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

Journal of Mammalogy, 84(2):729±752, 2003

A NEW BAT (CHIROPTERA: NATALIDAE) FROM THE EARLY MIOCENE OF FLORIDA, WITH COMMENTS ON NATALID PHYLOGENY

GARY S. MORGAN* AND NICHOLAS J. CZAPLEWSKI New Mexico Museum of Natural History, 1801 Mountain Road NW,

Albuquerque, NM 87104, USA (GSM) Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK 73072, USA (NJC)

We describe a new extinct genus and species of bat belonging to the endemic Neotropical family Natalidae (Chiroptera) from the Thomas Farm Local Fauna in northern peninsular Florida of early Miocene age (18±19 million years old). The natalid sample from Thomas Farm consists of 32 fossils, including a maxillary fragment, periotics, partial dentaries, isolated teeth, humeri, and radii. A proximal radius of an indeterminate natalid is reported from the I-75 Local Fauna of early Oligocene age (about 30 million years old), also from northern Florida. These fossils from paleokarst deposits in Florida represent the 1st Tertiary records of the Natalidae. Other extinct Tertiary genera previously referred to the Natalidae, including Ageina, Chadronycteris, Chamtwaria, Honrovits, and Stehlinia, may belong to the superfamily Nataloidea but do not ®t within our restricted de®nition of this family. Eight derived characters of the Natalidae sensu stricto are discussed, 5 of which are present in the new Miocene genus. Intrafamilial phylogenetic analysis by parsimony of the Natal- idae suggests that the 3 living subgenera, Natalus (including N. major, N. stramineus, and N. tumidirostris), Chilonatalus (including C. micropus and C. tumidifrons), and Nyctiellus (including N. lepidus), deserve full generic rank. The Natalidae apparently evolved in North America before the late Oligocene, went extinct in what is now the Nearctic region (i.e., Florida) after the early Miocene, and survived in tropical Middle America during the re- mainder of the Tertiary. The presence of 2 endemic genera and 4 endemic species suggests that natalids reached the West Indies by overwater dispersal early in their history (Oligocene or Miocene). The lack of a Tertiary fossil record, marginal distribution, and limited species richness and endemism of natalids in South America are suggestive of a comparatively late arrival on that continent, possibly in the late Pliocene after the beginning of the Great American Faunal Interchange.

Key words: bat, biogeography, Chiroptera, Florida, fossil, Miocene, Natalidae, Nataloidea, paleo- karst, phylogeny

We describe a new extinct genus and spe- West Indies. The species described in this cies of bat in the endemic Neotropical fam- study is the 1st known extinct member of ily Natalidae from the early Miocene (early the Natalidae, excluding 5 other extinct Ter- Hemingfordian land mammal age) Thomas tiary genera (Ageina, Chadronycteris, Farm Local Fauna in northern peninsular Chamtwaria, Honrovits, and Stehlinia) that Florida. The Natalidae are a small family have been considered natalids based on the composed of 6 living species restricted to concept of the family proposed by Van Val- Middle America, South America, and the en (1979). These 5 genera are referred to the superfamily Nataloidea following Sim- * Correspondent: [email protected]

729 730 JOURNAL OF MAMMALOGY Vol. 84, No. 2 mons and Geisler (1998) but are not con- ric tons of sediment screenwashed between 1981 sidered members of the Natalidae sensu and 1985 by Ann Pratt and Arthur Poyer as part stricto. of Pratt's dissertation on the taphonomy and pa- We also report a proximal radius of a na- leoecology of the Thomas Farm vertebrate fauna talid from the Oligocene I-75 Local Fauna, (Pratt 1989, 1990; Pratt and Morgan 1989). The use of ®ne-mesh screens (1-mm opening) re- located near Gainesville, Alachua County, sulted in the recovery of large numbers of iso- Florida, about 80 km southeast of Thomas lated bat teeth that passed through the standard Farm. The I-75 site is the oldest land-ver- window screening used by earlier workers. Bat tebrate locality in Florida (Patton 1969), fossils were recovered throughout the 3-m sec- dating to the Whitneyan land mammal age tion of sediments excavated at Thomas Farm in Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 (early Oligocene, about 30 ϫ 106 years the early 1980s, but the greatest number of bats ago). The radius from the I-75 site cannot occurred in a 1-m-thick unit of lime sand near be identi®ed below the family level but is the top of the section (unit 15; Pratt 1989, important because it represents the oldest 1990). fossil record of the Natalidae. All fossils cited are from the vertebrate pale- The discovery of a new genus and spe- ontology collection, FLMNH, University of cies of Natalidae from the early Miocene of Florida, Gainesville (UF). We follow the chirop- teran dental terminology of Legendre (1984) and Florida provides new information on the Menu and Sige (1971). We use standard abbre- historical biogeography and phylogeny of viations for tooth positions in mammals, with this small and enigmatic group of New uppercase letters for upper teeth and lowercase World bats. The identi®cation of a natalid letters for lower teeth: I and i (upper and lower nearly 20 million years older than any other incisors), C and c (upper and lower canines), P known member of this family prompted us and p (upper and lower premolars), and M and to examine the phylogenetic relationships m (upper and lower molars). The terminology of all species within the Natalidae. Al- for chiropteran postcranial elements follows though our primary aim was to determine Smith (1972) and Vaughan (1959). All measure- the systematic position of the new Florida ments of fossils are in millimeters. Miocene natalid, we also examined the re- We compared the Thomas Farm natalid fossils lationships among the 6 extant species of with at least 2 skulls and 2 complete skeletons natalids because no formal hypothesis of re- of each of the 6 extant species of Natalidae and the potentially related families Furipteridae, lationships has been proposed for the fam- Thyropteridae, and Vespertilionidae (see Appen- ily. dix I for comparative material examined). Methods of phylogenetic analysis.ÐThere is MATERIALS AND METHODS no previous intrafamilial phylogenetic analysis In the late 1950s and early 1960s, Clayton for the Natalidae. We developed a hypothesis of Ray of the Florida State Museum (now the Flor- relationships based on parsimony analysis (using ida Museum of Natural History [FLMNH]) and PAUP 4.0b10 softwareÐSwofford 2000) of the Pierce Brodkorb of the University of Florida De- Floridian fossils and all 5 extant species gener- partment of Zoology began screenwashing sed- ally recognized as members of the Natalidae iments from Thomas Farm for small vertebrates. (Koopman 1993, 1994; Nowak 1994; Appendix These early screenwashing efforts led to system- I) plus Natalus major, which we recognize as a atic studies of frogs (Holman 1965, 1967), sal- valid species following Morgan (1989b). Mor- amanders and lizards (Estes 1963), snakes (Auf- phological comparisons disclosed 50 characters fenberg 1963), birds (Brodkorb 1956), and ro- that may be phylogenetically informative (Ap- dents (Black 1963). Bat fossils also were recov- pendix II). We scored 24 cranial, dental, and hu- ered during this early screenwashing program, meral characters that are present in the fragmen- but early ®nds did not include specimens of the tary fossils as well as in extant natalids (Appen- new natalid. The majority of bat fossils from dix II, characters 1±24). The remaining 26 cra- Thomas Farm, including all natalid specimens nial and postcranial characters were available described here, were collected from about 2 met- only in the extant natalids (with missing values May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 731 coded for the fossil taxon). Characters and char- acter states are based on Dalquest (1950), Good- win (1959), Koopman (1994), Miller (1907), Ot- tenwalder and Genoways (1982), Silva Taboada (1979), Simmons (1998), Simmons and Geisler (1998), Van Valen (1979), and our personal ob- servations. There has been controversy surrounding the higher-level relationships of the Natalidae. In 1 study by Simmons and Geisler (1998), the Na- talidae were part of a clade (Nataloidea) that Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 also included the Furipteridae, Thyropteridae, and Myzopodidae. In their analysis (Simmons and Geisler 1998), this clade was sister to the Vespertilionoidea (Vespertilionidae and Molos- sidae) and farther removed from the Noctilion- oidea. In a contrasting analysis based on mito- chondrial DNA, Van Den Bussche and Hoofer (2001) found the nataloids (except Myzopodi- FIG. 1.ÐLocation of the 2 sites in Florida that dae) strongly associated with the Noctilionoidea have produced Tertiary fossils of Natalidae, I-75 (Noctilionidae, Mystacinidae, Mormoopidae, (Oligocene) and Thomas Farm (early Miocene). and Phyllostomidae) and far removed from the Vespertilionidae and Molossidae. More recently, cated 12 km north of Bell in Gilchrist County in using additional evidence from nuclear genes, northern peninsular Florida (Fig. 1), has pro- Hoofer et al. (in press) reexamined yangochirop- duced the best known early Miocene (early teran bats, including all the aforementioned fam- Hemingfordian land mammal age) vertebrate ilies. In their analysis Nataloidea was found to fauna in eastern North America. There are de- be polyphyletic; Natalidae clustered with Ves- tailed accounts of the discovery, excavation, ge- pertilionidae and Molossidae, whereas Furipter- ology, and stratigraphy of Thomas Farm (Auf- idae and Thyropteridae clustered with the Noc- fenberg 1963; Pratt 1989, 1990; Simpson 1932; tilionoidea. Based in part on the analysis of White 1942), as well as faunal lists and bibli- Hoofer et al. (in press) and in part on the tra- ographies (Olsen 1962; Ray 1957; Webb 1981). ditional association of Natalidae with Vesperti- The Thomas Farm Local Fauna is composed lionidae from morphological data, we used 2 of about 90 species of vertebrates (Webb 1981), vespertilionids (Kerivoula picta and Myotis lu- primarily consisting of terrestrial forms, al- cifugus) as a monophyletic out-group in our though freshwater taxa are present, including phylogenetic analysis. Moreover, in a recent ar- frogs, salamanders, pond turtles, alligators, and ticle, HoraÂcek (in press) noted that among living aquatic birds. There are 23 species of large vespertilionids, Myotis and Kerivoula had the mammals from Thomas Farm, the most abun- most primitive dentition. dant of which is the three-toed horse, Parahip- A simple branch-and-bound search yielded a pus leonensis (Hulbert 1984). This site has an single most parsimonious tree of 82 steps. We exceptionally rich small-vertebrate fauna of then used the bootstrap and jackknife methods nearly 70 species, including large and taxonom- with branch-and-bound search to examine trees ically diverse samples of frogs, lizards, snakes, that were 1 step to a few steps longer than the birds, bats, and rodents. In addition to bats, other most parsimonious tree in order to calculate Bre- small mammals from Thomas Farm include a mer support values for branches in the consen- didelphid, a soricid, 3 sciurids, 2 heteromyids, sus cladograms (Simmons 2000). Results from and an eomyid (Pratt 1989; Webb 1981). bootstrap and jackknife methods were very sim- The fossiliferous Miocene sediments at ilar, so only the bootstrap results are reported Thomas Farm consist of alternating layers of below. The number of bootstrap replicates was clay and sand ®lling a 30-m-deep sinkhole de- set to 10,000 but was similar when set to 1,000. veloped in Eocene marine limestone (Pratt 1989, Fossil localities.ÐThe Thomas Farm site, lo- 1990). The fossil assemblage from the lime sand 732 JOURNAL OF MAMMALOGY Vol. 84, No. 2 appears to represent a cave deposit that formed described genus similar to Corynorhinus. The through 2 processes: the natural accumulation of Vespertilionidae dominate the Thomas Farm chi- bat carcasses on a cave ¯oor and a coprocoe- ropteran fauna. nosis representing the fecal remains from small The I-75 Local Fauna is not nearly as well mammalian carnivores or the regurgitation pel- documented as Thomas Farm, with only 1 sum- lets of raptorial birds. Pratt (1989) concluded mary paper on the locality (Patton 1969). The that the Thomas Farm site consisted of a deep site was discovered in 1965 during the construc- sinkhole surrounded by a forested habitat. The tion of Interstate Highway 75 through Gaines- abundance of bats suggests that caves probably ville, Alachua County, in northern peninsular occurred in the walls of the sinkhole. Florida (Fig. 1). The I-75 site consists of fossil- The mammalian fauna from Thomas Farm, in iferous clays deposited in a small karst solution Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 particular the carnivores, equids, artiodactyls, cavity developed in marine Eocene limestone. and sciurids (ForsteÂn 1975; Maglio 1966; Pratt The I-75 site has a diverse vertebrate fauna con- and Morgan 1989; Tedford and Frailey 1976), taining about 45 species (Patton 1969). The indicates a late early Hemingfordian age (be- large mammal fauna is composed of 2 carni- tween 18 ϫ 106 and 19 ϫ 106 years ago). The vores, including the amphicyonid Daphoenus early Hemingfordian is de®ned in part by the 1st and a mustelid, the horse Miohippus, a tayas- appearance of the immigrant carnivores Amphi- suid, an oreodont, and the small artiodactyl Lep- cyon, Hemicyon (Phoberocyon), and Leptarctus, tomeryx. I-75 also has a rich microvertebrate the 1st occurrence of the canid Tomarctus and fauna, including frogs, salamanders, lizards, the camelid Floridatragulus, and the last occur- snakes, and small mammals (Holman 1999; Pat- rence of the small amphicyonid Cynelos and the ton 1969). Small mammals include the didelphid rhinocerotid Menoceras (Tedford et al. 1987), all Herpetotherium; the insectivore Centetodon cf. of which are present at Thomas Farm. Other cor- C. wolf®; 6 species of bats (see below); the le- relative early Hemingfordian faunas from the porid Palaeolagus; and at least 4 species of ro- western United States are the Garvin Gully Fau- dents, Eutypomys, a heteromyid, and 2 eomyids. na of Texas, the faunas from the Runningwater Previous workers (Emry et al. 1987; Patton Formation of Nebraska, the Flint Hill Local Fau- 1969) placed the I-75 Local Fauna in the Whit- na of South Dakota, and the Martin Canyon Lo- neyan land mammal age (early Oligocene, 30 ϫ cal Fauna of Colorado (Tedford et al. 1987). 106 to 32 ϫ 106 years agoÐWoodburne and Fossil bats were initially reported from Thom- Swisher 1995), making it the oldest land ver- as Farm by Lawrence (1943), who described 2 tebrate fauna known from Florida. Whitneyan new genera and species of Vespertilionidae, faunas are otherwise unknown outside the Suaptenos whitei and Miomyotis ¯oridanus. northern Great Plains (Tedford et al. 1996). Czaplewski and Morgan (2000) recently de- Most of the age-diagnostic mammals from I-75 scribed a 3rd extinct genus and species of ves- occur in the Whitneyan and early Arikareean pertilionid from Thomas Farm, Karstala silva, land mammal ages. The presence of Centeto- characterized by its large size. All the natalid don, Eutypomys, Leptomeryx, Miohippus, and fossils from Thomas Farm were collected in the Palaeolagus in I-75 establishes an early Ari- early 1980s as part of an intensive screenwash- kareean or older age (older than 24 ϫ 106 years ing project (Pratt 1989, 1990). Thomas Farm has agoÐTedford et al. 1987, 1996). The similarity the largest known sample of bats from any pre- of the bat faunas from I-75 and the early Ari- Pleistocene fossil deposit in North America, kareean (26 ϫ 106 to 28 ϫ 106 years ago) numbering almost 2,000 specimens. The chirop- Brooksville 2 Local Fauna from central Florida teran fauna is composed of at least 8 species, (Hayes 2000) suggests that these 2 faunas are including 1 species each in the Neotropical fam- close in age. The absence of mammals in the ilies Emballonuridae, Natalidae, and Molossidae I-75 Local Fauna that are restricted to the Ari- and 5 species of Vespertilionidae (Czaplewski kareean (e.g., the small artiodactyl Nanotra- and Morgan 2000; Morgan 1989a; Morgan and gulus) indicates a late Whitneyan age (about 30 Czaplewski 2000). In addition to Suaptenos whi- ϫ 106 years ago), making the I-75 natalid spec- tei, Miomyotis ¯oridanus, and Karstala silva, the imen about 12 million years older than the Thomas Farm vespertilionids include an unde- Thomas Farm natalid. scribed genus similar to Lasiurus and a 2nd un- The I-75 site provides the earliest record of May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 733 the taxonomic diversity and community struc- ture of North American middle Cenozoic chi- ropteran faunas. In addition to the natalid re- ported in this study, 5 other species of bats are known from I-75, including a new mormoopid, 2 new emballonurids, a new possible phyllos- tomid, and a vespertilionid. Species belonging to families now restricted to the Neotropics are the most abundant bats in the I-75 fauna, including a large emballonurid and a small mormoopid. Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 SYSTEMATIC PALEONTOLOGY Primonatalus, new genus

Type species.ÐPrimonatalus prattae. Included species.ÐOnly the type species is known. Diagnosis.ÐMandibular ramus is deep below molars. Ventral margin of mandible is straight between p4 and mandibular an- gle. Summit of coronoid process of man- dible is comparatively well developed with triangular-shaped dorsal tip. Anterior edge of coronoid process curves posterodorsally from alveolar margin to dorsal tip of coro- noid at an angle of about 70Њ. Coronoid pro- cess rises above level of articular condyle. Posterior portion of coronoid process slopes ventrally from tip of coronoid down to con- dyle. Distinct mandibular angle is present directly ventral to tip of coronoid. Base of angular process is located halfway between ventral edge of mandible and alveolar mar- gin in vertical dimension and about halfway FIG. 2.ÐDentaries of Primonatalus prattae n. between coronoid and condyle in antero- gen. et sp. A±C) Partial right dentary with m1± 3 (UF 108641, holotype) in labial, occlusal, and posterior dimension. Angular process does lingual views. D and E) Posterior fragment of not ¯are strongly laterally. Mandibular fo- right dentary with intact angular process (UF ramen opens level with alveolar margin. 121145) in labial and lingual views. F±H) Eden- Two features, the comparatively well-de- tulous left partial ramus with mandibular sym- veloped triangular-shaped coronoid process physis and alveoli for i1±3, c1, and p2±4 (UF that is taller than the articular condyle and 108647) in labial, occlusal, and lingual views. the ventral position of the mandibular fo- ramen and angular process, re¯ect a lesser degree of dorsal cranial ¯exion in Primon- height on m1 and m2. Carnassial-like atalus than in Natalus. Posterior mental fo- notches on cristid obliqua and postcristid on ramen located in deep concavity near al- lower molars (Figs. 2A, 2B, and 3C) pre- veolar margin between roots of c1 and p2. sent but weak. Lophid associated with car- The p3 is larger, longer, and more com- nassial-like notch extending posteriorly pressed with distinct concavity posterior to from cristid obliqua into talonid basin ab- main cusp. Metaconid and entoconid same sent. Talonid notch was deeper than in Re- 734 JOURNAL OF MAMMALOGY Vol. 84, No. 2

Primonatalus prattae, new species Holotype.ÐUF 108641, partial right den- tary with m1±3 (Figs. 2A±C and 3C). The holotype, paratypes, and all referred speci- mens are from the early Miocene Thomas Farm Local Fauna, Florida. Paratypes.ÐUF 121145, ascending ra- mus of right dentary (Figs. 2D and 2E); UF 108647, edentulous partial left dentary with Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 symphysis and alveoli for i1±p4 (Figs. 2F± H); UF 108642, partial left dentary with p3 and alveoli for p2 and p4 (Figs. 3A and 3B); UF 121136, right P4 (Figs. 4E±G); UF 108638, partial left maxilla with M1±2 (Figs. 4B±D); UF 121139, right M3 (Fig. 4A); UF 121141, periotic (Figs. 5A±D); UF 108650, distal half of left humerus (Fig. 6). Referred specimens.ÐUF 121137, partial left maxilla with M1; UF 108639, 108640, 2 right M1; UF 121138, left M1; UF FIG. 3.ÐLower teeth of Primonatalus prattae 121140±121142, 3 periotics (2 left, 1 right); n. gen. et sp. A) Left dentary fragment with p3, UF 121143, partial left dentary with m1±2; alveoli for c1, p2, and p4, and anterior root of UF 108643, partial right dentary with m1 m1 (UF 108642) in labial view. B) Same as A, and alveoli for p4 and m2; UF 108644, par- in occlusal view. C) Right lower molar series tial left dentary with m1; UF 108646, eden- (UF 108641, holotype) in occlusal view. tulous partial right dentary with alveoli for m1±3; UF 121144, edentulous partial left dentary with alveolus for m3; UF 108645, cent natalids because of taller metaconid right m3; UF 108648, 108649, 108651, and more ventral connection of entocristid 108652, 4 distal portions of humeri (3 right, to trigonid. Cristid obliqua also connects to 1 left); UF 108653, 108654, 2 right proxi- trigonid more ventrally. Labial cingular mal portions of radii; UF 108655±108658, cusp of P4 is low. Occlusal outline of P4 4 distal portions of radii (2 right, 2 left); has anterior indentation. Talon weakly de- UF 108660, right proximal metacarpal III; veloped on M1±2. Lingual cingulum weak UF 108659, distal portion of femur. on upper molars. Humerus has broad tri- Type locality and age.ÐThomas Farm angular-shaped epitrochlea with no notch Local Fauna (UF locality number GI01), 12 between weak medial process and distal km northeast of Bell, Gilchrist County, spinous process. Distal spinous process Florida, early Miocene, early Hemingfor- small, projecting only to edge of trochlea or dian. very slightly distal to it, and separated from Diagnosis.ÐSame as for genus. trochlea by narrow but distinct notch. Etymology.ÐNamed for Ann E. Pratt in Etymology.ÐPrimus (Latin), 1st, origi- recognition of her work on Miocene small nal, early, and Natalus (Latin), generic vertebrate faunas from Florida. Dr. Pratt's name for all currently recognized Recent screenwashing efforts at Thomas Farm pro- species in the family Natalidae. The name duced all the natalid fossils described in this indicates that this is the earliest known rep- study. resentative of the Natalidae. Descriptions and comparisons.ÐThe May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 735

Thomas Farm fossils of Primonatalus are compared with Recent Natalidae in consid- erable detail because of their importance in interpreting the phylogenetic relationships among all natalids. Most comparisons are with the 3 subgenera of Natalus: Natalus (including the species major, stramineus, and tumidirostris), Chilonatalus (including the species micropus and tumidifrons), and

Nyctiellus (including the species lepidus). Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 The subgenera of Natalus are discussed in the context of genera, and our phylogenetic analysis suggests that these 3 subgenera are best recognized as genera (see below). In- dividual species are mentioned only where signi®cant differences exist between the species in a subgenus. Primonatalus is in- termediate in size for the family, smaller than the species in the subgenus Natalus but larger than N.(Chilonatalus) micropus and N.(Nyctiellus) lepidus. The fossils are most similar in size to N.(Chilonatalus) tumidi- frons from the Bahamas. The most complete fossil of Primonatal- us prattae from Thomas Farm is the holo- type, a partial right dentary with all 3 mo- lars (UF 108641; Figs. 2A±C and 3C). The ascending ramus of the type is nearly com- plete, but the horizontal ramus is missing anterior to m1. The coronoid process and articular condyle are intact, and the angular process is missing only the tip. The hori- zontal ramus is comparatively deep below the molars and has a straight ventral edge. The ventral margin is nearly parallel to the alveolar margin of the toothrow from the p4 posteriorly to the mandibular angle. The horizontal ramus is also relatively deep FIG. 4.ÐUpper teeth of Primonatalus prattae with nearly parallel ventral and alveolar n. gen. et sp. A) Right M3 (UF 121139) in oc- margins in Natalus. The horizontal ramus clusal view. B±D) Left maxillary fragment with is more slender and curved ventrally below M1±2 and lateral alveoli for M3 (UF 108638) in the m1 and m2 in Chilonatalus and Nyc- occlusal, posterior, and labial views. E±G) Right tiellus. The coronoid process of Primona- P4 (UF 121136) in labial, occlusal, and lingual talus is triangular in shape and has a de®- views. nite dorsal tip. The tip of the coronoid is taller than the articular condyle so that the posterior portion of the coronoid process slopes posteroventrally from its tip to the condyle. In all Recent natalids, the coronoid 736 JOURNAL OF MAMMALOGY Vol. 84, No. 2 Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021

FIG. 5.ÐPeriotic bones of fossil and Recent Natalidae. A±D) Right periotic of Primonatalus prattae n. gen. et sp. (UF 121141) in A) lateral view (inverted), B) endocranial (dorsal) view, C) ventral view, and D) posterior view (inverted). E) Right periotic bone in skull of Chilonatalus tumidifrons (UF 24853 male from the Bahamas), posterior view of inverted skull (same view as D). Abbrevia- tions: ant. ϭ anterior; asc ϭ anterior semicircular canal; aub ϭ auditory bulla; bap ϭ basisphenoid pits; br ϭ broken semicircular canal exposing internal chamber; fec ϭ fenestra cochleae (ϭfenestra rotunda); fev ϭ fenestra vestibuli (ϭfenestra ovalis; for footplate of stapes); ff ϭ facial foramen; fom ϭ foramen magnum; fsm ϭ fossa for stapedius muscle; gic ϭ groove for internal carotid artery; gsa ϭ groove for stapedial artery; iam ϭ internal acoustic meatus; lat. ϭ lateral; lsc ϭ lateral semicircular canal; oc ϭ occipital condyle; pr ϭ small, hooked posteroventral tubercle on the periotic; psc ϭ posterior semicircular canal; pt ϭ pterygoid; saf ϭ subarcuate fossa; sc ϭ semicircular canal; sty ϭ stylohyal; va ϭ vestibular aqueduct; vent. ϭ ventral; za ϭ zygomatic arch. Labels are our provisional interpretations based on StaneÏk (1933) and Wible et al. (2001). process is more reduced and its anterior the tip of the coronoid in Recent natalids. edge forms a right angle with the horizontal The anterior edge of the coronoid in the ramus, whereas the posterior edge is nearly fossil is not vertical as in most other natal- horizontal or slopes very slightly dorsally. ids but curves posterodorsally from the al- The articular condyle is slightly higher than veolar margin to the tip of the coronoid at May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 737

an angle of about 70Њ. The presence of a well-developed coronoid with a distinct tri- angular dorsal portion is clearly a primitive feature of Primonatalus, probably re¯ecting its lesser degree of dorsal cranial ¯exion compared with modern natalids. Primonatalus has a distinct mandibular angle located on the ventral edge of the dentary anterior to the angular process and

directly below the dorsal tip of the coro- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 noid. There is a minor degree of dorsal ¯ex- ion of the ascending ramus posterior to the mandibular angle. A distinct mandibular angle is lacking in Natalus because of the sharp dorsal upturning of the mandible just posterior to m3. Chilonatalus has a weak mandibular angle located in a more dorsal position and farther posteriorly than in the fossil, posterior to the tip of the coronoid. Nyctiellus has a distinct mandibular angle located slightly posterior to the dorsalmost projection of the coronoid and dorsal to the ventral margin of the horizontal ramus. The angular process on the type dentary of Primonatalus is damaged, but it does not ¯are as strongly laterally as in other natal- ids, particularly Natalus. In lateral aspect, the base of the angular process of the fossil is situated at a level halfway between the alveolar margin and the ventral edge of the dentary and is also located farther anteriorly than in living natalids, approximately half- way between the articular condyle and the tip of the coronoid. The anteroventral po- sition of the angular process in Primona- talus also re¯ects the lesser dorsal ¯exion FIG. 6.ÐDistal portion of left humerus (UF of the ascending ramus in the fossil species. 108650) of Primonatalus prattae n. gen. et sp.: Viewed laterally, the angular process is lo- A) anterior view; B) medial view; C) posterior cated directly ventral to the articular con- view; D) lateral view; and E) distal view. Ab- dyle in Natalus and is positioned well dor- breviations: c ϭ capitulum; le ϭ lateral epicon- sal to the alveolar margin of the toothrow. dyle; lrc ϭ lateral ridge of capitulum; me ϭ epi- trochlea or medial epicondyle; mp ϭ medial pro- The angular process of Chilonatalus is lo- cess of epitrochlea; mrc ϭ medial ridge of ca- cated directly below, or very slightly ante- pitulum; rf ϭ radial fossa; sp ϭ spinous process rior to, the articular condyle and is at the of epitrochlea (ϭdistal spinous process); tr ϭ same level or slightly dorsal to the alveolar trochlea. margin of the toothrow. In Nyctiellus, the angular process is located anterior to the ar- ticular condyle and slightly ventral to the alveolar margin of the toothrow. In dorsal 738 JOURNAL OF MAMMALOGY Vol. 84, No. 2 and posterior views, the angular process of fossil species is clearly the most primitive Primonatalus does not ¯are laterally from member of the family in characters relating the vertical plane of the ascending ramus. to the dorsal ¯exion of the ascending ra- In Chilonatalus and Nyctiellus the angular mus. process ¯ares slightly, and in Natalus it The lower molars of Primonatalus are ¯ares widely. In lateral view, the angular most similar in size to teeth of Chilonatalus process of Primonatalus is curved and tumidifrons from the Bahamas. Measure- bears a small transverse crest along the dor- ments of P. prattae teeth are provided in sal edge of the tip. The angular process is Table 1. The cusps of the fossil teeth are more curved with a sharper hook in Nyc- less in¯ated or robust than in Natalus but Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 tiellus, strongly curved and hooked at the are more in¯ated than in Chilonatalus,in tip in Chilonatalus, and straight with a crest which the crests are more slender and the along the lateral edge of the tip in Natalus. cusps are sharp and spikelike. The talonids In dorsal view, the ascending ramus of are somewhat narrower in the fossil than in the dentary in Primonatalus is not inclined Chilonatalus. The metaconid and entoconid as far laterally as in Natalus and Chilona- are the same height on the m1 and m2 of talus. The coronoid process of the fossil is the type dentary of P. prattae, whereas on nearly vertical and located in the same m3 the metaconid is slightly taller than the plane as the labial edge of the toothrow. entoconid. The entoconid is slightly higher The coronoid process ¯ares much farther than the metaconid on m1±3 in all living laterally in Natalus and Chilonatalus. Nyc- natalids. The talonid notch is deeper in the tiellus is somewhat intermediate between fossil species owing to both the relatively Primonatalus and the 2 other living sub- higher metaconid and the more ventral con- genera in the location of the coronoid pro- nection of the entocristid on the trigonid be- cess. The relative position of the mandibu- low the metaconid. The cristid obliqua also lar foramen also re¯ects the degree of dor- connects nearer the base of the trigonid sal ¯exion of the braincase and ascending ventral to the protoconid in the fossils, thus ramus. The mandibular foramen is located forming a deeper notch than those in Na- at the same level as the alveolar margin in talus and Chilonatalus. The small carnas- Primonatalus and Nyctiellus and dorsal to sial-like notches in the cristid obliqua and the alveolar margin in Chilonatalus and Na- postcristid (Figs. 2A, 2B, and 3C) tend to talus. Among the 3 living subgenera of Na- form a small ``valley'' connecting the 2 talidae, Natalus shows the greatest degree crests in Primonatalus, but this feature is of dorsal ¯exion of the skull and conse- not as well developed as in Recent natalids. quently the strongest dorsal upturning of The ridge trending posteriorly from the the ascending ramus of the dentary. Nyc- cristid obliqua, just lingual to the ``valley'' tiellus shows the least dorsal ¯exion of the connecting the carnassial-like notches, is braincase and ascending ramus, whereas lacking in the fossils. Lower molars of Nyc- Chilonatalus is somewhat intermediate, al- tiellus are distinctly different from those of though more similar to Natalus. All living Primonatalus, Natalus, and Chilonatalus. natalids are more derived in this suite of Talonids are very broad and compose about characters than is Primonatalus. Compared two-thirds the length of the tooth on m1 and with living natalids, Primonatalus has a re- m2. In other natalids, the trigonid and tal- duced degree of dorsal upturning of the as- onid are approximately equal in size. Be- cending ramus of the dentary that is re¯ect- cause of the greater breadth of the talonids ed in the ventral position of the articular in Nyctiellus compared with other natalids, condyle, angular process, and mandibular the cristid obliqua forms a more acute angle foramen and the well-developed coronoid with the trigonid. The cristid obliqua attach- process with a distinct dorsal process. The es to the trigonid ventrally in Nyctiellus,as May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 739

TABLE 1.ÐMeasurements (in mm) of the teeth of Primonatalus prattae from the early Miocene (Hemingfordian) Thomas Farm Local Fauna, Florida.

Anteroposterior Transverse Trigonid Talonid Specimen number Tooth length width width width UF 121136 P4 1.09 1.09 UF 108639 M1 1.38 UF 108640 M1 1.25 1.28 UF 121137 M1 1.25 1.34 UF 121138 M1 1.28 1.44 UF 108638 M1 1.25 Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 UF 108638 M2 1.22 1.31 UF 121139 M3 1.13a 1.31a UF 108642 p3 1.16 0.50 UF 108641 (type) m1 1.25 0.75 0.78 UF 108641 (type) m2 1.19 0.78 0.78 UF 108641 (type) m3 1.19 0.72 0.63 UF 108644 m1 1.25 0.73 0.75 UF 121143 m1 1.22 0.69 0.75 UF 121143 m2 1.19 0.69 0.72 UF 108645 m3 1.06 0.59 0.56 a Estimated measurement of a broken tooth. in the fossil species. Both the paraconid and Nyctiellus than in Primonatalus, but it has metaconid are very small in Nyctiellus com- a distinct concavity between 2 crests pos- pared with other extant natalids and Pri- terior to the main cusp. monatalus. Overall, the lower molars of A 2nd paratype dentary of Primonatalus Primonatalus are most similar to those of prattae (UF 108647) is edentulous but con- Natalus. tains the alveoli for the lower incisors, ca- One of the paratype dentaries of Primo- nine, and p2±4 (Figs. 2F±H). This speci- natalus prattae (UF 108642) possesses the men has a large mental foramen located in p3 and the alveoli for p2 and p4 (Figs. 3A a deep concavity positioned high on the and 3B), providing additional information dentary between the roots of c and p2. In on the dental morphology not evident in the Natalus and Chilonatalus, the region be- type specimen. Like all living natalids, P. tween the alveoli of c and p2 is not as con- prattae has a single-rooted p2 and a double- cave as in the fossil, and the mental fora- rooted p3 and p4. The p3 of Primonatalus men is more ventrally located, about half- is relatively larger than the p3 of Natalus, way between the alveolar margin of the even though the species of Natalus are larg- toothrow and the ventral margin of the den- er than the fossil in most other dimensions. tary and ventral to the alveoli of both c and The p3 of the fossil species is comparative- p2. Nyctiellus has a smaller mental foramen ly longer and more laterally compressed in a shallower concavity. The anterior men- than in any living natalid. The p3 in the tal foramen is comparatively large in Pri- fossil possesses a distinct concavity be- monatalus and is located at the mandibular tween 2 vertical crests posterior to the pri- symphysis, immediately ventral to i1 and mary cusp. This concavity is present in extending posteriorly to the level of i2. The Chilonatalus, but absent in Natalus,in anterior mental foramen is very small and which the p3 is symmetrical anterior and inconspicuous in Natalus and Chilonatalus posterior to the main cusp with only 1 crest and extends posteriorly only to the middle descending posterior to the main cusp. The of i1. The anterior mental foramen is well p3 is comparatively shorter and broader in developed in Nyctiellus and extends poste- 740 JOURNAL OF MAMMALOGY Vol. 84, No. 2 riorly to i2. The mandibular symphysis in funnel-shaped cavity with large lachrymal the fossil is a short, inclined oval with a and infraorbital foramina separated by a nearly ¯at articular surface. The ventral thin ridge of bone. The overall size of the edge of the symphysis of Primonatalus pro- anterior orbital fossa and the 2 foramina are jects slightly below the lower margin of the reduced in N. tumidirostris because of the horizontal ramus. The symphysis in Chilon- in¯ation of the rostrum on the medial wall atalus and most Natalus (except N. stra- of the orbit. In Chilonatalus, the anterior mineus) has a narrower, more elongate oval orbital fossa is reduced in size compared outline and projects much more posteroven- with the fossil and most species of Natalus. trally than in Primonatalus, Nyctiellus, and The posterior opening of the infraorbital fo- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 Natalus stramineus. This process is small ramen in Chilonatalus is large and round in Nyctiellus, like the fossil. The fossil den- and widely separated from the small dorsal tary possesses alveoli for 3 lower incisors; lachrymal foramen. The anterior orbital fos- the ®rst 2 are coalesced and are separated sa and the openings of both foramina lo- from the 3rd by a tiny ridge. This situation cated within it are much reduced in Nyc- matches that in Chilonatalus, in which i1 tiellus owing to the greatly in¯ated rostrum. and i2 are close to one another but are sep- The anterior edge of the orbit in Chilona- arated from i3 by a minute diastema. In talus is inclined slightly anteriorly as in the Nyctiellus and Natalus the lower incisors fossil, and the maxilla below the orbits is are equally spaced. broader anteriorly and thinner posteriorly A partial left maxilla with M1 and M2 compared with Primonatalus. The anterior (UF 108638) is the only portion of the skull portion of the zygomatic arch above the M3 of Primonatalus preserved (Figs. 4B±D). is considerably thinner in Chilonatalus than This specimen retains the anteriormost re- in the fossil. The anterior base of the zy- gion of the zygomatic arch and the anterior gomatic arch begins at the anterior edge of and ventral margins of the orbit, as well as M3 in Chilonatalus but posterior to the M3 the posterior opening for the infraorbital fo- in the fossil. In Natalus, the anterior edge ramen. The zygomatic arch is ¯ared some- of the orbit is vertical, the anterior portion what laterally in the fossil and the anterior of the zygomatic arch is thinner and more edge of the muscle scar for the masseter parallel-sided than in Primonatalus, and the muscle is located at the level of the poste- masseteric muscle scar extends anteriorly to rior root of the M3. The anterior edge of about the middle of M3. In Nyctiellus, the the orbit in Primonatalus is located above anterior edge of the orbit is oriented at the mesostyle of M2 and is nearly vertical, about a 45Њ angle to the toothrow, the max- inclined only slightly anteriorly. The por- illa below the orbit is very deep, and the tion of the maxilla between the ventral edge masseteric muscle scar extends to the mid- of the orbit and the alveolar margin of the dle of the M3. The anterior process of the toothrow above M1±3 is relatively deep in zygomatic arch is more robust in Nyctiellus the fossil. Primonatalus possesses a deep, than in any other natalid. funnel-shaped fossa at the anterior edge of The upper molars of Primonatalus prat- the orbit that incorporates both the lachry- tae are similar to those of living members mal foramen and the posterior opening of of the Natalidae. Apparently, dental mor- the infraorbital foramen. The dorsal lach- phology of both the upper and lower molars rymal foramen and ventral infraorbital fo- has remained persistently conservative in ramen are both large in Primonatalus and this family because only minor changes are are separated only by a thin splint of bone. observed to have occurred since the early The anterior orbital fossa of Natalus (ex- Miocene. The M1 and M2 of Primonatalus cluding N. tumidirostris) is similar in mor- have a narrower trigon basin than in Recent phology to that of the fossil, being a deep natalids owing to the reduced talon and May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 741 metaconule in the terminology of Legendre rather cuspate, often giving a twinned ap- (1984). An ``incipient hypocone'' appears pearance to the mesostyle. In M2 the crest as a small cusp on the cingulum of the tal- is less prominent with no tendency to ap- on. This cusp is present on M1, but not on pear cuspate; instead, it is more smoothly M2, in Primonatalus, Natalus major, and curved inward toward the postparacrista. In N. stramineus but not in N. tumidirostris. Primonatalus this mesostylar crest is irreg- The M1 talon lacks the ``incipient hypo- ularly cristate in M1 and is abruptly ter- cone'' in Nyctiellus, Chilonatalus, and N. minated at its anterior end by a small cusp tumidirostris. The talon is very small on separating it from an anterior remnant of

M1 and virtually absent on M2. The par- the ectocingulum that extends backward Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 astyle is reduced on M2, and the protocone from the parastyle. Often a miniscule stylar is located closer to the paracone on both cuspule appears at the posterior end of this M1 and M2. The lingual cingulum is re- anterior ectocingulum, across the small gap duced in the fossil teeth, especially ventral from the mesostylar crest. In M2 of Pri- to the protocone. In Natalus, M1 is squar- monatalus the mesostylar crest is more ish, and M2 is more rectangular in occlusal smoothly cristate and more strongly curved outline. Both upper molars in Natalus are inward (lingually). A small posterior rem- squared off lingually because of the broad nant of the ectocingulum occurs in both M1 trigon basin and the well-developed talon. and M2 of Primonatalus, extending for- Both teeth are nearly as broad lingually as ward from the metastyle halfway across the labially. In Primonatalus, the 2 upper mo- posteroexternal valley and terminating in a lars are noticeably narrower lingually than very small stylar cuspule. Attributes of the labially. The M1 is broader lingually than mesostylar crest and stylar shelf of M1 and the M2 owing to the slight development of M2 in Chilonatalus and Natalus are similar a talon, whereas M2 is essentially triangu- to Primonatalus. Nyctiellus differs from the lar. In Chilonatalus, M2 is more similar to other genera in having mesostylar crests the fossil, with a very weak to nearly absent that are very small and weak and in lacking talon. The parastyle and talon are both pre- the remnants of the ectocingulum posterior sent on M2 in Nyctiellus but are very small. to the parastyle and anterior to the meta- The M1 and M2 are both narrow antero- style. M3 has a weak mesostylar crest in posteriorly in Nyctiellus as in the fossils Primonatalus and Chilonatalus. This ridge owing to the weakly developed talon. In is absent in M3 in Nyctiellus and Natalus. Primonatalus and Nyctiellus there is no Another dental synapomorphy uniting all connection between the posterior end of the Natalidae is the presence of a cingular (or postprotocrista and the metacingulum in ``stylar'') cusp labial to the primary cusp on M1 and M2. In Chilonatalus and Natalus P4 (Figs. 4E and 4F). This cusp is absent there is a weak-to-moderate connection of in the extinct nataloids Honrovits, Chad- these 2 structures. The lingual cingulum in ronycteris, and Chamtwaria. Stehlinia has M1 and M2 is complete in Natalus, inter- P4 with a strong rise in the labial cingulum, rupted around the base of the protocone in but no cingular cusp occurs in that genus Primonatalus and Chilonatalus, and more either. In Primonatalus the tip of the cin- widely interrupted in Nyctiellus. gular cusp is broken in the sole available A dental synapomorphy of natalids that specimen of P4 (UF 121136), but the cusp is absent in other extant and extinct nata- appears to have been low. In extant genera loids is the presence in M1 and M2 of a the labial cingular cusp is low in Nyctiellus, small curved crest extending forward from intermediate in height in Natalus, and high- the mesostyle into the anteroexternal valley est in Chilonatalus. The P4 in all natalids (Fig. 4B). In M1 this crest is straighter and is 3-rooted. The P4 of Primonatalus is dis- more prominent and has a tendency to be tinct from that of extant natalids in having 742 JOURNAL OF MAMMALOGY Vol. 84, No. 2 a narrow anterolabial projection in occlusal well as the lateral hook. The shallow view; the portion of P4 supported by the grooves along which the internal carotid ar- anterolabial root is separated from the lin- tery and stapedial artery run are apparent in gual portion of the tooth by an indentation the fossil periotics and follow paths across in the cingulum. The lingual moiety of P4 the cochlear surface that are identical to the (``talon'') is broadly rounded in Primona- paths in the modern natalids. talus. In the extant natalids there is no an- Five distal ends of the humerus referable terolingual indentation, and the anterolabial to the Natalidae are known from Thomas corner of the P4 is rather broadly quadrate. Farm, but no proximal humeri are present

The overall outline of P4 in occlusal view in the sample. The fossil humeri are readily Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 is roughly squarish in the labial half and identi®ed as natalids and separated from broadly rounded in the lingual half in Chi- those of vespertilionids by the laterally off- lonatalus and Natalus. Nyctiellus is distinct set distal articular surface and the broad, tri- from the others in being relatively shorter angular-shaped epitrochlea (ϭmedial epi- anteroposteriorly and in having a narrower condyle; Fig. 6). The following range of and more pointed lingual half, giving the measurements (mm) were made of the dis- tooth a roughly triangular occlusal outline. tal humeri of Primonatalus prattae from In Chilonatalus the rounded lingual portion Thomas Farm (UF 108648±108652): max- does not extend so far lingually as in Pri- imum distal width 2.8±2.9 (n ϭ 4); width monatalus so that the tooth is transversely of distal articular surface 1.9±2.0 (n ϭ 4); narrower. and shaft diameter 1.0±1.1 (n ϭ 4). The Three periotics of Primonatalus (Figs. distal humeri of P. prattae have a broad 5A±D) are virtually identical to the periot- triangular-shaped epitrochlea that has no ics of modern natalids. All are broken, but notch or concavity between the spinous 1 of the fossil periotics preserves 2 of the process of the epitrochlea (ϭdistal spinous semicircular canals. The cochlea of Pri- process) and the medial process of the epi- monatalus makes about 2¼ turns. The co- trochlea. The medial process is broad and chlea of Natalus stramineus appears to blunt in the fossil humeri and is barely sep- make about 2¼ turns, but in all other living arable from the large epitrochlea of which species of natalids the cochlea makes about it is a part. In Natalus the medial process 2½ turns. A small tubercle arises from the is better developed, primarily resulting smooth, rounded posteroventral surface of from the stronger notch located on the me- the cochlea of all natalids. This small tu- dial margin of the epitrochlea halfway be- bercle is situated just ventral to the fenestra tween the medial process and the spinous cochleae. It bears a tiny hooklike process at process. The medial process is better de- its lateral end that partly encircles the in- veloped in Chilonatalus, in which it is ternal carotid artery in life and contacts part prominent and elongated. The medial pro- of the posterior end of the stylohyal. The cess is separated from the spinous process tubercle and hook are transversely short in by a deep rounded notch in the epitrochlea Primonatalus. The transverse width of this that forms nearly a right angle. The entire tubercle varies in Natalus;inN. stramineus epitrochlea is greatly reduced in Nyctiellus, it is short and very similar to that in Pri- and the medial process is absent. The spi- monatalus,inN. tumidirostris it is longer nous process is small in Primonatalus, pro- and rather more ¯attened on its posteroven- jects slightly laterally, and is somewhat tral surface, and in N. major it is still longer pointed distally. In the fossils this process and ¯attened. In Chilonatalus the tubercle projects distally only to the level of the dis- forms a taller, larger hook. In Nyctiellus the tal edge of the trochlea or slightly distal to tubercle is transversely wide and distinctive it in several specimens. The spinous process in possessing a medial pointed projection as is separated from the medial edge of the May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 743 trochlea by a narrow but rather deep notch. tween it and the medial surface of the ca- A small spinous process is also character- pitulum. The medial edge of the trochlea on istic of Natalus, but this process is con- the posterior surface is a well-developed nected to the trochlea by a ridge, and it thus ridge extending proximally to the ¯ared lacks the distinct notch found in the fossil part of the shaft in the fossils and Natalus. humeri. Because this notch is lacking in This ridge is weakly developed in Chilo- Natalus, the spinous process does not ap- natalus and absent in Nyctiellus. In lateral pear to be as prominent as in Primonatalus, view, the articular surface of the lateral ca- even though it projects the same distance pitulum of the Primonatalus humeri is nar- distally in both genera. The spinous process row anteroposteriorly and is ¯attened dis- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 is very well developed in Chilonatalus, ex- tally, and the lateral surface of the capitu- tending noticeably distal to the medial edge lum has a deep elliptically shaped concavity of the trochlea and separated from it by a that opens proximally. In other natalids, the deep notch. The spinous process of Chilo- articular surface of the lateral capitulum is natalus also bears a sharp laterally project- more rounded, and the lateral concavity is ing process. Nyctiellus has a very large spi- present but shallower. nous process that extends far distal to the Two proximal radii from Thomas Farm articular surface. In characters of the spi- are very similar in size and morphology to nous process, Primonatalus is intermediate the proximal radius of extant natalids. The between Natalus, in which this process is only difference is that the process extending small, short, blunt distally, and connected proximal to the articular surface for the hu- to the medial edge of the trochlea, and Chi- merus is more rounded in the fossils. This lonatalus, in which the process is very large proximal process is more pointed and tri- and broad, extends well distal to the artic- angular-shaped in living natalids. Despite ular surface, projects noticeably laterally, the slight difference in the proximal pro- and is separated from the medial edge of cess, the fossil radii are more similar to the the trochlea by a deep notch. radius of natalids than to that of any other In anterior aspect, the medial ridge of the bat present at Thomas Farm. Four distal ra- capitulum is bulbous and almost spherical dii from Thomas Farm are essentially iden- in Primonatalus and Natalus. In these 2 tical to the distal radius of living natalids. genera, this rounded portion of the capitu- As with most other elements of the Thomas lum is separated from the lateral ridge of Farm natalid, the fossil radii are most sim- the capitulum and the trochlea by relatively ilar in size to the radius of Chilonatalus deep, broadly rounded grooves. The medial tumidifrons. ridge of the capitulum is more ¯attened in The proximal end of a chiropteran radius Chilonatalus, and the grooves separating it (UF 121723) from the Oligocene I-75 Local from the trochlea and lateral capitulum are Fauna is very similar to the 2 proximal radii shallow. The medial ridge of the capitulum from Thomas Farm in its morphology, is reduced in Nyctiellus, and the entire distal small size, and delicate shaft. The tip of the articular surface is relatively narrower com- proximal process is slightly damaged in the pared with all other natalids. I-75 fossil; thus, it is not possible to deter- In posterior aspect, the proximal margin mine whether this specimen differs from of the lateral ridge of the capitulum is ¯at- the living natalids in the same feature as tened in the fossil, Chilonatalus, and Nyc- does Primonatalus. Although diagnostic at tiellus and is separated from the medial ca- the family level, the proximal end of the pitulum by a shallow groove. In Natalus the radius is not useful for distinguishing dorsal surface of the lateral capitulum is ex- among species in the Natalidae, except for panded into a large rounded process; con- size. The I-75 radius is referred to the Na- sequently, there is a deep rounded notch be- talidae, but the genus and species are in- 744 JOURNAL OF MAMMALOGY Vol. 84, No. 2 determinate. The identi®cation of this spec- imen as a member of the Natalidae is sig- ni®cant because it represents the oldest known record for the family, some 12 mil- lion years older than Primonatalus from Thomas Farm. The distal end of a femur of Primona- talus from Thomas Farm is similar in shaft FIG. 7.ÐPotential relationships within the diameter and distal articular width to the family Natalidae, based on an analysis of 50 femora of Chilonatalus micropus and Na- morphological characters of the teeth and skel- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 talus stramineus. No characters distinctive eton. Topology represents the single most par- at the genus or species level were found in simonious tree from a branch-and-bound search the fossil or in the distal femur in living (tree length 82 steps; consistency index ϭ 0.7439; retention index ϭ 0.6769). Numbers natalids. The relative length of the femur above branches are decay values (Bremer sup- varies signi®cantly among extant natalids, port values); numbers below branches are boot- but it is not possible to determine length of strap values representing the percentage of the femur from the incomplete fossil. The 10,000 bootstrap replicates in which each clade femur is longer than the humerus (about appeared. 105%) in Chilonatalus tumidifrons, slightly shorter than the humerus (about 90±95%) in C. micropus and Natalus, and much member of the family and is distinct from shorter than the humerus (about 80%) in the other natalids, but its membership in the Nyctiellus. The variation in relative length family is strongly supported. These results of the humerus and femur among living na- are remarkably concordant with aspects of talids indicates differences in size and, pos- the traditional classi®cation scheme for the sibly, function of the hind limb and tail living natalids and suggest that there may membrane. be a phylogenetic signal in the dental±cra- nial±postcranial morphology in this family. PHYLOGENETIC ANALYSIS All living members of the Natalidae typ- We ®nd strong support for the monophy- ically are placed in the genus Natalus ly of the family Natalidae (sensu stricto), (Koopman 1993). We suggest, however, including Primonatalus prattae (Fig. 7). that the 6 extant species should be separated Among the extant taxa, the 3 species of Na- into 3 genera, Natalus (including the spe- talus are monophyletic, and this clade is cies N. major, N. stramineus, and N. tumi- strongly supported. Within this clade, Na- dirostris), Chilonatalus (including the spe- talus stramineus and Natalus tumidirostris cies C. micropus and C. tumidifrons), and are sister taxa (not surprising, because the Nyctiellus (including the species N. lepi- 2 species are dif®cult to tell apart in hand dus), each previously recognized as a sub- and are very similar in skeletal morpholo- genus of Natalus (Dalquest 1950; Goodwin gy), with strong support for the inclusion of 1959; Koopman 1993). Most workers con- Natalus major. The 2 species of Chilona- sider N. major to be a junior synonym of talus are monophyletic, supporting the no- N. stramineus (Koopman 1993; Silva Ta- tion that Chilonatalus is distinct from Na- boada 1979; Varona 1974), but Morgan talus. Chilonatalus and Natalus cluster to- (1989b) noted several morphological differ- gether, but support for this clade is moder- ences between the 2 species that he thought ate. There is moderate support for justi®ed recognition of N. major as a dis- distinction of the traditional subgenera Nyc- tinct species, including large size (there is tiellus, Chilonatalus, and Natalus. The fos- no overlap in measurements with the small- sil Primonatalus is the most primitive er N. stramineus); more in¯ated braincase May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 745 that meets the rostrum at a steeper angle; tinguish them from other chiropteran fami- more strongly constricted interorbital re- lies. gion; stronger sagittal crest; and less in¯at- The derived features of the Natalidae are ed rostrum that is more tapered anteriorly, as follows (characters in our data set are constricting the size of the narial opening. identi®ed by their character number in pa- Among the 3 living genera, Nyctiellus is rentheses followed by the character state in the most distinct. We recognize Nyctiellus brackets, from Appendix II): elongated, as a separate genus on the basis of numer- dorsoventrally ¯attened rostrum; reduced ous cranial and dental characters that differ orbit with anterior edge located far poste- substantially from other natalids (character rior above M2; deep funnel-shaped fossa at Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 numbers [character states] from Appendix anterior edge of orbit enclosing lachrymal II), including small size; broader, deeper, foramen and posterior opening of infraor- and more in¯ated rostrum (character bital foramen (39[1]); greatly elongated in- 31 [state 1]); lower, less in¯ated braincase fraorbital canal originating above M2 and (25 [1]) and less dorsally upturned braincase opening anteriorly above P3 (38[1]); elon- (26 [0]); deeper, more robust zygomatic arch gated styliform process of ectotympanic (34 [1]); deep midline (unpaired) basisphe- that extends anteromedially and connects noidal pit (36 [2]); in¯ated auditory bullae with pterygoid process (33[1]); presence on (37 [1]); cleft between premaxillae reduced P4 of cingular or stylar cusp labial to pri- so that I1s nearly meet at midline (35 [1]); mary cusp (14[1]); presence on M1 and M2 reduced upper canines (27 [1]); and greatly of small curved crest extending anteriorly reduced P2 (29 [1]) and p2 (50 [1]). Chilon- from mesostyle into anteroexternal valley atalus is distinguished by having P4 with a (17[1]); manubrium (presternum) very strong labial cingular cusp (14 [2]) and a broad with strong posteriorly oriented me- highly derived degree of fusion in the axial dian keel (43[1]); last thoracic and all lum- skeleton, with only 1 free (unfused) poste- bar vertebrae except the last 1 or 2 fused rior lumbar vertebrae (42 [2]) and ribs that into rigid, laterally compressed structure are greatly coalesced with each other and with prominent dorsal and ventral ridges with the sternum (44 [2]). Such axial fusion (41[1]); exceptionally elongated caudal ver- is found in varying degrees in several fam- tebrae 3 through 6; femur and tibia ex- ilies of bats and is probably related to the tremely elongated and slender; proximal energetics of echolocation (Lancaster et al. end of femur strongly bent laterally (46[1]); 1995, 2001). Natalus sensu stricto is distin- proximal end of femur with greater and guished by having a straight angular process lesser trochanters reduced (48[1]); and pres- (4 [0]) that ¯ares strongly outward (5 [2]) ence of natalid organ, a subcutaneous and a distal spinous process on the humerus glandlike organ on forehead and muzzle of that is not separated from the medial edge males. Simmons (1998) listed the presence of the trochlea by a notch (23 [1]). of the natalid organ as an apomorphy di- The fossil material of Primonatalus prat- agnosing the Natalidae. tae is not nearly so complete as that avail- able for the 6 living species of natalids. Al- DISCUSSION though many of the derived characters of Five extinct genera have been referred to the Natalidae are unknown in Primonatal- the Natalidae, including Ageina from the us, enough of the cranial and dental anat- early Eocene of Europe (Russell et al. omy of Primonatalus is preserved to con- 1973), Stehlinia from the Eocene and Oli- ®rm that this taxon is unquestionably refer- gocene of Europe (Sige 1974), Honrovits able to the Natalidae. The combination of from the early Eocene of Wyoming (Beard the derived character states listed below et al. 1992), Chadronycteris from the late unite all members of the Natalidae and dis- Eocene of Nebraska (Ostrander 1983), and 746 JOURNAL OF MAMMALOGY Vol. 84, No. 2

Chamtwaria from the Miocene of Africa has been followed by most subsequent chi- (Butler 1984). On considering these 5 ex- ropteran systematists. An analysis of chi- tinct genera to be members of the Natalidae, ropteran dentitions supported Miller's sug- Beard et al. (1992) followed an expanded gestion of a close relationship between the de®nition of the family proposed by Van Natalidae, Furipteridae, and Thyropteridae Valen (1979), which included, in addition (Slaughter 1970). Van Valen (1979) ex- to the Natalidae sensu stricto, the Furipter- panded the concept of the Natalidae by in- idae, Thyropteridae, Myzopodidae, and cluding not only the Furipteridae and Thy- Kerivoulidae as subfamilies. Excluding the ropteridae but also the Myzopodidae and

Kerivoulidae, this is similar to the concept Kerivoulidae as subfamilies. Van Valen's Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 of the superfamily Nataloidea proposed by (1979) expanded concept of the Natalidae Simmons (1998) and Simmons and Geisler was supposedly based on shared derived (1998), consisting of the families Natalidae, characters; however, the supporting data for Furipteridae, Thyropteridae, and Myzopod- his phylogenetic hypothesis were not pro- idae. These authors considered the Kerivou- vided, and most subsequent authors have lidae to be a subfamily of the Vespertilion- not followed his arrangement. Recently, idae. McKenna and Bell (1997) placed Van Den Bussche and Hoofer (2001) pro- Honrovits in the Natalidae sensu stricto, vided molecular evidence that Nataloidea is Stehlinia and Chamtwaria in the Vesperti- not a monophyletic lineage. Their mito- lionidae, and Ageina and Chadronycteris in chondrial DNA sequence data suggested in- Microchiroptera incertae sedis. Simmons stead that Myzopodidae represents a basal and Geisler (1998) regarded Ageina, Chad- microchiropteran lineage and that Furipter- ronycteris, Honrovits, and Stehlinia as in- idae, Thyropteridae, and Natalidae are suc- certae sedis within the Nataloidea (they cessive out-groups to the Noctilionoidea. considered only Eocene forms and thus did However, subsequent to that analysis, Hoof- not mention the Miocene Chamtwaria). All er et al. (in press) extended their analysis 5 extinct Tertiary genera referred to the Na- by examining DNA sequences from a nu- talidae or Nataloidea by previous authors, clear gene in all families of Yangochirop- including Ageina, Chadronycteris, Cham- tera. Their goal speci®cally was to reconcile twaria, Honrovits, and Stehlinia, fall out- controversies surrounding 5 families whose side the strict de®nition of the Natalidae, as phylogenetic positions have been unstable recognized in this study (i.e., they lack the (Furipteridae, Mystacinidae, Myzopodidae, derived features of the Natalidae listed Natalidae, and Thyropteridae). On combin- above). Only the 6 extant species previous- ing their new nuclear sequence data with ly placed in the genus Natalus and the Mio- the previous mitochondrial data, Hoofer et cene Primonatalus, described in this study, al. (in press) found support for an associa- belong to the Natalidae sensu stricto. tion of Natalidae with Vespertilionidae and Many authors have discussed the close Molossidae in the Vespertilionoidea (but relationship between the Natalidae, Furip- Furipteridae and Thyropteridae remained teridae, and Thyropteridae (Dalquest 1950; with Noctilionoidea). Accordingly, and Miller 1899, 1907; Simmons and Geisler based on our results, we adhere to the con- 1998; Slaughter 1970). Miller (1899) orig- ventional arrangement of the Natalidae, in- inally placed the Furipteridae and Thyrop- cluding only 3 living genera (or subgenera) teridae within the Natalidae, but in his clas- Natalus, Chilonatalus, and Nyctiellus, along sic study of the families and genera of bats, with the extinct Miocene genus Primona- Miller (1907) assigned the genera Furipte- talus. rus and Amorphochilus to the Furipteridae The presence of natalids in the middle and Thyroptera to the Thyropteridae. Mill- Cenozoic of Florida provides important er's (1907) arrangement of these 3 families new information pertaining to the evolu- May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 747 tionary history of this small family of bats. Natalids are widespread throughout tropical Before the discovery of Primonatalus in Middle America (Hall 1981), but they have Florida, the Natalidae sensu stricto had no a restricted distribution in South America, pre±late Pleistocene fossil record. The fos- where they occur primarily in the Carib- sils from the Oligocene I-75 Local Fauna bean and Atlantic coastal regions in the and the early Miocene Thomas Farm Local northern and northeastern portions of the Fauna extend the record of the Natalidae continent (Koopman 1982). Natalus stra- back nearly 30 million years in North mineus is the most widely distributed spe- America. Furthermore, the Florida fossils cies in the family, occurring in the mainland occur in what is now the Nearctic region, Neotropics from northern Mexico, through- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 whereas no species in the Natalidae are cur- out Middle America, and as far south as rently found outside the Neotropical region. Brazil, with a disjunct population in the Late Pleistocene fossils representing extant northern Lesser Antilles (Hall 1981; Koop- species of Natalidae are known from many man 1982, 1989). Natalus tumidirostris is islands in the West Indies (Morgan 2001) found in northern Colombia and Venezuela and from Brazil (Czaplewski and Cartelle and on the southern Caribbean islands of 1998). Tertiary fossils of natalids are un- Trinidad, Margarita, and CuracËao (Koop- known from South America, although the man 1982). Although these islands are lo- fossil record of bats on this continent is cated in the West Indies, they were exclud- very sparse (Czaplewski 1997). Of the 2 ed from the West Indian subregion by other Neotropical families supposedly Koopman (1958, 1989) because they lack closely related to the Natalidae, the Furip- endemic species of Antillean bats. teridae occur in Pleistocene deposits in Bra- All living natalids are obligate cave zil (Czaplewski and Cartelle 1998) and dwellers. In the West Indies, most species Peru (Morgan and Czaplewski 1999) and in this family roost in chambers deep within the Thyropteridae are known from the mid- large caves that are characterized by a sta- dle Miocene of Colombia (Czaplewski ble microenvironment with high tempera- 1997). ture and relative humidity (Goodwin 1970; The Natalidae are currently restricted to Silva Taboada 1979). The roosting ecology the Neotropical region, although several of natalids suggests that their present (and species occur at the northern limits of the presumably past) distribution is limited by Neotropics in northern Mexico, the Baha- the availability of extensive cave systems mas, and Cuba. Natalus stramineus is (Morgan 2001). known from the states of Sonora and Nuevo The Tertiary fossil record of the Natali- Leon in northern Mexico (Hall 1981), Chi- dae is limited to the early Oligocene and lonatalus tumidifrons occurs on Abaco and early Miocene of Florida. The family pre- Andros in the Bahamas, and C. micropus sumably survived in tropical North Ameri- and Nyctiellus lepidus are found in Cuba ca (i.e., Middle America) between the early (Morgan 2001). The family attains its great- Miocene and the late Pleistocene, although est species richness in the West Indies, natalids have no fossil record during that where all 3 genera and 6 species of the fam- time period. The presence in the West In- ily occur. Two of the genera (Chilonatalus dies of 2 endemic genera and 4 endemic and Nyctiellus) and 4 of the species (Na- species of natalids suggests this group prob- talus major from Jamaica and Hispaniola; ably reached the Antilles by overwater dis- Chilonatalus micropus from Cuba, Jamaica, persal from tropical North America early in and Hispaniola; C. tumidifrons from the Ba- their evolutionary history, presumably in hamas; and Nyctiellus lepidus from Cuba the Oligocene or Miocene. Conversely, in and the Bahamas) are endemic to the West South America, the Natalidae have a mar- Indian subregion of the Neotropical region. ginal distribution, limited species richness 748 JOURNAL OF MAMMALOGY Vol. 84, No. 2

(2 species of Natalus, 1 of which is endem- cene bats from cave deposits in Bahia, Brazil. Jour- nal of Mammalogy 79:784±803. ic), and lack a Tertiary fossil record. These CZAPLEWSKI,N.J.,AND G. S. MORGAN. 2000. A new factors suggest that natalids may not have vespertilionid bat (Mammalia: Chiroptera) from the entered South America until comparatively early Miocene (Hemingfordian) of Florida, USA. Journal of Vertebrate Paleontology 20:736±742. late in time, perhaps in the Pliocene after DALQUEST, W. W. 1950. The genera of the chiropteran the formation of the Panamanian Isthmus family Natalidae. Journal of Mammalogy 31:436± and the beginning of the Great American 443. EMRY, R. J., P. R. BJORK, AND L. S. RUSSELL. 1987. Faunal Interchange. The Chadronian, Orellan, and Whitneyan North American land mammal ages. Pp. 118±152 in Ce-

ACKNOWLEDGMENTS nozoic mammals of North America: geochronology Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 and biostratigraphy (M. O. Woodburne, ed.). Uni- We thank A. E. Pratt of Georgia Southern versity of California Press, Berkeley. University and A. R. Poyer of the Florida Mu- ESTES, R. 1963. Early Miocene salamanders and liz- seum of Natural History for their help in the ards from Florida. Quarterly Journal of the Florida ®eld and with screenwashing and sorting matrix. Academy of Sciences 26:234±256. Â All the Miocene natalid fossils described in this FORSTEN, A. 1975. The fossil horses of the Texas Gulf Coastal Plain: a revision. Texas Memorial Museum, article were recovered through their efforts. We Pearce-Sellards Series 22:1±86. thank S. D. Webb and B. J. MacFadden of the GOODWIN, G. G. 1959. Bats of the subgenus Natalus. Florida Museum of Natural History for loans American Museum Novitates 1977:1±22. and permission to study the Thomas Farm and GOODWIN, R. E. 1970. The ecology of Jamaican bats. Journal of Mammalogy 51:571±579. I-75 bats. For access to Recent comparative HALL, E. R. 1981. The mammals of North America. specimens, we thank L. Wilkins of the Florida 2nd ed. John Wiley & Sons, Inc., New York 1:1± Museum of Natural History, L. K. Gordon of the 600 ϩ 90. National Museum of Natural History, Washing- HAYES, F. G. 2000. The Brooksville 2 Local Fauna ton, D.C., N. B. Simmons of the American Mu- (Arikareean, latest Oligocene): Hernando County, Florida. Bulletin of the Florida Museum of Natural seum of Natural History, New York, and W. L. History 43:1±47. Gannon of the Museum of Southwestern Biol- HOLMAN, J. A. 1965. Early Miocene anurans from ogy, University of New Mexico, Albuquerque. Florida. Quarterly Journal of the Florida Academy We greatly appreciate the advice of R. L. Nydam of Sciences 28:68±82. HOLMAN, J. A. 1967. Additional Miocene anurans from and N. B. Simmons on the parsimony analyses. Florida. Quarterly Journal of the Florida Academy N. B. Simmons and an anonymous reviewer pro- of Sciences 30:121±140. vided helpful comments on the manuscript. HOLMAN, J. A. 1999. Early Oligocene (Whitneyan) Funding for this study was provided by a grant snakes from Florida (USA), the second oldest col- from the National Science Foundation (DEB ubrid snakes in North America. Acta Zoologica Cra- coviensia 42:447±454. 9981512) to N. J. Czaplewski and G. S. Morgan. HOOFER, S. R., S. A. REEDER,E.W.HANSEN, AND R. A. VAN DEN BUSSCHE. In press. Molecular phylo- LITERATURE CITED genetics and taxonomic review of noctilionoid and AUFFENBERG, W. 1963. The fossil snakes of Florida. vespertilionoid bats (Chiroptera: Yangochiroptera). Tulane Studies in Zoology 10:131±216. Journal of Mammalogy. Â BEARD, K. C., B. SIGEÂ , AND L. KRISHTALKA. 1992. A HORACEK, I. In press. On the early history of vesper- primitive vespertilionoid bat from the early Eocene tilionid bats in Europe: the lower Miocene record of central Wyoming. Comptes Rendus des SeÂances from the Bohemian Massif. Myotis. de l'AcadeÂmie des Sciences Paris 314:735±741. HULBERT, R. C., JR. 1984. Paleoecology and popula- BLACK, C. C. 1963. Miocene rodents from the Thomas tion dynamics of the early Miocene (Hemingfordian) Farm Local Fauna, Florida. Bulletin of the Museum horse Parahippus leonensis from the Thomas Farm of Comparative Zoology 128:483±501. Site, Florida. Journal of Vertebrate Paleontology 4: BRODKORB, P. 1956. Two new birds from the Miocene 547±558. of Florida. Condor 58:367±370. KOOPMAN, K. F. 1958. Land bridges and ecology in bat BUTLER, P. M. 1984. Macroscelidea, Insectivora, and distribution on islands off the northern coast of Chiroptera from the Miocene of East Africa. Pa- South America. Evolution 12:429±439. laeovertebrata 14:117±200. KOOPMAN, K. F. 1982. Biogeography of the bats of CZAPLEWSKI, N. J. 1997. Chiroptera. Pp. 410±431 in South America. Pp. 273±302 in Mammalian biology Vertebrate paleontology in the Neotropics: the Mio- in South America (M. A. Mares and H. H. Geno- cene fauna of La Venta, Colombia (R. F. Kay, R. H. ways, eds.). Special Publication Series, Pymatuning Madden, R. L. Cifelli, and J. J. Flynn, eds.). Smith- Laboratory of Ecology, University of Pittsburgh, sonian Institution Press, Washington, D.C. Pittsburgh, Pennsylvania 6:1±539. CZAPLEWSKI,N.J.,AND C. CARTELLE. 1998. Pleisto- KOOPMAN, K. F. 1989. A review and analysis of the May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 749

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ropteran dentitions. Pp. 51±83 in About bats: a chi- Submitted 13 August 2001. Accepted 19 August 2002. ropteran symposium (B. H. Slaughter and D. W. Walton, eds.). Southern Methodist University Press, Associate Editor was Thomas J. O'Shea. Dallas, Texas. SMITH, J. D. 1972. Systematics of the chiropteran fam- ily Mormoopidae. University of Kansas Museum of APPENDIX I Natural History, Miscellaneous Publication 56:1± 132. Modern comparative specimens of natalids STANEÏ K, V. J. 1933. K topogra®cke a srovnaÂvacõ ana- and other nataloids are from the mammal col- tomii sluchoveÂho orgaÂnu nasÏich chiropter (Toward a lections of the Florida Museum of Natural His- topographic and comparative anatomy of the audi- tory, University of Florida (UF), Gainesville, Ï tory organ of bats). NaÂkladem Ceske Akademie VeÏd Florida; United States National Museum of Nat- a UmeÏni, Prague, Czech Republic (in Czech). Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 ural History (USNM), Smithsonian Institution, SWOFFORD, D. L. 2000. PAUP: phylogenetic analysis using parsimony. Version 4.02b. Sinauer Associates, Washington, D.C.; Museum of Southwestern Bi- Inc., Publishers, Sunderland, Massachusetts. ology (MSB), University of New Mexico, Al- TEDFORD, R. H., AND C. D. FRAILEY. 1976. Review of buquerque, New Mexico; Field Museum of Nat- some Carnivora (Mammalia) from the Thomas Farm ural History (FMNH), Chicago, Illinois; and Local Fauna (Hemingfordian, Gilchrist County, Florida). American Museum Novitates 2610:1±9. Oklahoma Museum of Natural History TEDFORD, R. H., ET AL. 1987. Faunal succession and (OMNH), Norman, Oklahoma. biochronology of the Arikareean through Hemphil- Recent comparative specimens examined.Ð lian interval (late Oligocene through earliest Plio- Natalidae: Natalus (Natalus) major, UF 13691 cene epochs) in North America. Pp. 153±210 in Ce- (&), UF 13692 ((), St. Clair Cave, Jamaica; Na- nozoic mammals of North America: geochronology talus (Natalus) stramineus, UF 9847 ((), Rio and biostratigraphy (M. O. Woodburne, ed.). Uni- versity of California Press, Berkeley. Frio Caves, Belize, MSB 53771 ((), MSB TEDFORD, R. H., J. B. SWINEHART,C.C.SWISHER III, 53776 (&), La Aduana, Sonora, Mexico; Natalus D. R. PROTHERO,S.A.KING, AND T. E. TIERNEY. (Natalus) tumidirostris, UF 22512 (&), Pueblo 1996. The Whitneyan-Arikareean transition in the Nuevo, Venezuela, USNM 441670 ((), USNM High Plains. Pp. 312±334 in The terrestrial Eocene- 441671 ((), Miranda, Venezuela; Natalus (Chi- Oligocene transition in North America (D. R. Proth- ero and R. J. Emry, eds.). Cambridge University lonatalus) micropus, UF 13698 (&), UF 19076 Press, Cambridge, United Kingdom. ((), St. Clair Cave, Jamaica; Natalus (Chilon- VAN DEN BUSSCHE, R. A., AND S. R. HOOFER. 2001. atalus) tumidifrons, UF 24850 ((), UF 24853 Evaluating monophyly of Nataloidea (Chiroptera) ((), Eight Mile Cave, Abaco, Bahamas; Natalus with mitochondrial DNA sequences. Journal of (Nyctiellus) lepidus, UF 12835 ((), Pinder's Set- Mammalogy 82:320±327. VAN VALEN, L. 1979. The evolution of bats. Evolu- tlement, Long Island, Bahamas, USNM 300528 tionary Theory 4:103±121. ((), Havana, Cuba. Furipteridae: Furipterus VARONA, L. S. 1974. CataÂlogo de los mamõÂferos vi- horrens, USNM 315737 ((), Changuinola, Pan- vientes y extinguidos de las Antillas. Academia de ama, USNM 405785 ((), T. F. Amazonas, Ve- Ciencias de Cuba, La Habana, Cuba. nezuela, USNM 549507 ((), Rio Xingu, Brazil; VAUGHAN, T. A. 1959. Functional morphology of three bats: Eumops, Myotis, and Macrotus. University of Amorphochilus schnablii, USNM 152261 (sex Kansas Publications, Museum of Natural History 12: unknown), Dintorni di Lima, Peru, USNM 1±153. 269981 (&), Arequipa, Peru. Thyropteridae: WEBB, S. D. 1981. The Thomas Farm fossil site. Plas- Thyroptera discifera, USNM 102922 ((), ter Jacket 37:6±25. USNM 105422 (&), San Julian, Venezuela, WHITE, T. E. 1942. The lower Miocene mammal fauna of Florida. Bulletin of the Museum of Comparative USNM 457965 (&), Barro Colorado Island, Pan- Zoology 92:1±49. ama; Thyroptera tricolor, USNM 281200 ((), WIBLE, J. R., G. W. ROUGIER,M.J.NOVACEK, AND M. Colonia Agricola, Colombia, USNM 564337 C. MCKENNA. 2001. Earliest eutherian ear region: a (&), Beni, Bolivia, USNM 582082 ((), Barro petrosal referred to Prokennalestes from the early Cretaceous of Mongolia. American Museum Novi- Colorado Island, Panama; Thyroptera lavali, tates 3322:1±44. FMNH 89118 ((), FMNH 89120 (&), Loreto, WOODBURNE,M.O.,AND C. C. SWISHER III. 1995. Peru. Vespertilionidae: Kerivoula (Phoniscus) Land mammal high-resolution geochronology, inter- jagori, USNM 155837 (&), Java, Indonesia; continental overland dispersals, sea level, climate, Kerivoula hardwickii, USNM 17909 (&), para- and vicariance. Geochronology, time scales, and global stratigraphic correlation. Society of Economic type, Cambodia; Kerivoula picta, USNM Paleontologists and Mineralogists, Special Publica- 356429 (&), Da Nang, Viet Nam; Myotis luci- tion 54:336±364. fugus, OMNH uncatalogued (sex unknown), May 2003 MORGAN AND CZAPLEWSKIÐNEW MIOCENE NATALID FROM FLORIDA 751

Killdeer, North Dakota. Specimens consist of upper molars: (0) normal; (1) extended antero- skulls and mandibles, most with complete post- medially as a curved mesostylar crest. 18. Ec- cranial skeletons. tocingulum of upper molars posterior to para- style and anterior to metastyle: (0) present but APPENDIX II weak; (1) absent. 19. M1 postprotocrista: (0) ex- tends only to ``metaconule'' region (i.e., a true Characters used in phylogenetic analysis, metaconule is absent), does not reach metacone with the states for each character. nor metacingulum; (1) becomes con¯uent with 1. Coronoid process of dentary, height rela- tive to articular condyle: (0) higher than con- distal cingulum (metacingulum); (2) joins lin- dyle; (1) same level as condyle; (2) lower than gual base of metacone; (3) extends as ridge onto Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 condyle. 2. Angular process of dentary in lateral relatively well-developed talon. 20. Posteroven- view projects: (0) at or below level of occlusal tral tubercle of cochlea: (0) absent; (1) present, plane of toothrow; (1) above level of occlusal with lateral hook; (2) present, with lateral and plane. 3. Base of angular process of dentary in medial projections; (3) present but weakly de- lateral view is situated: (0) ventral to condyle; veloped. 21. Cochlea: (0) makes about 2¼ turns; (1) at a level between condyle and tip of coro- (1) makes about 2½ turns; (2) makes more than noid process. 4. Shape of angular process: (0) 2½ turns. 22. Epitrochlea of humerus: (0) with straight; (1) slightly curved; (2) strongly hooked. no medial process; (1) with weak medial pro- 5. Angular process in dorsal or posterior view: cess; (2) with prominent medial process. 23. (0) in same vertical plane as condyle and as- Distal spinous process of humerus: (0) separated cending ramus; (1) slightly ¯ared laterally; (2) from medial edge of trochlea by small notch; (1) strongly ¯ared laterally. 6. Position of mandib- no notch. 24. Shape of spinous process of hu- ular foramen: (0) opens above level of alveolar merus: (0) short with blunt tip; (1) intermediate margin of lower toothrow; (1) opens at same in length with lateral projection at tip; (2) long level as alveolar margin. 7. Size of anterior men- and curved laterad; (3) long and nearly straight. tal foramen (ventral to lower incisors): (0) large, 25. Braincase shape: (0) in¯ated; (1) relatively extending distally beneath i2; (1) small, extend- unin¯ated. 26. Cranial ¯exion (rostral±forehead ing distally beneath i1. 8. Spacing of lower in- angle): (0) forehead not sharply elevated above cisor alveoli: (0) alveoli of i1 and i2 nearly co- rostrum (Ͼ135Њ); (1) more sharply elevated alesced, small gap between i2 and i3; (1) alveoli above rostrum (Ͻ135Њ). 27. Upper canine height of i1, i2, and i3 evenly spaced. 9. Shape of ar- relative to P4: (0) taller than P4; (1) about equal ticular surface of mandibular symphysis in me- to P4. 28. Lower canine height relative to p4: dial view: (0) rounded oval with small ventral (0) taller than p4; (1) about equal to p4. 29. P2 projection; (1) narrow, elongate oval with long size: (0) normal; (1) greatly reduced. 30. Ros- posteroventral projection; (2) elongate oval with trum length: (0) longer than or equal to brain- no ventral projection. 10. Crown of p3: (0) hav- case; (1) shorter than braincase. 31. Sides of ros- ing 2 posterior crests separated by a concavity; trum: (0) not markedly in¯ated; (1) markedly in- (1) having 1 posterior or lingual crest. 11. Oc- ¯ated. 32. Posterior border of palate: (0) not or clusal outline of p3: (0) long and strongly lat- slightly emarginated; (1) deeply emarginated. erally compressed (length divided by width 33. Elongated styliform process of ectotympan- greater than 1.7); (1) shorter and broader (length ic: (0) absent; (1) present, extends anteromedi- divided by width less than 1.7). 12. Alignment ally and contacts pterygoid process. 34. Anterior of paraconid of m1: (0) in line with other lingual portion (maxillary branch) of zygomatic arch: cusps; (1) situated slightly labial in relation to a (0) shallow and gracile; (1) deeper and more ro- line drawn through metaconid and entoconid. bust. 35. Cleft between premaxillae: (0) deep, 13. Notches in cristid obliqua and postcristid of widely separating I1s; (1) tiny, I1s nearly meet lower molars: (0) absent; (1) weakly developed; at midline. 36. Basisphenoid pits: (0) absent or (2) strongly developed, carnassial-like. 14. La- shallow, double; (1) moderately deep, double; bial cingular cusp of P4: (0) absent; (1) weak; (2) deep and steep-sided, single. 37. Auditory (2) strong. 15. P4 occlusal outline: (0) indented bullae: (0) not enlarged, covers half or less of anterolingually; (1) not indented anterolingually. the periotic; (1) greatly enlarged, covers most of 16. P4 lingual outline: (0) broadly rounded; (1) the periotic. 38. Infraorbital canal length: (0) rel- narrowed anteroposteriorly. 17. Mesostyle on atively short, extending from level of P4 to M1; 752 JOURNAL OF MAMMALOGY Vol. 84, No. 2

(1) elongated, extending from level of P2 to M2. moderately reduced; (2) highly reduced. 49. P3 39. Fossa at anterior edge of orbit (a deep, size: (0) normal; (1) greatly reduced; (2) absent. sharp-edged, funnel-shaped cavity that houses 50. p2 size: (0) normal, about equal to p3; (1) the lacrimal foramen and posterior end of the greatly reduced. infraorbital canal): (0) absent; (1) present. 40. APPENDIX III Fusion of 7th cervical and anterior thoracic ver- tebrae: (0) 7 C not fused to 1 T; (1) 7 C and 1 Data matrix used in the phylogenetic analysis. T fused; (2) 7 C and 1±2 T fused; (3) 7 C and Numbers after each species name denote char- 1±3 T fused. 41. Fusion of thoracic and lumbar acter states given in Appendix II. Question mark vertebrae (to form what is often termed a lumbar indicates that no data were available for a par- ``shield'' or ``tube,'' consisting of 10th and 11th ticular character. Dash indicates that this char- Downloaded from https://academic.oup.com/jmammal/article/84/2/729/2373805 by guest on 29 September 2021 thoracic and all but the last 1±2 lumbar verte- acter is absent. brae): (0) lumbar ``shield'' absent; (1) lumbar Primonatalus prattae.Ð00110, 10000, 01110, ``shield'' present. 42. Number of unfused pos- 01101, 0101?, ?????, ???0?, ??11?, ?????, ????0. terior lumbar vertebrae: (0) all lumbar vertebrae Chilonatalus micropusÐ10121, 01010, 01121, 01101, 12010, 10000, 00100, 10113, 12121, free and unfused; (1) only last 2 lumbar verte- 11100. Chilonatalus tumidifronsÐ10121, brae unfused; (2) only last lumbar vertebra un- 11010, 00221, 01001, 12010, 10000, 00100, fused. 43. Shape of manubrium of sternum: (0) 10113, 12121, 11100. Natalus majorÐ10002, not expanded laterally; (1) expanded laterally 11110, 01211, 01001, 12100, 10000, 00100, and posteriorly into platelike structure. 44. 00111, 11111, 11100. Natalus stramineusÐ Shape of ribs: (0) normal, not expanded or 10002, 01101, 01221, 01011, 02100, 10000, fused; (1) broadly expanded but not fused to one 00100, 00111, 11111, 11100. Natalus tumidi- another; (2) coalesced with each other and with rostrisÐ10002, 01111, 01221, 01011, 12100, sternum, especially ventrally. 45. Length of cau- 10000, 11100, 00111, 11111, 11100. Nyctiellus dal vertebrae 3 through 6: (0) not elongated; (1) lepidusÐ10111, 11000, 11211, 11002, 10121, greatly elongated. 46. Proximal end of femur: 01010, 10111, 21113, 11111, 11101. Kerivoula (0) not sharply bent laterally; (1) sharply bent pictaÐ01011, 01130, 10000, 00000, 10± ± 0, laterally. 47. Shape of femur head: (0) rounded; 00000, 00000, 00200, 00011, 000??. Myotis lu- (1) squarish. 48. Greater and lesser trochanters cifugusÐ00121, 11130, 10100, 00100, 01001, of femur: (0) unreduced, typical for bats; (1) 00011, 00000, 10000, 00101, 00010.