. Another new genus and species of emballonurid, Karstopteryx gunnelli, occurs in the latest Oligocene (~24 Ma) Buda LF in Florida. The early Miocene (~18 Ma) Thomas Farm LF adds four species with Neotropi- cal affinities (Czaplewski et al., 2003a; Morgan and Czaplewski, 2003; this paper): the diclidurine emballonurid Floridopteryx poyeri, described here; the extinct genus and species of natalid Primona- talus prattae; and two molossids near Tadarida or Mormopterus. Floridopteryx is the earliest diclidu- rine and Primonatalus prattae is the oldest named natalid. The early Miocene Miller LF from Florida has an unidentified molossid that may be similar to one of the two molossids from the slightly younger Thomas Farm LF (Morgan and Czaplewski, 2012). As discussed in more detail above under Paleoecol- ogy, the occurrence of a diverse fauna of bats with tropical affinities corroborates a tropical or sub- tropical climate in peninsular Florida during the Oligocene and early Miocene. The disappearance of Neotropical bats from Florida after the early Miocene suggests a change to a more temperate climate as indicated by the presence of a middle Miocene paleoflora in the Florida panhandle with warm temperate affinities (Jarzen et al., 2010; Lott et al., 2019). The rarity of bats with tropical affini- ties from North American Tertiary sites outside of Florida probably reflects the more temperate cli- mate in western North America after the Eocene (Czaplewski et al., 2008; Morgan and Czaplewski, 2012). Excluding a rather diverse fauna of vesper- tilionids from Oligocene, Miocene, and Pliocene sites from Florida and the western US of presumed temperate affinities, only about half a dozen other Tertiary records of bats with Neotropical affinities are known from North America (Czaplewski et al., 2008, 2018; Morgan et al., 2013). The oldest is the molossid Wallia scalopidens from the middle Eo- cene (late Uintan) Swift Current Creek LF in Sas- katchewan, Canada (Storer, 1984; Legendre, 1985), but also see Smith et al. (2012) who questioned the molossid affinities of Wallia. A somewhat older re- cord, from the middle Eocene (Bridgerian NAL- MA) Tabernacle Butte LF in Wyoming (McKenna et al., 1962), has been questionably allied with the Molossidae (Legendre, 1985). Two new genera and three new species of early middle Eocene (Bridg- erian) bats were recently described from the El- derberry Canyon LF in Nevada (Czaplewski et al., 2022). Among these three species, Volactrix sim- monsae and Palaeochiropteryx sambuceus belong to extinct, archaic bat families and Sonor handae appears to be an early member of the Vespertil- ionidae. None of these new middle Eocene species has obvious Neotropical affinities. An undescribed genus and species of phyllostomine phyllostomid from the early Miocene Lirio Norte and Cente- nario local faunas in Panama is one of the earli- est known members of the Phyllostomidae and the only Tertiary bat reported from Central America (Morgan et al., 2013). Three other North Ameri- can records of bats with Neotropical affinities are post-Interchange molossids (Table 5; Czaplewski, 1993; Czaplewski et al., 2003a, 2018): Eumops cf. perotis from McRae Wash, Arizona and Tadarida sp. from Deer Park, Kansas are both late Pliocene (early Blancan NALMA) in age, whereas a record of Tadarida sp. from Macasphalt Shell Pit, Florida is early Pleistocene (late Blancan). Two bats from late Pliocene and early Pleis- tocene Interchange sites in North America appear MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 221 to be of South American origin and were prob- ably participants in the Plio-Pleistocene phase of the GABI, the large extant molossid Eumops cf. perotis from the late Pliocene McRae Wash LF in Arizona (Czaplewski, 1993) and the extinct des- modontine vampire bat Desmodus archaeodaptes (Phyllostomidae) from three early Pleistocene sites in Florida (Morgan et al., 1988). Eumops sp. is first known from the middle Miocene La Venta Fauna in Colombia (Czaplewski, 1997; Czaplewski et al., 2003b). The genus Desmodus is now known from five sites that date to the late Pliocene/early Pleisto- cene phase of the GABI, a large species, Desmodus sp., similar to the extinct D. draculae, from Uru- guay and Venezuela (Ubilla et al., 2019; Czaplews- ki and Rincon, 2020) and a smaller extinct species, D. archaeodaptes, from the early Pleistocene (~1-2 Ma) Inglis 1A, Haile 16A, and Haile 21A LFs (type locality) in Florida (Morgan et al., 1988). It seems most likely that vampire bats evolved in South America and followed their favored prey/blood donor species (large xenarthans?) northward into North America during the Interchange (Morgan et al., 1988; Morgan, 1991). Two Neotropical chirop- teran families, Mormoopidae and Natalidae, are present in several Oligocene and/or early Miocene faunas in Florida but are unknown in South Ameri- ca before the late Pleistocene. Mormoopids and na- talids apparently evolved in North America in the Oligocene if not earlier, and were also participants in the GABI, dispersing southward across the Pan- amanian isthmus to South America sometime after the early Pliocene (Morgan and Czaplewski, 2003; Morgan et al., 2019). Previous hypotheses on the origins and evolutionary histories of the various families of Neotropical bats were based primarily on the geo- graphic distribution of the modern chiropteran fauna (Koopman, 1970, 1976, 1982), and more re- cently including phylogenetic relationships based on molecular genetics, with minimal input from the fossil record (Teeling et al., 2005; Lim, 2009, 2010). These hypotheses suggested that members of the six endemic Neotropical families, includ- ing five families in the superfamily Noctilionoidea (Furipteridae, Mormoopidae, Noctilionidae, Phyl- lostomidae, Thyropteridae) and the Natalidae, as well as two pantropical families (Emballonuridae, Molossidae), either evolved in South America or reached South America early in their evolution- ary history (Oligocene or Miocene), and then dis- persed northward overwater to the West Indies or Central America before the Pliocene, or overland to Central America following the connection of the two continents at the Panamanian isthmus in the early Pliocene (O’Dea et al., 2016). Prior to their arrival in South America, an African or Gondwa- nan origin has been hypothesized for seven of these bat families (Teeling et al., 2005; Lim, 2009), in- cluding Emballonuridae, the five families in the Noctilionoidea and Molossidae. Among the nine families of bats found in South America, only the New World representatives of the Vespertilionidae were considered to have a North American origin. Discoveries of large samples of fossil bats from the Oligocene and early Miocene of Florida over the past three decades establish that more than half of the New World families with tropical affinities now have a Tertiary fossil record in North America (Czaplewski et al., 2003a; Morgan and Czaplewski, 2003, 2012; Czaplewski and Morgan, 2012; Mor- gan et al., 2013, 2019; this paper), complicating the evolutionary history and biogeography of these groups. With the earliest Western Hemisphere re- cords for the Emballonuridae, Mormoopidae, and Natalidae, as well as the extinct basal noctilionoid family Speonycteridae, from the early Oligocene of Florida, a South American or African/Gondwa- nan origin for the New World members of these groups is in question. The Oligocene to Pliocene fossil record of the Chiroptera from the Western Hemisphere now consists of 11 genera and 17 species from South America and 22 genera and 28 species from North America (Czaplewski, 2005; Czaplewski et al., 2008; Morgan and Czaplewski, 2012; Antoine et al., 2016; Ubilla et al., 2019; Czaplewski and Rincon, 2020; this paper). The numbers of genera in these two faunas, particularly North America, are com- parable to the generic diversity of Oligocene, Mio- cene, and Pliocene chiropteran faunas from Africa and Australia, and are considerably more diverse 222 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) than the post-EoceneTertiary bat fauna from Asia (Gunnell and Simmons, 2005). The North Ameri- can post-Eocene Tertiary bat fauna consists of 10 genera and 14 species belonging to 5 families now primarily tropical in their distribution, mostly from the Oligocene and early Miocene of Florida (Table 5), as well as 12 genera and 15 species of Vesper- tilionidae from Florida and western North America that lack obvious Neotropical affinities (Czaplews- ki et al., 2008; Morgan and Czaplewski, 2012; Morgan et al., 2019; this paper). The combined Tertiary New World chiropteran fauna with Neo- tropical affinities consists of 7 families, 18 genera, and 30 species (Table 5), compared to 2 families, 3 genera, and 3 species known prior to 1990, a nearly four-fold increase in families, six-fold increase in genera, and ten-fold increase in species. Only the molossid genus Eumops is known from the Ter- tiary of both continents, with a middle Miocene record from Colombia and a post-Interchange late Pliocene record from Arizona (Czaplewski, 1993, 1997; Czaplewski et al., 2003b). The significant additions to the New World Tertiary chiropteran record over the past 30 years reflect a concerted effort by a large number of pa- leontologists to collect and screenwash sediments from fossil sites in both North America and South America, in particular from the middle Miocene of La Venta, Colombia (Czaplewski, 1997; Czaplews- ki et al., 2003b; and recent fieldwork by Nancy Sim- mons, Camilo López-Aguirre, and associates), the Eocene through late Miocene of the western Ama- zon Basin in Peru and Brazil (Czaplewski, 1996; Czaplewski and Campbell, 2004, 2017; Antoine et al., 2016), and Oligocene and early Miocene karst deposits in Florida (Pratt, 1989, 1990; Czaplewski and Morgan, 2000, 2012; Czaplewski et al., 2003a; Morgan and Czaplewski, 2003, 2012; Morgan et al., 2019; this paper). With a few notable excep- tions, bats are generally rare in Tertiary fossil sites in the Western Hemisphere, primarily owing to the scarcity of pre-Pleistocene karst deposits in North America, outside of Florida, and the absence of Tertiary karst deposits in South America contain- ing bats. In comparison, widespread karst and la- custrine deposits in Europe have produced the most diverse Tertiary chiropteran fauna known from any of the continents (Sigé and Legendre, 1983; Gun- nell and Simmons, 2005). The ongoing efforts of paleontologists in North America and South Amer- ica to screenwash Tertiary vertebrate fossil depos- its will continue to improve the fossil record of the Chiroptera in the Western Hemisphere. ACKNOWLEDGMENTS Many people have assisted us in our work on the Tertiary bats of Florida. We are indebted to Clayton E. Ray and the late Pierce Brodkorb who initiated a screenwashing program at the Miocene Thomas Farm site in Florida in the 1950s and early 1960s, demonstrating the potential this site held for pro- ducing significant samples of fossil bats, as well as many other groups of microvertebrates. Ann Pratt conducted an intensive screenwashing effort at Thomas Farm in the early 1980s as part of her doctoral dissertation research. Art Poyer has been instrumental in washing and sorting large volumes of microvertebrate-bearing matrix from Thomas Farm over the past 40 years. For almost two de- cades beginning in the early 2000s, David Stead- man of the FLMNH led a major screenwashing program at Thomas Farm as part of his research on the avian fauna from this site, and also collected considerable new material of fossil bats. The com- bined efforts of these dedicated paleontologists in screenwashing and sorting sediments from Thomas Farm have produced the largest Tertiary bat fauna from North America, including the new genus and species of emballonurid, Floridopteryx poyeri, de- scribed herein, and named in honor of Art Poyer. Erika Simons and Richard Hulbert provided field photos of the Thomas Farm site. Ann Pratt and Art Poyer also helped collect, screenwash, and sort fos- sil bats from Brooksville 2. Glynn Hayes provided helpful information on the Brooksville 2 fauna, as well as several field photos. Richard Hulbert of the Florida Museum of Natural History loaned us the fossil samples of em- ballonurid bats from Florida described here. Nancy Simmons of the American Museum of Natural His- tory, Laurie Wilkins of the Florida Museum of Nat- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 223 ural History, Joe Cook, John Dunnum, and William Gannon of the Museum of Southwesten Biology, Linda Gordon and Helen Kafka of the National Museum of Natural History, Smithsonian Institu- tion, and Brandi Coyner and Janet Braun of the Oklahoma Museum of Natural History helped us during our visits to their respective museums and/ or arranged loans of modern comparative speci- mens. Charles Baker made some of the scanning electron micrographs. Steve Westrop and Roger Burkhalter allowed the use of their bellows cam- era, Stackshot rig, focus-stacking software, and provided help in taking photographs of specimens. Tom Gunning gave advice on coating specimens and casts with ammonium chloride for micropho- tography. Michael A. Mares, previous director of the Oklahoma Museum of Natural History, gener- ously provided funds to NJC for travel relevant to this project. We are especially grateful to Suzanne Hand and Nancy Simmons for their careful reviews of our paper. Their suggestions greatly improved the manuscript. Early aspects of this study were funded in part by a National Science Foundation grant (# DEB 99-81512) to N. J. Czaplewski and G. S. Morgan. We dedicate this paper to our good friend and colleague Dr. Gregg F. Gunnell, whose untimely passing in 2017 saddened all who knew him. We honor his memory with the description of a new genus and species of emballonurid bat, Karstopter- yx gunnelli. Gregg was the Director of the Division of Fossil Primates at the Duke Lemur Center. He made numerous contributions to chiropteran pale- ontology, in particular from Africa, including late Eocene and early Oligocene bats from the Fayum Depression in Egypt, Pliocene bats from Kenya and Morocco, and early Pleistocene bats from Old- uvai Gorge in Tanzania. We are especially grateful for his meticulous editing of our two chapters for the 2012 book on the Evolutionary History of Bats. We greatly miss Gregg’s legendary knowledge of fossil mammals, and above all his friendship. LITERATURE CITED Albright, L. B., III. 1998. The Arikareean land mam- mal age in Texas and Florida: southern exten- sion of Great Plains faunas and Gulf Coastal Plain endemism. Pp. 167–183 in D. O. Terry, Jr., H. E. LaGarry, and R. M. Hunt, Jr. (eds.), Depositional Environments, Lithostratigra- phy, and Biostratigraphy of the White River and Arikaree Groups (Late Eocene to Early Miocene, North America). Geological Society of America, Special Paper 325. Albright, L. B., III, M. O. Woodburne, T. Fremd, C. C. Swisher, B. J. MacFadden, and G. R. Scott. 2008. Revised chronostratigraphy and biostra- tigraphy of the John Day Formation (Turtle Cove and Kimberly members), Oregon, with implications for updated calibration of the Ari- kareean North American Land Mammal Age. Journal of Geology 116:211–237. Allen, G. M. 1932. A Pleistocene bat from Florida. Journal of Mammalogy 13:256–259. Alvarez-Castañeda, S. T., and J. L. Patton, eds. 1999. Mamíferos del Noroeste de México, vol. I. Centro de Investigaciones Biológicas de Noroeste, La Paz, Baja California Sur, 583 pp. Antoine, P.-O., M. A. Abello, S. Adnet, A. J. Al- tamirano Sierra, P. Baby, G. Billet, M. Boivin, Y. Calderón, A. Candela, J. Chabain, F. Cor- fu, D. A. Croft, M. Ganerod, C. Jaramillo, S. Klaus, L. Marivaux, R. E. Navarrete, M. J. Or- liac, F. Parra, M. E. Pérez, F. Pujos, J.-C. Rage, A. Ravel, C. Robinet, M. Roddaz, J. V. Tejada- Lara, J. Vélez-Juarbe, F. P. Wesselingh, and R. Salas-Gismondi. 2016. A 60-million-year Ce- nozoic history of western Amazonian ecosys- tems in Contamana, eastern Peru. Gondwana Research 31:30–59. Antoine P.-O., L. Marivaux, D. A. Croft, D. A., G. Billet, M. Ganerød, C. Jaramillo, T. Martin, M. J. Orliac, J. Tejada, F. Duranthon, G. Fanjat, S. Rousse, and R. Salas-Gismondi. 2012. Mid- dle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography. Proceedings of the Royal Society B: Biology 279:1319–1326. Arnal, M., A. G. Kramarz, M. G. Vucetich, C. D. Frailey, and K. E. Campbell, Jr. 2019. New Paleogene caviomorphs (Rodentia, Hystri- 224 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) cognathi) from Santa Rosa, Peru: systemat- ics, biochronology, biogeography and early evolutionary trends. Papers in Palaeontology 2019:1-24. DOI: 10.1002/spp2.1264. Arroyo-Cabrales, J. 1992. Sinopsis de los murcié- lagos fosiles de México. Revista de la Socie- dad Mexicana de Paleontología 5(1):1–14. Arroyo-Cabrales, J., and O. J. Polaco. 2003. Caves and the Pleistocene vertebrate paleontology of Mexico. Pp. 273-291 in B. W. Schubert, J. I. Mead, and R. W. Graham (eds.), Ice Age Cave Faunas of North America, Indiana University Press, Bloomington. Assemat, A., M. Boivin, L. Marivaux, F. Pujos, A. Benites-Palomino, R. Salas-Gismondi, J. V. Tejada-Lara, R. M. Varas-Malca, F. R. Ne- gri, A. M. Ribeiro, and P.-O. Antoine. 2019. Restes inédits de rongeurs caviomorphes du Paléogène de la region de Juanjui (Amazonie péruvienne): systématique, implications mac- ro-évolutives et biostratigraphiques. Geodi- versitas 41:699-730. https://doi.org/10.5252/ geodiversitas2019v41a20. Bambini, L., A. Blyth, T. Bradford, R. Bristol, S. Burthe, L. Craig, N. Downs, S. Laying, L. Marshall-Ball, D. McGowan, T. Vel, and P. Racey. 2006. Another Seychelles endemic close to extinction: the emballonurid bat Co- leura seychellensis. Oryx 40:1–9. Barbour, T. 1936. Eumops in Florida. Journal of Mammalogy 17:414. Barghoorn, S. F. 1977. New material of Vespertil- iavus Schlosser (Mammalia, Chiroptera) and suggested relationships of emballonurid bats based on cranial morphology. American Mu- seum Novitates 2618:1–29. Bartlett, S. N., M. M. McDonough, and L. K. Am- merman. 2013. Molecular systematics of bon- neted bats (Molossidae: Eumops) based on mitochondrial and nuclear DNA sequences. Journal of Mammalogy 94:867–880. Berry E. W. 1916. The physical conditions and age indicated by the flora of the Alum Bluff For- mation. U. S. Geological Survey Professional Paper, Report: P 0098-E, 41–59. Bloch, J. I., E. D. Woodruff, A. R. Wood, A. F. Rincon, A. R. Harrington, G. S. Morgan, D. A. Foster, C. Montes, C. A. Jaramillo, N. A. Jud, D. S. Jones, and B. J. MacFadden. 2016. First North American fossil monkey and early Mio- cene tropical biotic interchange: Nature 533: 243–246. Bond M., M. F. Tejedor, K. E. Campbell, L. Chor- nogubsky, N. Novo, and F. Goin. 2015. Eo- cene primates of South America and the Af- rican origins of New World monkeys. Nature 520:538–541. Brooke, A. P. 1990. Tent construction and social or- ganization in Vampyressa nymphaea (Chirop- tera: Phyllostomidae) in Costa Rica. Journal of Tropical Ecology 3:171–175. Bryant J. D., B. J. MacFadden, and P. A. Muel- ler. 1992. Improved chronologic resolution of the Hawthorn and the Alum Bluff Groups in northern Florida: Implications for Miocene chronostratigraphy. Geological Society of America Bulletin, 104:208–218. Butler, P.M., and A. T. Hopwood. 1957. Insectivora and Chiroptera from the Miocene rocks of Ke- nya Colony. Fossil Mammals of Africa. British Museum (Natural History) 13:1–35. Ceballos, G., and R. Medellín., 1988. Diclidurus albus. Mammalian Species 316:1–4. Ceballos, G., J. Arroyo-Cabrales, and D. Vasquez. 2014. Order Chiroptera. Pp. 667–838 in G. Ceballos (ed.), Mammals of Mexico. Johns Hopkins University Press, Baltimore. Cirranello, A., N. B. Simmons, S. Solari, and R. J. Baker. 2016. Morphological diagnoses of higher-level phyllostomid taxa (Chirop- tera: Phyllostomidae). Acta Chiropterologica 18:39–71. Coombs, M. C., R. M. Hunt, Jr., E. Stepleton, L. B. Albright III, and T. J. Fremd. 2001. Stratigra- phy, chronology, biogeography, and taxonomy of early Miocene small chalicotheres in North America. Journal of Vertebrate Paleontology 21:607–620. Czaplewski, N. J. 1993. Late Tertiary bats (Mamma- lia, Chiroptera) from the southwestern United States. Southwestern Naturalist 38:111–118. Czaplewski, N. J. 1996a. Opossums (Didelphidae) https://doi.org/10.5252/geodiversitas2019v41a20 https://doi.org/10.5252/geodiversitas2019v41a20 MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 225 and bats (Noctilionidae and Molossidae) from the late Miocene of the Amazon Basin. Journal of Mammalogy 77:84–94. Czaplewski, N. J. 1996b. Thyroptera robusta Czaplewski, 1996 is a junior synonym of Thy- roptera lavali Pine, 1993 (Mammalia: Chirop- tera). Mammalia 60:153–156. Czaplewski, N. J. 1997. Chiroptera. Pp. 410-431 in R. F. Kay, R. H. Madden, R. L Cifelli, and J. J. Flynn (eds.), Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Ven- ta, Colombia. Smithsonian Institution Press, Washington, DC. Czaplewski, N. J. 2005. A review of the pre-Pleis- tocene fossil record of bats (Chiroptera) in South America. II Congresso Latino-America- no de Paleontologia de Vertebrados, Boletim de Resumos, Rio de Janeiro, Museu Nacional, Serie Livros 12: 87–89. Czaplewski, N. J. 2010. Colhuehuapian bats (Mam- malia: Chiroptera) from the Gran Barranca, Chubut province, Argentina. Pp. 240–252 in R. H. Madden, A. A. Carlini, M. G. Vucetich and R. F. Kay (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change through the Middle Cenozoic of Pa- tagonia. Cambridge University Press, Cam- bridge. Czaplewski, N. J., and K. E. Campbell, Jr. 2004. A possible bat (Mammalia: Chiroptera) from the ?Eocene of Amazonian Peru. Natural History Museum of Los Angeles County, Science Se- ries 40:141–144. Czaplewski, N. J., and K. E. Campbell, Jr. 2017. Late Miocene bats from the Juruá River, state of Acre, Brazil, with a description of a new genus of Thyropteridae (Chiroptera, Mamma- lia). Natural History Museum of Los Angeles County, Contributions in Science 525: 55–60. Czaplewski, N. J., and C. Cartelle. 1998. Pleisto- cene bats from cave deposits in Bahia, Brazil. Journal of Mammalogy 79:784–803. Czaplewski, N. J., and G. S. Morgan. 2000. A new vespertilionid bat (Mammalia: Chiroptera) from the early Miocene (Hemingfordian) of Florida, USA. Journal of Vertebrate Paleontol- ogy 20:736–742. Czaplewski, N. J., and G. S. Morgan. 2012. New basal noctilionoid bats (Mammalia: Chirop- tera) from the Oligocene of subtropical North America. Pp. 162–209 in G. F. Gunnell and N. B. Simmons (eds.), Evolutionary History of Bats: Fossils, Molecules, and Morphology. Cambridge University Press, Cambridge. Czaplewski, N. J., and G. S. Morgan. 2015. A late- surviving apatemyid (Mammalia: Apatothe- ria) from the latest Oligocene of Florida, USA. PeerJ 3:e1509; DOI 10.7717/peerj.1509 (22 pages). Czaplewski, N. J., G. S. Morgan, R. J. Emry, P. M. Gignac, and H. D. O’Brien. 2022. Three new early middle Eocene bats (Mammalia: Chirop- tera) from Elderberry Canyon, Nevada, USA. Smithsonian Contributions to Paleobiology 106:1–26. Czaplewski, N. J., G. S. Morgan, and R. A. Mar- tin. 2018. New specimens of Pliocene bats (Mammalia, Chiroptera, Vespertilionidae and Molossidae) from the Meade Basin, Kansas, U.S.A., Journal of Vertebrate Paleontology DOI: 10.1080/02724634.2018.1430034 (14 pages). Czaplewski, N. J., G. S. Morgan, and S. A. McLeod. 2008. Chiroptera. Chapter 12. Pp. 174–197 in C. M. Janis, G. F. Gunnell, and M. D. Uhen (eds.), Evolution of Tertiary Mammals of North America, Vol. 2: Small Mammals, Xen- arthrans, and Marine Mammals. Cambridge University Press, Cambridge. Czaplewski, N. J., G. S. Morgan, and T. Naeher. 2003a. Molossid bats from the Late Tertiary of Florida with a review of the Tertiary Molos- sidae of North America. Acta Chiropterologica 5:61–74. Czaplewski, N. J., and A. D. Rincon. 2020. A gi- ant vampire bat (Phyllostomidae, Desmodon- tinae) from the Pliocene-Pleistocene El Breal de Orocual asphaltic deposits (tar pits), Ven- ezuela. Historical Biology doi.org/10.1080/08 912963.2020.1800684. Czaplewski, N. J., M. Takai, T. M. Naeher, N. Shigehara, and T. Setoguchi. 2003b. Addi- 226 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) tional bats from the middle Miocene La Venta Fauna of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas, y Naturales 27 (103):263–282. Dalquest, W. W., and E. Roth. 1970. Late Pleis- tocene mammals from a cave in Tamaulipas, Mexico. Southwestern Naturalist 15:217–230. Dawson, M. R. 1976. Paleogene terrestrial verte- brates: northernmost occurrence, Ellesmere Island, Canada. Science 192:781–782. Dawson, M. R. 1991. Early Eocene rodents (Mam- malia) from the Eureka Sound Group of Elles- mere Island, Canada. Canadian Journal of Earth Sciences 28: 364–371. Eisenberg, J. F. 1989. Mammals of the Neotropics, Volume 1: The Northern Neotropics: Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana. University of Chicago Press, Chicago, 550 p. Eberle, J. J., and M. C. McKenna. 2002. Early Eo- cene Leptictida, Pantolesta, Creodonta, Car- nivora, and Mesonychidae (Mammalia) from the Eureka Sound Group, Ellesmere Island, Nunavut. Canadian Journal of Earth Sciences 39: 899–910. Emry, R. J., P. R. Bjork, and L. S. Russell. 1987. The Chadronian, Orellan, and Whitneyan North American Land Mammal Ages. Pp. 118–152 in M. O. Woodburne (ed.), Cenozoic Mammals of North America: Geochronology and Biostratigraphy. University of California Press, Berkeley. Eshelman, R. E., and G. S. Morgan. 1985. Tobagan Recent mammals, fossil vertebrates, and their zoogeographical implications. Natonal Geo- graphic Society, Research Reports 21:137– 143. Estes, R., and J. H. Hutchison. 1980. Eocene lower vertebrates from Ellesmere Island, Canadian Arctic Archipelago. Palaeogeography, Palaeo- climatology, Palaeoecology 30:325–347. Florea, L. J. 2008. Geology and hydrology of karst in west-central and north-central Florida. Pp. 225–239 in L. J. Florea (ed.), Caves and Karst of Florida. A Guidebook for the 2008 National Speleological Society National Convention. National Speleological Society, Huntsville, Alabama. Flynn, J. J., and A. R. Wyss. 1998. Recent advances in South American mammalian paleontology. Trends in Ecology and Evolution 13:449–454. Forstén, A. 1975. The fossil horses of the Texas Gulf Coastal Plain: a revision. Texas Memo- rial Museum, Pearce-Sellards Series 22:1–86. Frailey, C. D. 1978. An early Miocene (Arika- reean) fauna from northcentral Florida (the SB-1A Local Fauna). Occasional Papers, Mu- seum of Natural History, University of Kansas 75:1–20. Frailey, C. D. 1979. The large mammals of the Buda Local Fauna (Arikareean: Alachua County, Florida). Bulletin of the Florida State Museum, Biological Sciences 24:123–173. Frank, P. A. 1997a. First record of Artibeus jamai- censis Leach (1821) from the United States. Florida Scientist 60:37–39. Frank, P. A. 1997b. First record of Molossus molossus tropidorhynchus Gray (1839) from the United States. Journal of Mammalogy 78:103–105. Gaudioso, P. J., R. M. Barquez, and M. M. Díaz. 2020. Esqueleto Postcranial de Chiroptera: Atlas para la Identificación de las Partes. Pub- licaciones Especiales no. 3, Programa de In- vestigaciones de Biodiversidad Argentina, Tucuman. Genoways, H. H., C. J. Phillips, and R. J. Baker. 1998. Bats of the Antillean island of Grenada: a new zoogeographic perspective. Museum of Texas Tech University, Occasional Papers 177:1–28. Giannini, N. P., and N. B. Simmons. 2007. Element homology and the evolution of dental formu- lae in megachiropteran bats (Mammalia: Chi- roptera: Pteropodidae). American Museum Novitates 3559:1–27. Giannini, N. P., J. R. Wible, and N. B. Simmons. 2006. On the cranial osteology of Chiroptera. I. Pteropus (Megachiroptera: Pteropodidae). Bulletin of the American Museum of Natural History 295:1–134. Goodman, S. M., S. J. Puechmaille, N. Friedli- https://en.wikipedia.org/wiki/Canadian_Journal_of_Earth_Sciences https://en.wikipedia.org/wiki/Canadian_Journal_of_Earth_Sciences MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 227 Weyeneth, J. Gerlach, M. Ruedi, M. C. Schoe- man, W. T. Stanley, and E. C. Teeling. 2012. Phylogeny of the Emballonurini (Emballon- uridae) with descriptions of a new genus and species from Madagascar. Journal of Mam- malogy 93:1440–1455. Goodwin, G. G., and A. H. Greenhall. 1961. A re- view of the bats of Trinidad and Tobago. Bul- letin of the American Museum of Natural His- tory 122:187–302. Griffiths, T. A., and A. L. Smith. 1991. Systematics of emballonuroid bats (Chiroptera: Emballon- uridae and Rhinopomatidae), based on hyoid morphology. Bulletin of the American Muse- um of Natural History 206:62–83. Gunnell, G. F. 2010. Chiroptera. Pp. 581–597 in L. Werdelin and W. J. Sanders (eds.), Cenozoic Mammals of Africa. University of California Press, Berkeley. Gunnell G. F, and F. K. Manthi. 2020. Pliocene bats (Chiroptera) from Kanapoi, Turkana Ba- sin, Kenya. Journal of Human Evolution 140: 1–18. Gunnell, G. F., and N. B. Simmons. 2005. Fos- sil evidence and the origin of bats. Journal of Mammalian Evolution 12:209–246. Gunnell, G. F, N. B. Simmons, and E. R. Seiffert. 2014. Myzopodidae (Chiroptera) from the Late Paleogene of Egypt: emended family di- agnosis and biogeographic origins of Noctil- ionoidea. PLoS ONE 9(2): e86712. Gunnell G. F., E. L. Simons, and E. R. Seiffert. 2008. New bats (Mammalia: Chiroptera) from the late Eocene and early Oligocene, Fayum Depression, Egypt. Journal of Vertebrate Pale- ontology 28:1–11. Gunnell, G. F., S. R. Worsham, E. R. Seiffert, and E. L. Simons. 2009. Vampyravus orientalis Schlosser (Chiroptera) from the Early Oligo- cene (Rupelian), Fayum, Egypt—body mass, humeral morphology and affinities. Acta Chi- ropterologica 11:271–278. Hall, E. R. 1981. The Mammals of North Ameri- ca, 2nd ed., vol. 1. John Wiley and Sons, New York, 600 p. Hand, S. J., M. Archer, and H. Godthelp. 2005. Australian Oligo-Miocene mystacinids (Mi- crochiroptera): upper dentition, new taxa and divergence of New Zealand species. Geobios- Lyon 38:339–352. Hand, S. J., B. Sigé, M. Archer, G. F. Gunnell, and N. B. Simmons. 2015. A new early Eo- cene (Ypresian) bat from Pourcy, Paris Basin, France, with comments on patterns of diversi- ty in the earliest chiropterans. Journal of Mam- malian Evolution 22:343–354. Hayes, F. G. 2000. The Brooksville 2 Local Fau- na (Arikareean, latest Oligocene): Hernando County, Florida. Bulletin of the Florida Mu- seum of Natural History 43:1–47. Hayes, F. G. 2005. Arikareean (Oligocene-Mio- cene) Herpetotherium (Marsupialia, Didel- phidae) from Nebraska and Florida. Bulletin of the Florida Museum of Natural History 45:335–353. Head, J. J., A. F. Rincon, C. Suarez, C. Montes, and C. Jaramillo. 2012. Fossil evidence for earliest Neogene American faunal interchange: Boa (Serpentes, Boinae) from the early Miocene of Panama. Journal of Vertebrate Paleontology 32:1328–1334. Henson, O. W., Jr. 1970. The ear and audition. Pp. 181–263 in W. A. Wimsatt (ed.), Biology of Bats, Volume II. Academic Press, New York. Hill, J. E., and J. D. Smith, 1984. Bats: A Natural History. British Museum (Natural History), London, 243 p. Holman, J. A. 1999. Early Oligocene (Whitneyan) snakes from Florida (USA), the second old- est colubrid snakes in North America. Acta Zoologica Cracoviensia 42:447–454. Holman, J. A., and D. L. Harrison. 2000. Early Oligocene (Whitneyan) snakes from Florida (USA), a unique booid. Acta Zoologica Craco- viensia 43:127–134. Holman, J. A., and D. L. Harrison. 2001. Early Oligocene (Whitneyan) snakes from Florida (USA): remaining boids, indeterminate colu- broids, summary and discussion of the I-75 Local Fauna snakes. Acta Zoologica Cracovi- ensia 44:25–36. Hooker, J. J. 1996. A primitive emballonurid bat 228 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) (Chiroptera, Mammalia) from the earliest Eo- cene of England. Palaeovertebrata 25:287– 300. Horáček, I. 2001. On the early history of vesper- tilionid bats in Europe: the lower Miocene re- cord from the Bohemian Massif. Lynx (Praha), n.s. 32, 123–154. Hulbert, R. C., Jr. 1984. Paleoecology and popula- tion dynamics of the early Miocene (Heming- fordian) horse Parahippus leonensis from the Thomas Farm Site, Florida. Journal of Verte- brate Paleontology 4:547–558. Hulbert, R. C., Jr., and B. J. MacFadden. 1991. Morphological transformation and cladogen- esis at the base of the adaptive radiation of Miocene horses. American Museum Novitates 3000:1–61. Jaramillo, C., E. Moreno, V. Ramirez, S. da Silva, A. Barrera, B. Adhara, S. Moron, F. Herrera, J. Escobar, J. R. Koll, S. Manchester, and N. Hoyos. 2014. Palynological record of the last 20 million years in Panama. Pp. 134–253 in W. D. Stevens, O. M. Montiel, P. H. Raven (eds.), Paleobotany and Biogeography: A Festschrift for Alan Graham in His 80th Year. Missouri Botanical Garden Press, St. Louis, Mono- graphs in Systematic Botany 128. Jarzen, D. M., S. L. Corbett, and S. R. Manches- ter. 2010. Palynology and paleoecology of the middle Miocene Alum Bluff flora, Liberty County, Florida, USA. Palynology 34:261– 286. Jones, D. S., P. A. Mueller, D. A. Hodell, and L. A. Stanley. 1993. 87Sr/86Sr geochronology of Oli- gocene and Miocene marine strata in Florida. Florida Geological Survey, Special Publica- tion 37:55–66. Keith, D. A., and 40 other authors. 2022. A func- tion-based typology for Earth’s ecosystems. Nature 610:513–518. King, T. R., T. J. Myers, K. N. Armstrong, M. Ar- cher, and S. J. Hand. 2020. Sheath-tailed bats (Chiroptera: Emballonuridae) from the early Pleistocene Rackham’s Roost Site, Riversleigh World Heritage Area, and the distribution of northern Australian emballonurid species. PeerJ 9: e10857 http://dx.doi.org/10.7717/ peerj.10857. Koopman, K. F. 1970. Zoogeography of bats. Pp. 29–50 in B. H. Slaughter and D. W. Walton (eds.), About Bats. Southern Methodist Uni- versity Press, Dallas, Texas. Koopman, K. F. 1971. The systematic and histori- cal status of the Florida Eumops (Chiroptera, Molossidae). American Museum Novitates 2478:1–6. Koopman, K. F. 1976. Zoogeography. Pp. 39–47 in R. J. Baker, J. K. Jones, Jr., and D. C. Cart- er (eds.), Biology of Bats of the New World Family Phyllostomatidae, Part 1. Special Pub- lications, The Museum Texas Tech University, Number 10. Koopman, K. F. 1982. Biogeography of the bats of South America. Pp. 273–302 in M. A. Mares and H. H. Genoways (eds.), Mammalian Bi- ology in South America. Pymatuning Labora- tory of Ecology, University of Pittsburgh, Spe- cial Publication Series, Vol. 6. Koopman, K. F. 1989. A review and analysis of the bats of the West Indies. Pp. 635–644 in C. A. Woods (ed.), Biogeography of the West In- dies: Past, Present, and Future. Sandhill Crane Press, Gainesville, Florida. Lane, E. 1986. Karst in Florida. Florida Geological Survey, Special Publication 29:1–100. Lawrence, B. 1943. Miocene bat remains from Florida, with notes on the generic characters of the humerus of bats. Journal of Mammalogy 24:356–369. Legendre, S. 1980. Un chiroptère emballonuridé dans le Néogène d’Europe Occidentale; con- sidérations paléobiogéographiques. Geobios, no. 13, fasc. 6:839–847. Legendre, S. 1985. Molossidés (Mammalia, Chi- roptera) cénozoïques de l’Ancien et du Nou- veau Monde; statut systématique; intégration phylogénique de données. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 170:205–227. Lessa, G., C. Cartelle, and M. P. de A. Fracasso. 2005. The bat (Mammalia, Chiroptera) fossils of Brazil. II Congresso Latino-Americano de https://gcc02.safelinks.protection.outlook.com/?url=https%3A%2F%2Furldefense.proofpoint.com%2Fv2%2Furl%3Fu%3Dhttp-3A__dx.doi.org_10.7717_peerj.10857%26d%3DDwMF-g%26c%3DsJ6xIWYx-zLMB3EPkvcnVg%26r%3DfDN_lbWzanKmXSwJY9r9Ril88TLtELA8DNk1aK09qDI%26m%3DlspG2SzVdX3RqKF4MNMChSCQF2F-E76hoevGq9f2kOXSy1UvvDGAHsWww7MZDa2s%26s%3DZJixrnFUTrWanUzdgNWS9a_eD-G-llsako0BbQbORpU%26e%3D&data=04%7C01%7Cgary.morgan1%40state.nm.us%7Cecc0da52947444c7042908d9c3dacdf1%7C04aa6bf4d436426fbfa404b7a70e60ff%7C0%7C0%7C637756169323937107%7CINCLUDEPICTURE https://gcc02.safelinks.protection.outlook.com/?url=https%3A%2F%2Furldefense.proofpoint.com%2Fv2%2Furl%3Fu%3Dhttp-3A__dx.doi.org_10.7717_peerj.10857%26d%3DDwMF-g%26c%3DsJ6xIWYx-zLMB3EPkvcnVg%26r%3DfDN_lbWzanKmXSwJY9r9Ril88TLtELA8DNk1aK09qDI%26m%3DlspG2SzVdX3RqKF4MNMChSCQF2F-E76hoevGq9f2kOXSy1UvvDGAHsWww7MZDa2s%26s%3DZJixrnFUTrWanUzdgNWS9a_eD-G-llsako0BbQbORpU%26e%3D&data=04%7C01%7Cgary.morgan1%40state.nm.us%7Cecc0da52947444c7042908d9c3dacdf1%7C04aa6bf4d436426fbfa404b7a70e60ff%7C0%7C0%7C637756169323937107%7CINCLUDEPICTURE MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 229 Paleontologia de Vertebrados, Boletim de Re- sumos, Rio de Janeiro, Museu Nacional, Serie Livros 12:153–155. Lillegraven, J. A., M. C. McKenna, and L. Krishtal- ka. 1981. Evolutionary relationships of middle Eocene and younger species of Centetodon (Mammalia, Insectivora, Geolabididae) with a description of the dentition of Ankylodon (Adapisoricidae). University of Wyoming Publications 45:1–115. Lim, B. K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (Tribe Dicli- durini) in South America. Molecular Phyloge- netics and Evolution 45:777–791. Lim, B. K. 2008. Historical biogeography of New World emballonurid bats (tribe Diclidurini): taxon pulse diversification. Journal of Bioge- ography 35:1385–1401. Lim, B. K. 2009. Review of the origins and bioge- ography of bats in South America. Chiroptera Neotropical 15:391–410. Lim, B. K. 2010. Adaptive radiation of Neotropi- cal emballonurid bats: molecular phylogenet- ics and evolutionary patterns in behavior and morphology. Pp. 283–299 in P. Pontarotti (ed.), Evolutionary Biology – Concepts, Mo- lecular and Morphological Evolution. Spring- er-Verlag, Berlin. Lim, B. K., and J. M. Dunlop. 2008. Evolution- ary patterns of morphology and behavior as inferred from a molecular phylogeny of New World emballonurid bats (Tribe Diclidurini). Journal of Mammalian Evolution 15:79–121. Lim, B. K., M. D. Engstrom, J. W. Bickham, and J. C. Patton. 2008. Molecular phylogeny of New World sheath-tailed bats (Emballonuridae: Di- clidurini) based on loci from the four genetic transmission systems in mammals. Biological Journal of the Linnean Society 93:189–209. López-Aguirre, C., S. J. Hand, S. W. Laffan, and M. Archer. 2019. Zoogeographical regions and geospatial patterns of phylogenetic diversity and endemism of New World bats. Ecography 42:1188–1199. López-Forment, W., and G. Tellez-Giron. 2014. Peters’ Sac-winged Bat. Pp. 669–670 in G. Ceballos (ed.), Mammals of Mexico. Johns Hopkins University Press, Baltimore. Lott, T. A., S. R. Manchester, and S. L. Corbett. 2019. The Miocene flora of Alum Bluff, Lib- erty County, Florida. Acta Palaeobotanica 59:75–129. Macdonald, J.R. 1963. The Miocene faunas from the Wounded Knee area of western South Da- kota. Bul letin of the American Museum of Natural History 125(3):141–238. Macdonald, J.R. 1970. Review of the Miocene Wounded Knee faunas of southwestern South Dakota. Los Angeles County Museum of Nat- ural History Bulle tin 8:1–82. MacFadden, B. J., J. I. Bloch, H. Evans, D. A. Foster, G. S. Morgan, A. F. Rincon, and A. R. Wood. 2014. Temporal calibration and biochronology of the Centenario Fauna, ear- ly Miocene of Panama. Journal of Geology 122:113–135. MacFadden, B. J., and G. S. Morgan. 2003. New oreodont (Mammalia, Artiodactyla) from the late Oligocene (early Arikareean) of Florida. Bulletin of the American Museum of Natural History 279:368–396, Maitre, E. 2014. Western European middle Eo- cene to early Oligocene Chiroptera: system- atics, phylogeny and paleoecology based on new material from the Quercy (France). Swiss Journal of Palaeontology 133:141–242. Marivaux, L., S. Adnet, A. J. Altamirano-Sierra, F. Pujos, A. Ramdarshan, R. Salas-Gismondi, J. V. Tejada-Lara, and P. O. Antoine. 2016. Den- tal remains of cebid platyrrhines from the ear- liest late Miocene of western Amazonia, Peru: macroevolutionary implications on the extant capuchin and marmoset lineages. American Journal of Physical Anthropology 161:478– 493. Marks, C. S., and G. E. Marks. 2006. Bats of Flor- ida. University Press of Florida, Gainesville, 176 p. McDonough, M. M., L. K. Ammerman, R. M. Timm, H. H. Genoways, P. A. Larsen, and R. J. Baker. 2008. Speciation within bonneted bats (genus Eumops): the complexity of morpho- 230 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) logical, mitochondrial, and nuclear data sets in systematics. Journal of Mammalogy 89:1306– 1315. McKenna, M. C. 1975. Fossil mammals and early Eocene North Atlantic land continuity. Annals of the Missouri Botanical Garden 62:335–353. McKenna, M. C., P. Robinson, and D. W. Tay- lor. 1962. Notes on Eocene Mammalia and Mollusca from Tabernacle Butte, Wyoming. American Museum Novitates 2102:1–33. Mead, J. I. 2013. Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene, North America, and a fossil history overview. Geo- bios 46:225–231. Medellín, R. A., H. T. Arita, and H. Ó. Sánchez 2008. Identificación de Los Murciélagos de México: Clave de Campo. Segunda edición. Instituto de Ecología, Universidad Nacional Autónoma de Mexico. 80 p. Mezzalira, S. 1966. Os fósseis do Estado de São Paulo. São Paulo, Instituto Geográfico do Ge- ológico, Boletim 45 1–132. Miller, G. S., Jr. 1907. The families and genera of bats. U. S. National Museum Bulletin 57:1– 282. Montes, C., A. Cardona, R. McFadden, S. E. Mo- ron, C. A. Silva, S. Restrepo-Moreno, D. A. Ramirez, N. Hoyos, J. Wilson, D. Farris, G. A. Bayona, C. A. Jaramillo, V. Valencia, J. Bryan, and J. A. Flores. 2012. Evidence for middle Eocene and younger emergence in Central Panama: implications for Isthmus clo- sure. Geological Society of America Bulletin 124:780–799. Morgan, G. S. 1991. Neotropical Chiroptera from the Pliocene and Pleistocene of Florida. Bul- letin of the American Museum of Natural His- tory 206:176–213. Morgan, G. S. 1993. Mammalian biochronology and marine-nonmarine correlations in the Neogene of Florida. Florida Geological Sur- vey, Special Publication 37:55–66. Morgan, G. S. 2001. Patterns of extinction in West Indian bats. Pp. 369–407 in C. A. Woods and F. E. Sergile (eds.), Biogeography of the West Indies: Patterns and Perspectives, Second ed., CRC Press, Boca Raton, Florida. Morgan, G. S. 2002. Late Rancholabrean mammals from southernmost Florida, and the Neotropi- cal influence in Florida Pleistocene Faunas. Smithsonian Contributions to Paleobiology 93:15–38. Morgan, G. S. 2008. Vertebrate fauna and geochro- nology of the Great American Biotic Inter- change in North America. New Mexico Mu- seum of Natural History and Science Bulletin 44:93–140. Morgan, G. S., and N. J. Czaplewski. 2003. A new bat (Chiroptera: Natalidae) from the early Miocene of Florida, with comments on natalid phylogeny. Journal of Mammalogy 84:729– 752. Morgan, G. S., and N. J. Czaplewski. 2012. Evolu- tionary history of the Neotropical Chiroptera: the fossil record. Pp. 105–161 in G. F. Gunnell and N. B. Simmons (eds.), Evolutionary His- tory of Bats: Fossils, Molecules, and Morphol- ogy. Cambridge University Press, Cambridge. Morgan, G. S., N. J. Czaplewski, A. F. Rincon, A. R. Wood, and B. J. MacFadden. 2013. An ear- ly Miocene bat (Chiroptera: Phyllostomidae) from Panama and mid Cenozoic chiropteran dispersals between the Americas. Journal of Vertebrate Paleontology, October 2013, Pro- gram and Abstracts, p. 180. Morgan, G. S., N. J. Czaplewski, and N. B. Sim- mons. 2019. A new mormoopid bat from the Oligocene (Whitneyan and Arikareean) of Florida, and phylogenetic relationships of the major clades of Mormoopidae (Mammalia: Chiroptera). Bulletin of the American Muse- um of Natural History 434:1–141. Morgan, G. S., and S. D. Emslie. 2010. Tropical and western influences in vertebrate faunas from the Pliocene and Pleistocene of Florida. Quaternary International 217:143–158. Morgan, G. S., and R. C. Hulbert, Jr. 2008. Ceno- zoic vertebrate fossils from paleokarst depos- its in Florida. Pp. 248–271 in L. J. Florea (ed.), Caves and Karst of Florida. A Guidebook for the 2008 National Speleological Society Na- tional Convention. National Speleological So- MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 231 ciety, Huntsville, Alabama. Morgan, G. S., O. J. Linares, and C. E. Ray. 1988. New species of fossil vampire bats (Mamma- lia: Chiroptera: Desmodontidae) from Florida and Venezuela. Proceedings of the Biological Society of Washington 101:912–928. Nkrumah, E. E., H. J. Baldwin, E. K. Badu, P. Anti, P. Vallo, S. Close, S., E. K. V. Kalko, S. K. Op- pong, and M. Tschapka. 2021. Diversity and conservation of cave-roosting bats in central Ghana. Tropical Conservation Science 14:1– 10. Novacek, M. J. 1985. Comparative morphology of the bat auditory region. Fortschritte der Zoolo- gie 30:149–151. Novacek, M. J. 1991. Aspects of morphology of the cochlea in microchiropteran bats: an inves- tigation of character transformation. Bulletin of the American Museum of Natural History 206:84–100. Nowak, R. M. 1994. Walker’s Bats of the World. The Johns Hopkins University Press, Balti- more, 287 p. O’Dea, A., and 35 other authors. 2016. Formation of the Isthmus of Panama. Science Advances 2: e1600883. O’Leary, M. A., J. I. Bloch, J. J. Flynn, T. J. Gaud- in, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z.-X. Luo, J. Meng, X. Ni, M. J. Novacek, F. A. Perini, Z. S. Ran- dall, G. W. Rougier, E. J. Sargis, M. T. Silcon, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible, and A. L. Cirranello. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339:662–667. Olson, D. M., et al., 2001. Terrestrial ecoregions of the world: a new map of life on Earth. BioSci- ence 51:933–938. Patton, T. H. 1969a. An Oligocene land vertebrate fauna from Florida. Journal of Paleontology 43:543–546. Patton, T. H. 1969b. Miocene and Pliocene artio- dactyls, Texas Gulf Coastal Plain. Florida State Museum Bulletin, Biological Sciences 14:115–226. Paula Couto, C. de. 1956. Une chauve-souris fos- sile des argiles feuilletées Pléistocènes de Tremembé, État de São Paulo (Brèsil). Actes IV Congrès International du Quaternaire, Ro- ma-Pise, Août-Septembre 1953, vol. 1, 343– 347. Paula Couto, C. de and S. Mezzalira. 1971. Nova conceituação geocronológica de Tremem- bé, Estado de São Paulo, Brasil. Anais da Academia Brasileira de Ciências, 43 (Sup- pl.):473–488. Pratt, A. E. 1989. Taphonomy of the microverte- brate fauna from the early Miocene Thomas Farm locality, Florida (U.S.A.). Palaeogeog- raphy, Palaeoclimatology, and Palaeoecology 76:125–151. Pratt, A. E. 1990. Taphonomy of the large verte- brate fauna from the Thomas Farm locality (Miocene, Hemingfordian), Gilchrist County, Florida. Bulletin of the Florida Museum of Natural History 35(2):35–130. Pratt, A. E., and G. S. Morgan. 1989. New Sci- uridae (Mammalia: Rodentia) from the early Miocene Thomas Farm Local Fauna, Florida. Journal of Vertebrate Paleontology 9:89–100. Prothero, D. R., and R. J. Emry. 2004. The Chadro- nian, Orellan, and Whitneyan North American Land Mammal Ages. Pp. 156–168 in M. O. Woodburne (ed.), Late Cretaceous and Ceno- zoic Land Mammals of North America: Bio- stratigraphy and Geochronology. Columbia University Press, New York. Prothero, D. R., and N. Shubin. 1989. The evolu- tion of Oligocene horses. Pp. 142-175 in D. R. Prothero and R. M. Schoch (eds.), The Evolu- tion of Perissodactyls. Oxford Monographs in Geology and Geophysics 15, Oxford Univer- sity Press, Oxford. Ravel, A., M. Adaci, M. Bensalah, A.-L. Charru- ault, E. M. Essid, H. K. Ammar, W. Marzou- gui, M. Mahboubi, F. Mebrouk, G. Merzeraud, M. Vianey-Liaud, R. Tabuce, and L. Marivaux. 2016. Origine et radiation initiale des chauve- souris modernes: nouvelles découvertes dans l’Éocène d’Afrique du Nord. Geodiversitas 38:355–434. 232 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) Rich, T. H. V., and T. H. Patton. 1975. First re- cord of a fossil hedgehog from Florida (Eri- naceidae: Mammalia). Journal of Mammalogy 56:692–695. Robbins, L. W., and V. M. Sarich. 1988. Evolution- ary relationships in the Family Emballonuridae (Chiroptera). Journal of Mammalogy 69:1–13. Rodríguez-Herrera, B., R. A. Medellín, and R. M. Timm. 2007. Murciélagos neotropicales que acampan en hojas: Guía de Campo / Neotropi- cal tent-roosting bats: Field Guide. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. Rosina, V. V., and M. Pickford. 2019. Preliminary overview of the fossil record of bats (Chi- roptera, Mammalia) from the Miocene sites of Otavi Mountainland (northern Namibia). Communications of the Geological Survey of Namibia 21:48–58. Rosina, V. V., and M. Pickford. 2020. Miocene emballonurids (Chiroptera, Mammalia) from Berg Aukas I of Namibia (Africa) and their odontological features. Communications of the Geological Survey of Namibia 22:47–64. Rosina, V. V., and M. Pickford. 2021. The new small emballonurid (Emballonuridae, Chi- roptera, Mammalia) from the Miocene of Africa: its phylogenetic and palaeogeograph- ic implications. Historical Biology, DOI: 10.1080/08912963.2021.1973451. Ruedi, M., N. Friedli-Weyeneth, E. C. Teeling, S. J. Puechmaille, and S. M. Goodman. 2012. Bio- geography of Old World emballonurine bats (Chiroptera: Emballonuridae) inferred with mitochondrial and nuclear DNA. Molecular Phylogenetics and Evolution 64:204–211. Rydell, J., M. B. Fenton, E. Seamark, P. W. Webala, and T. C. Michaelsen. 2019. White and clear wings in bats. Abstracts of Papers presented at the 49th Annual Symposium of the North American Society for Bat Research, Kalama- zoo, Michigan, USA, 23-26 October 2019. Bat Research News 60(4):178. Savage, D. E. 1951. A Miocene phyllostomatid bat from Colombia, South America. University of California Publications, Bulletin of the De- partment of Geological Sciences 28:357–366. Savage, D. E., and D. E. Russell. 1983. Mammali- an Paleofaunas of the World. Addison-Wesley Publishing Company, London, 432 p. Seiffert, E. R., M. F. Tejedor, J. G. Fleagle, N. M. Novo, F. M. Cornejo, M. Bond, D. de Vries, and K. E. Campbell, Jr. 2020. A parapithecid stem anthropoid of African origin in the Paleo- gene of South America. Science 368:194–197. Doi: 10.1126/science.aba1135 Sigé, B. 1990. Nouveaux chiroptères de l’Oligocène moyen des phosphorites du Quercy, France. Compte Rendu de l’Académie des Sciences de Paris 310:1131–1137. Sigé, B., and S. Legendre. 1983. L’histoire des peuplements de chiroptères du bassin médi- terranéen: l’apport comparé des remplissages karstiques et des dépôts fluviolacustres. Mé- moires de Biospéologie 10:207–224. Sigé, B., H. Thomas, S. Sen, E. Gheerbrant, J. Rog- er, and Z. Al- Sulaimani. 1994. Les Chirop- tères de Taqah (Oligocène Inférieur, Sultanat D’Oman). Premier inventaire systématique. Münchner Geowissenschaftliche Abhandlun- gen 26:35–48. Simmons, N. B. 2005. Order Chiroptera. Pp. 312- 529 in D. E. Wilson and D. M. Reeder (eds.), Mammal Species of the World: A Taxonomic and Geographic Reference. Johns Hopkins University Press, Baltimore. Simmons, N. B., and A. L. Cirranello. 2019. Bat Species of the World: A taxonomic and geo- graphic database. Accessed on 19 August 2019. www.batnames.org. Simmons, N. B., and T. M. Conway. 2001. Phylo- genetic relationships of mormoopid bats (Chi- roptera: Mormoopidae) based on morphologi- cal data. Bulletin of the American Museum of Natural History 258:1-97. Simmons, N. B., and J. H. Geisler. 1998. Phylo- genetic relationships of Icaronycteris, Archae- onycteris, Hassianycteris, and Palaeochirop- teryx to extant bat lineages, with comments on the evolution of echolocation and foraging strategies in Microchiroptera. Bulletin of the American Museum of Natural History 235:1– http://www.batnames.org MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 233 182. Simmons, N. B., G. F. Gunnell, and N. J. Czaplews- ki. 2020. Fragments and gaps: the fossil record. Pp. 63–86 in T. H. Fleming, L. M. Dávalos, and M. A. R. Mello (eds.), Phyllostomid Bats: A Unique Mammalian Radiation. University of Chicago Press, Chicago. Simmons, N. B., and R. S. Voss. 1998. The mam- mals of Paracou, French Guiana: a Neotropical lowland rainforest fauna. Part 1. Bats. Bulletin of the American Museum of Natural History 237:1–219. Smith, J. D. 1972. Systematics of the chiropteran Family Mormoopidae. University of Kansas Museum of Natural History, Miscellaneous Publication 56: 1–132. Smith, T., J. Habersetzer, N. B. Simmons, and G. F. Gunnell. 2012. Systematics and paleobiogeog- raphy of early bats. Pp. 23–66 in G. F. Gunnell and N. B. Simmons (eds.), Evolutionary His- tory of Bats: Fossils, Molecules and Morphol- ogy. Cambridge University Press, Cambridge. Solari, S., C. G. Sotero-Caio, and R. J. Baker. 2019. Advances in systematics of bats: towards a consensus on species delimitation and classifi- cations through integrative taxonomy. Journal of Mammalogy 100:838–851. Staněk, V. J. 1933. K topografické a srovnávací anatomii sluchového orgánu našich chiropter. Prague: Nákladem České Akademie Věd a Umění, 67 pp. + 9 plates. Storch, G., B. Sigé, and J. Habersetzer. 2002. Ta- chypteron franzeni n. gen., n. sp., earliest emballonurid bat from the Middle Eocene of Messel (Mammalia, Chiroptera). Paläontolo- gische Zeitschrift 76:189–199. Storer, J. E. 1984. Mammals of the Swift Current Creek local fauna (Eocene: Uintan, Saskatch- ewan). Saskatchewan Culture and Recreation, Museum of Natural History, Natural History Contribution 7:1–158. Sulser, R. B., B. D. Patterson, D. J. Urban, A. I. Ne- ander, and Z.-X. Luo. 2022. Evolution of inner ear neuroanatomy of bats and implications for echolocation. Nature 602:449–454. Tedford, R. H. 1970. Principles and practices of mammalian geochronology in North America. North American Paleontological Convention Proceedings F (Chicago, 1969):666–703. Tedford, R. H., L. B. Albright, III, A. D. Barnosky, I. Ferrusquia-Villafranca, R. M. Hunt, Jr., J. E. Storer, C. C. Swisher, III, M. R. Voorhies, S. D. Webb, and D. P. Whistler. 2004. Mamma- lian biochronology of the Arikareean through Hemphillian interval (late Oligocene through earliest Pliocene epochs). Pp. 169–231 in M. O. Woodburne (ed.), Late Cretaceous and Ce- nozoic Mammals of North America: Biostra- tigraphy and Geochronology. Columbia Uni- versity Press, New York. Tedford, R. H., and D. Frailey. 1976. Review of some Carnivora (Mammalia) from the Thomas Farm Local Fauna (Hemingfordian, Gilchrist County, Florida). American Museum Novi- tates 2610:1–9. Tedford, R H., M. P. Skinner, R. W. Fields, J. M. Rensberger, D. P. Whistler, T. Galusha, B. E. Taylor, J. R. Macdonald, and S. D. Webb. 1987. Faunal succession and biochronology of the Arikareean through Hemphillian inter- val (late Oligocene through earliest Pliocene epochs) in North America. Pp. 153–210 in M. O. Woodburne (ed.), Cenozoic Mammals of North America: Geochronology and Bio- stratigraphy. University of California Press, Berkeley. Tedford, R. H., J. B. Swinehart, C. C. Swisher, III, D. R. Prothero, S. A. King, and T. E. Tierney. 1996. The Whitneyan-Arikareean transition in the High Plains. Pp. 312–334 in D. R. Prothero and R. J. Emry (eds.), The Terrestrial Eocene- Oligocene Transition in North America. Cam- bridge University Press, Cambridge. Teeling, E. C., M. S. Springer, O. Madsen, P. Bates, S. J. O’Brien, and W. J. Murphy. 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307:580–584. Teeling, E.C., S. C. Vernes, L. M. Dávalos, D. A. Ray, M. T. P. Gilbert, E. Myers, and Bat1k Con- sortium. 2018. Bat biology, genomes, and the Bat1k project: to generate chromosome-level 234 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3) genomes for all living bat species. Annual Re- view of Animal Biosciences 2018.6:23–46. Tejedor, A. 2011. Systematics of funnel-eared bats (Chiroptera: Natalidae). Bulletin of the Ameri- can Museum of Natural History 352: 1–140. Tejedor, M. F., N. J. Czaplewski, F. J. Goin, and E. Aragón. 2005. The oldest record of South American bats. Journal of Vertebrate Paleon- tology 25:990–993. Tessman, N.T. 1969. The fossil sharks of Florida. M.S. thesis, University of Florida, Gainesville. Thomas, O. 1908. The missing premolar of the Chiroptera. Annals and Magazine of Natural History 1(4): 346–348. Timm, R.M., and H. H. Genoways. 2004. The Florida bonneted bat, Eumops floridanus (Chi- roptera: Molossidae): distribution, morpho- metrics, systematics, and ecology. Journal of Mammalogy 85:852–865. Ubilla M., P. Gaudioso, and D. Perea. 2019. First fossil record of a bat (Chiroptera, Phyllostomi- dae) from Uruguay (Plio-Pleistocene, South America): a giant desmodontine. Historical Biology. doi:10.1080/08912963.2019.159035 2. Uvizl, M., J. Šmíd, T. Aghová, Z. Kotyková Varadínová, and P. Benda. 2019. Molecular phylogeny and systematics of the sheath-tailed bats from the Middle East (Emballonuridae: Taphozous and Coleura). Acta Chiropterolog- ica 21:23–34. Vannatta, J. M., J. A. Gore, V. L. Mathis, and B. D. Carver. 2021. Eumops floridanus (Chiroptera: Molossidae). Mammalian Species 1009:125– 133. Vaughan, T. A. 1959. Functional morphology of three bats: Eumops, Myotis, Macrotus. Univer- sity of Kansas Publications, Museum of Natu- ral History 12:1–153. Wang, X., R. H. Tedford, and B. E. Taylor. 1999. Phylogenetic systematics of the Borophaginae (Carnivora: Canidae). Bulletin of the Ameri- can Museum of Natural History 243:1–391. Webb, S. D. 1981. The Thomas Farm fossil site. Plaster Jacket 37:6–25. Wesselman, H. B. 1984. The Omo micromammals – systematics and paleoecology of early man sites from Ethiopia. Pp. 1–219 in M. K. Hecht and F. S. Szalay (eds.), Contributions to Verte- brate Evolution. Karger, Basel. Woodburne, M. O. (ed.) 2004. Late Cretaceous and Cenozoic Mammals of North America: Bio- stratigraphy and Geochronology. Columbia University Press, New York. Woodburne, M. O. 2010. The Great American Biotic Interchange: Dispersals, tectonics, cli- mate, sea level and holding pens. Journal of Mammal Evolution 17:245–264. javascript:; javascript:; javascript:; javascript:; javascript:; A1-1MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida Emballonuridae. New World. Balantiopteryx io: USNM 267343 (♂), Guatemala, Izabal, Bohos, skull and mandible. Balantiopteryx plicata: USNM 146943 (♂), Mexico, Baja California, Santa Anita, skull and mandible; USNM 559432 (♂), Mexico, Nayarit, San Blas, skull and mandible; OMNH 26646 (♂), Mexico: Colima: Colima: 1 km NE Cardona; OMNH 26681 (♀), Mexico: Colima: Manzanillo: Playa de Oro. Centronycteris centralis: USNM 503827 (♀), Panama, Canal Zone, Barro Colorado Island, skull and skeleton; USNM 535021 (♂), Panama, Canal Zone, Barro Colorado Island, skull and mandible. Cormura brevirostris, USNM 315139 (♂), Panama, Almirante, skull and mandible; USNM 464551 (♀), Panama, Bocas del Toro, Cayo Nancy, skull and mandible. Cyttarops alecto: USNM 566432 (♀), Costa Rica, Li- mon, Tortuguero, skull and skeleton. Diclidurus albus: USNM 407098 (♂), Venezuela, Amazonas, San Juan, skull and mandible; USNM 418688 (♂), Venezuela, Zulia, El Rosari, skull and mandible; OMNH 26706 (♂), Mexico: Colima: Manzanillo: Playa de Oro. Diclidurus ingens: USNM 407091 (♀), Venezu- ela, Amazonas, San Juan, skull and mandible. Diclidurus isabellus: USNM 388542 (♂), Venezuela, Ama- zonas, Boca Mavaca, skull and mandibls; USNM 388548 (♀), Venezuela, Amazonas, 68 km SE of Es- meralda, skull and mandible. Peropteryx macrotis: UF-M 6935 (sex unknown), Guatemala, Petén, Tikal, skull and skeleton; USNM 313146 (♂), Panama, Buena Vista, skull and mandible; USNM 393000 (♂), Brazil, Para, Belem, Mocambo, skull and mandible. Peropteryx kappleri: USNM 418630 (♀), Venezuela, Falcon, Cerro Caridad, skull and mandible; USNM 461844 (♀), Colombia, Valle, SW of Jamundi, skull and mandible. Peropteryx trinitatis: USNM 338936 (♂), Guyana, Rupununi, Dananawa, skull and man- dible. Rhynchonycteris naso: USNM 315101 (♂), Panama, Almirante, skull and mandible; USNM 562211 (♀), Peru, Madre de Dios, skull and skeleton. Saccopteryx bilineata: UF 6971-M (♀), Guatemala, Petén, Tikal, skull and skeleton; USNM 549311 (♀), Brazil, Para Altamira, skull and skeleton; OMNH 14334 (♀), Mexico: Colima: Cuauhtémoc: El Cobano; Saccopteryx canescens: USNM 392996 (♂), Brazil, Para, Belem, Mocambo, skull and mandible. Saccopteryx gymnura: USNM 460080 (♀), Brazil, Para, Belem, Mocambo, skull and mandible. Saccopteryx leptura: USNM 392999 (♂), Brazil, Para, Belem, Mocambo, skull and skeleton; USNM 513430 (♂), Ecuador, Zamora-Chinchipe, Los Encuentros, skull and mandible. Old World. Coleura afra: USNM 350817 (♂), Kenya, Diana, skull and mandible; USNM 350831 (♂), Kenya, Diana, skull and skeleton. Emballonura alecto: USNM 458529 (♂), Philippines, Leyte, In- opacan, skull and mandible; USNM 458549, (♂), Philippines, Leyte, Inopacan, skull and mandible. Mosia nigrescens: USNM 277114 (♀), Papua New Guinea, Emirau Island, skull and mandible. Saccolaimus flaviventris: USNM 284163 (♂), Australia, Northern Territory, Port Langde, skull and mandible. Sacco- laimus peli: USNM 481709 (♀), Liberia, Grand Gedeh, Tars Town, skull and mandible. Saccolaimus plu- to: USNM 458550 (♂), Philippines, Negros, Dumaguete, skull and mandible. Saccolaimus saccolaimus: USNM 294813 (♀), Thailand, Ratchaburi, Tapa, skull and mandible. Taphozous melanopogon: USNM 251744 (♀), Thailand, Bangkok, skull and mandible; USNM 252225 (♂), Thailand, Bangkok, skull and mandible. Taphozous melanopogon: USNM 458557 (♀), Philippines, Maripipi Island, Maripipi, skull and skeleton. Taphozous nudiventris: USNM 300209 (♂), Egypt, Saqqara, skull and mandible. Non-Emballonuridae examined. Nycteridae. Nycteris grandis: USNM 411849, Ghana, Volta, Kpeve. Nycteris hispida: AMNH 184478, Sudan, Bahr el Ghazal, Yirol Dist. Nycteris thebaica: AMNH 168140, Botswana, Ngamiland, Mohembo. Rhinopomatidae. Rhinopoma microphyllum: AMNH 244388, Pakistan. APPENDIX 1. Modern comparative specimens of Emballonuridae and outgroups examined in this study. Species identification, museum acronym, catalog number, sex, basic locality information, and nature of specimen (e.g., cranium and mandible, skull and skeleton) are provided for each specimen. The taxonomy follows Simmons (2005). A2-1MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida APPENDIX 2. Selected characters and character states of the upper and lower dentition and dentary of Emballonuridae. Characters 1-65 are from Ravel et al. (2016), translated from French; characters 66-85 are new to this study. The numbered characters (1-85) and character states within each character (i.e., 0, 1, 2) listed below are analyzed in Table 4 for 14 species of extinct and extant species of Emballonuridae, including three extinct species from Florida described in this paper. The dental terminology used in the characters below, in Table 4, and in the text follows Czaplewski et al. (2008) and differs somewhat from that used by Ravel et al. (2016). Equivalent dental terms are both used in this appendix. Our term is listed first followed by the Ravel et al. (2016) term in parentheses (e.g., labial = buccal; anterior = mesial; pos- terior = distal; paracingulum= precingulum; metacingulum = postcingulum, etc.). CHARACTERS FROM RAVEL ET AL. (2016) Dentary 1. Coronoid process height 0. high (about twice the height of the dental row) 1. low 2. Coronoid process apex 0. sharp 1. rounded 3. Horizontal ramus 0. gracile (lower than height of molars) 1. robust (greater than height of molars) 4. Coronoid process angle 0. vertical (almost perpendicular to horizontal ramus) 1. oriented posteriorly Lower dentition 5. Number of lower incisors 0. three incisors on each mandible 1. two incisors on each mandible 6. Number of lower premolars 0. three premolars on each mandible 1. two premolars on each mandible 7. Lower incisors lobes 0. bilobed 1. trilobed 8. Lower incisors relative size 0. incisors of equivalent size BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3)A2-2 8. Lower incisors relative size (cont.) 1. i3 widest of lower incisors 2. i1 widest of lower incisors 9. c1 labial (=buccal) cingulum 0. strong (well developed and continuous) 1. weak (very thin or absent) 10. c1 root 0. straight 1. curved posteriorly 11. c1 posterior basin 0. extended posteriorly 1. reduced 12. p2 size 0. reduced 1. not reduced (size equivalent to p4) 2. larger than p4 13. p3 roots 0. one root 1. two roots 14. p3 crown 0. well developed 1. very small 15. p4 size 0. well developed (size larger than the anterior premolar, with a protoconid and talonid) 1. reduced (size smaller than the previous premolar 16. p4 talonid 0. extended posteriorly 1. reduced 2. absent 17. p4 paraconid 0. present 1. absent APPENDIX 2. Cont. A2-3MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 18. p4 metaconid 0. present 1. absent 19. Lower molar trigonid cusps 0. high and well separated 1. low 20. m1 trigonid 0. lingually open 1. compressed anteroposterioly (=mesiodistally) 21. m2 trigonid 0. lingually open 1. compressed anteroposterioly (=mesiodistally) 22. m1: distance between the paraconid and metaconid 0. less than distance between metaconid and entoconid 1. greater than or equal to distance between metaconid and entoconid 23. m2: distance between the paraconid and the metaconid 0. less than distance between metaconid and entoconid 1. greater than or equal to distance between metaconid and entoconid 24. m3: distance between the paraconid and the metaconid 0. less than distance between metaconid and entoconid 1. greater than or equal to distance between metaconid and entoconid 25. m1-m2 trigonid width 0. less than that of talonid 1. greater to that of talonid 2. equivalent to width of talonid 26. m1-m2 entoconid 0. high and inclined posteriorly (similar in size to metaconid) 1. absent or very reduced 2. moderate and upright (smaller than metaconid) 27. m1-2 entocristid 0. straight 1. slightly curved 2. strongly curved/sharply V-shaped APPENDIX 2. Cont. BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3)A2-4 28. m1-m2 cristid obliqua meets postcristid 0. medial; meets posterior wall of trigonid (= protocristid) at junction between metacristid and postprotocristid (dental terminology of Ravel et al., 2016). 1. lingual; meets posterior wall of trigonid (= protocristid) in more lingual position than junction between metacristid and postprotocristid. 2. labial; meets posterior wall of trigonid (= protocristid) in more labial position than junction between metacristid and postprotocristid 29. m1-m2 hypoconulid position 0. central and posterior position on postcristid between entoconid and hypoconid 1. on postcristid slightly more labial than the entoconid 2. very close and posterior to entoconid 30. m1-m2 postcristid 0. straight 1. curved posteriorly 31. m3 talonid width 0. equivalent to width of trigonid 1. less than the width of trigonid 32. m3 hypoconulid 0. present 1. weak/absent Upper dentition 33. C1 labial (=buccal) cingulum 0. present 1. absent 2. very thin and discontinuous 34. C1 lingual cingulum 0. strongly curved 1. almost straight 35. C1 posterior break in cingulum 0. present 1. absent 36. Number of upper premolars 0. three 1. two 2. one APPENDIX 2. Cont. A2-5MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 37. P4 talon basin 0. elongated or expanded lingually 1. compressed labiolingually (=buccolingually) 38. P4 crown, anterolabial (=buccomesial) lobe 0. with pronounced anterolabial (=buccomesial) lobe 1. without anterolabial (=buccomesial) lobe 2. with weak anterolabial (=buccomesial) lobe 39. P4 anterolingual (=mesiolingual) tubercle/root 0. present 1. absent 40. M1-M2 ectoflexus 0. double 1. simple and anterior to mesostyle 2. simple and central 41. M1-M2 “V” of ectoloph asymmetry 0. weak 1. pronounced 42. M1 inclination of labial (=buccal) margin 0. strong 1. weak 43. M1-M2 position of mesostyle 0. lingual, set back from labial (=buccal) margin 1. labial, extends outward from labial (=buccal) margin 44. M1-M2 labial (=buccal) cingulum 0. discontinuous along labial (=buccal) border 1. continuous along labial (=buccal) border 45. M1-M2 labial (=buccal) cingulum at mesostyle 0. present 1. absent 46. M1 parastyle 0. connected to the preparacrista 1. isolated from preparacrista 2. absent APPENDIX 2. Cont. BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3)A2-6 47. M1-M2 paraloph 0. present 1. absent 48. M1-M2 metaloph 0. present 1. absent 49. M1-M2 paracingulum (=precingulum) 0. wide 1. very thin to absent 50. M1-M2 metacingulum (=postcingulum) 0. wide 1. thin 51. M1-M2 protocone 0. well developed; larger than metacone and paracone 1. reduced; equal to or smaller than metacone and paracone 52. M1-M2 connection of posterior extension of postprotocrista 0. connects with metacingulum (=postcingulum) 1. connects with lingual cingulum 2. does not connect with any other structure 3. connects with hypocone 53. M1-M2 connection of preprotocrista 0. connects to paracingulum (=precingulum) 1. meets base of paracone 54. M1-M2 protofossa extension 0. extends along anteroposterior (=mesiodistal) axis 1. short 55. M1-M2 posterior (=distal) opening of protofossa 0. closed posteriorly (distally) by postprotocrista 1. open 56. M1-M2 level of protofossa 0. low and opening onto talon basin 1. high and without a connection with talon basin APPENDIX 2. Cont. A2-7MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 57. M1-M2 hypocone 0. present and well developed 1. present but small/low 2. absent 58. M1-M2 posterior (=distal) flattening of talon 0. strong 1. weak 2. absent 59. M1-M2 posterior (=distal) border of talon 0. rounded 1. with posterolingual (=distolingual) lobe or extension 60. M1-M2 orientation of talon 0. posterior (=distal) 1. posterolingual (=distolingual) 61. M1-M2 lingual (=buccal) cingulum 0. thick 1. thin/absent 62. M3 size 0. more than half the width of M2 1. less than half the width M2 63. M3 metacone 0. small 1. well developed (equivalent to paracone) 2. absent 64. M3 lingual (=buccal) cingulum 0. present 1. absent 65. M3 premetacrista 0. present 1. absent NEW CHARACTERS FROM THIS STUDY 66. c1 lingual cingulum 0. well developed 1. thin or absent APPENDIX 2. Cont. BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 60(3)A2-8 67. c1 anterolingual cusp 0. present 1. absent 68. c1 posterolingual cusp 0. present 1. absent 69. p2 occlusal shape 0. rounded 1. elongated anteroposteriorly 70. p4 occlusal shape 0. rounded 1. elongated anteroposteriorly 71. m1 and m2 morphology 0. similar, both with lingually open trigonids 1. similar, both with lingually compressed trigonids 2. different, m1 with open trigonid, m2 with more compressed trigonid 72. m1 trigonid 0. open lingually 1. compressed 73. m1/m2 protoconid and metaconid placement 0. protoconid anterior to metaconid 1. metaconid anterior to protoconid 2. protoconid and metaconid at same level 74. m1/m2 protocristid 0. transverse or horizontal to toothrow 1. posterolingually inclined from protoconid to metaconid 2. anterolingually inclined from protoconid to metaconid 75. m1/m2 entoconid 0. smaller than metaconid 1. equal in size to metaconid 2. larger than metaconid 76. m1/m2 hypoconulid 0. well developed 1. present but small APPENDIX 2. Cont. A2-9MORGAN AND CZAPLEWSKI: Bats from the Oligocene and early Miocene of Florida 77. m1/m2 hypoconid and entoconid placement 0. hypoconid anterior to entoconid 1. entoconid anterior to hypoconid 2. hypoconid and entoconid at same level 78. m1/m2 postcristid 0. transverse or horizontal to toothrow 1. posterolingually inclined from hypoconid to entoconid 2. anterolingually inclined from hypoconid to entoconid 79. m1/m2 labial cingulum 0. thick/strong 1. thin, continuous 2. thin, discontinuous 80. m3 talonid 0. very narrow, less than half the width of trigonid 1. narrow, about half the width of trigonid 2. broad, greater than half to almost equal to width of trigonid 81. m3 entoconid 0. well developed, about equal to metaconid 1. small to absent, much smaller than metaconid 82. C1 anterolingual cusp 0. present 1. absent 83. C1 posterolingual cusp 0. present 1. absent 84. M1 parastylar region, including parastyle and parafossa 0. weakly reduced, parastyle connected to ectoflexus 1. reduced, parastyle isolated 2. reduced, parastyle absent 85. M1 preparacrista 0. weakly reduced, ~½ the length of M2 preparacrista 1. short, less than ½ the length of M2 preparacrista 2. absent APPENDIX 2. Cont.