Lnovttattes, AMERICAN MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y

LNovttattes, AMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2618 APRIL 28, 1977

STEVEN F. BARGHOORN New Material of Vespertiliavus Schiosser (Mammalia, Chiroptera) and Suggested Relationships of Emballonurid Bats Based on Cranial Morphology -, .,. IIIII-,I-1..-..1..II-"I .` :," ., I.i -; - ": "-1 :- "",-, -I...... - I,,,,I,1.1....I.,-,,.....'. .. .' .:. ,. t. ,:" ..-.,.... .-,-. .II"....-.. 1, .:'-, :,...., ,'-,". ..II.II.I..,II,.,.,I,I.,,.IIII.I.-..I-:-.I--1II- .I,I.II".1 -'. --.,I ..!.,,-I..,.I ...1I...:II I..;1.I..1.";.,,,..... "I.,.nll .,1;.,'...... I.14.,I.::, I;;. .,.,,."I.,..,. ,... I.I...,..III-,I",.-..""..,,.-....,I,,.I..,I".7'.I-...,.i.I; ,.1.,1. ...-I .I .I.I..II.,-;.II.-I....III-;.,.,.I.-.,,.-.II..III.I.,..I.I. .,..I--II-I....II,..I-.II. I.I....Im,.I..I.i.I-II-. 1,.I;.I ....III1,.....,I.1;..I,..:;1-I,.,-;I..II-I".I..I-.I1 7.;I.,I .II.II.,I-1..:I..--..II.I.: ...Im;..I..q.:.1. -,.e."I.--;liI I;I,I-i.tf.I;...... 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I-.II...j1...;.:.;II;..o."..411..-i.-: -,.-,.. .:..w:..I1.IIII I.III.I-., ,.II.-Ie v;, IIIII-II.,,.,.II.j.III,"1,-.-I,f,II,..I.,..,-I.I., .,-,,.I..-1.P,.,.-1,.I..:..I1,. I.I .I....I..,..-..IIII..,.I. I.I.I,1,III.II. .;-...,., .I.1. I,.. ,.1. ,;.II Ir-,..1 ...I-. ,I. II11II...IIII:I,I.I.,I:.. .., .,..;I.I...!,"...,II..I.,::. .I-,,..I IIIII.:....I.I.-..II.i:-, ,i'. .. ,,, .:.,.II,II AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2618, pp. 1-29, figs. 1-12, tables 1-3 April 28, 1977

New Material of Vespertiliavus Schiosser (Mammalia, Chiroptera) and Suggested Relationships of Emballonurid Bats Based on Cranial Morphology

STEVEN F. BARGHOORN1

ABSTRACT Two previously undescribed skulls of Ves- in closely related genera, are ancient and have pertiliavus Schlosser have occasioned close changed little in the course of later Tertiary evo- examination of emballonurid and other chi- lution. The distribution of skull characters leads ropteran cranial morphology in order to assess its to a somewhat tentative hypothesis of re- usefulness for elucidating phylogenetic relation- lationships, allying the fossil genus more closely ships within the Emballonuridae. A Ludian to to some living emballonurids than to others. Sannoisian age for the fossils, from the Quercy Skull characters can be used to relate these and Phosphorites of France, suggests that many fea- other chiropterans, but with difficulty. tures of the basicranium and ear region, seen also INTRODUCTION Compared with many other mammalian was in general distinctly established in the early groups, new finds of well-preserved, pre-Pleis- Tertiary (see Jepsen, 1970). Paleocene and tocene bats are rare in paleontology. The fossil Eocene material which is not distinctly chi- record has seemingly left students of chiropteran ropteran, if indeed it does belong to the earliest evolution three times short-changed. For one, bats, has yet to be distinguished and removed coverage is scanty and uneven: with a recent from the contemporaneous insectivores (partic- diversity of about 130 genera of bats recognized ularly shrews) and possibly primates (McKenna, (Koopman and Jones, 1970), only a little more personal commun.). Thirdly, within the order the than 30 genera are known from the pre-late Plio- record offers no progressive evolutionary stages cene Tertiary (Romer, 1966), and the majority that might be used to delineate the taxonomy of these are from Europe. For another, the rec- and illustrate the development of the present ord shows no intermediate stages in the origin diversity of bats against the framework of time. of the order: the oldest material which is com- For many taxa morphology has changed little plete enough to assign to the bats with certainty during the later Tertiary; some Eocene and Oligo- shows only that (micro)chiropteran morphology cene material has been acceptably referred to

I Department of Geology, Graduate student, Columbia University, New York; Department of Vertebrate Paleon- tology, the American Museum of Natural History.

Copyright © The American Museum of Natural History 1977 ISSN 0003-0082 / Price $2.40 2 AMERICAN MUSEUM NOVITATES NO. 2618 living genera. More immediately than workers in however, affect their potential applicability to other groups, then, bat paleontologists are forced the problem at hand, namely, relationships to base their methods on comparative mor- within this one family of bats. The outcome of phology, without recourse to stratigraphic argu- the study seems to show, as discussed later, that ments. Good fossil material can both extend the these characters do not enable one to discern the range of known morphology and help clarify pre- relationships easily. The several alternative phy- viously poorly known character distributions and logeny reconstructions all imply that embal- associations. lonurid (and, generally, chiropteran) evolution The present paper describes two new speci- has involved to a considerable degree the inde- mens of emballonurid bats from the Quercy pendent acquisition of characters. Phosphorites, both well-preserved skulls referred to the genus Vespertiliavus Schlosser. Other ma- ACKNOWLEDGMENTS terial, also including skulls, was described by Revilliod in 1920. Apart from diagnosing Ves- I thank Dr. Karl F. Koopman, Department of pertiliavus as a primitive member of the Em- Mammalogy (AMNH), for allowing me free use ballonuridae, Revilliod did not consider relation- of the Recent bat collections and for thoroughly ships among all the genera of the family although reading and criticizing the manuscript; Dr. Mal- his conclusions were based on careful examina- colm C. McKenna, Department of Vertebrate tion of morphology. An attempt is made here Paleontology (AMNH), for allowing the use of the to elucidate those relationships; in general Revil- fossil emballonurids in the collections and for in- liod's conclusions on Vespertiliavus are sup- itially suggesting this project; Mr. G. F. Engel- ported, although difficulties with them are mann for reading the manuscript and pointing pointed out. out several errors; and Mr. Chester Tarka and Cladistic analysis is the method of phylo- Miss Lorraine Meeker, Department of Vertebrate genetic reconstruction used here. This has Paleontology (AMNH), for much patient advice been extensively discussed by Hennig (1966), on the illustrations. Columbia University funds Brundin (1966, 1972), Nelson (1973), and to the author are from a graduate degree program. others, and its theoretical basis need not be defended further in this paper. It is important, Institutional Abbreviations however, that the nature of a cladistic hypothesis AMNH, Department of Vertebrate Paleontology, of relationships makes it possible to state ex- the American Museum of Natural History plicitly its internal inconsistencies-viz., character AMNH (Mam), Department of Mammalogy, the distributions that conflict with the hypothesis as American Museum of Natural History proposed. Conflicting character distributions PU, Geology Museum, Princeton University imply that at least one of the features being in- vestigated has been acquired independently in Figure Abbreviations more than one group and, depending on the ab, ? line of attachment of membranous bulla hypothesis of relationships preferred, is not ac, anterior cingulum homologous among forms that appear to possess alf, alveolar foramina it. Presumably any character (in the most general al + of, confluent anterior lacerate plus optic fo- sense) can be used to adduce or test such a hy- ramen pothesis. This has been assumed in making obser- app, anterior process of petrosal bp, basisphenal pit vations on the material ofthis study and in formu- cf, condyloid fossa lating the morphology in terms of the characters cpb, cochlear process of basisphenoid presented in part in table 2. In fact, at a higher egl, ectotympanic groove lateral part level of taxonomy involving morphologically egm, ectotympanic groove medial part more divergent forms, many of these characters er, epitympanic recess might be considered trivial beside more striking fl VII, flange over path of VII contrasts and unique features. This does not, fme, foramen for ?medial entocarotid artery 1 977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 3 fmma, separate opening associated with foramen SYSTEMATICS ovale fo, foramen ovale ORDER CHIROPTERA fov, fenestra ovalis SUBORDER MICROCHIROPTERA fpg, postglenoid foramen frisa, foramen for ?ramus inferior of stapedial SUPERFAMILY EMBALLONUROIDEA artery frot, fenestra rotundum FAMILY EMBALLONURIDAE DOBSON, 1875 f ?v, foramen for ?vein VESPERTILIA VUS SCHLOSSER, 1887 g, grooves in interpterygoid trough gmp, groove medial to petrosal Vespertiliavus cf. bourguignati Filhol, 1877, g VII, groove for VII lateral to promontorium p. 45, figs. 5-8. hb, hypoconal basin in, internal nares Referred Specimen. AMNH 55349, skull, jf, jugular foramen without jaws, complete except for parts of denti- lap, lamina extension of petrosal tion and middle ear ossifications; somewhat lbp, lateral basal (= pterygoid) pit damaged. lc, lingual cingulum Locality and Horizon. Mouillac, near Caylux, lf, lacrimal foramen Quercy Phosphorites, south central France. mal, malleus Age. Late Eocene (Ludian) or early Oligocene mc, median canal mca, metacrista (Sannoisian) (Thaler and Hartenberger, 1974). me, metacone Diagnosis. Filhol, 1877, pp. 4548; but see "mes,"' "mesostyle" Discussion below. mf, maxillary foramen Measurements. See table 1. mr, molar roots Discussion. The specific assignment of AMNH ms, mesostyle 55349 is not entirely clear, pending revision msn, median septum of nares of the genus. Revilliod (1920) was the first mt, metastyle to describe skulls of Vespertiliavus, of which he occ, occipital condyle had two apart from fragments, one referred to V. pa, paracone bourguignati, the other designated V. schlosseri. paca, paracrista These and the other two species, V. wingei and V. pars mas, pars mastoideus of petrosal on the par proc, paroccipital process gracilis, Revilliod distinguished largely pas, parastyle basis of size. But his diagnoses (and other de- pf, pterygoid flange scriptions) give only one measurement of skull pgp, postglenoid process material for all: the linear distance from Pl to M3 pob, postorbital bar (see table 1, no. 1). AMNH 55349 is larger than pop, prominence on promontorium Revilliod's species V. wingei, the form largest in pp, pterygoid process this measurement, and is in fact smaller than ppf, posterior palatine foramen Filhol's type specimen for V. bourguignati be- ppp, posterior process of petrosal cause its measurement of 11 mm. encompasses pr, protocone only P1 to M2 (Filhol, 1877, p. 47). Were size the prca, precentrocrista only character used for specific taxonomy, it prem, premaxillary be re- prom, promontorium would appear that AMNH 55349 might pt, tubercle on promontorium ferred to V. bourguignati; and that Revilliod's ptca, postcentrocrista material for that species was incorrectly referred ptp, posttympanic process and belongs to a new species intermediate be- pyf, pyriform fenestra tween V. wingei and V. schlosseri. Revilliod, con- sf, stapedial fossa cerned largely with the morphology of the genus spf, sphenopalatine foramen as a whole, did not give fully comparative treat- st, stylocone ment of possible species differences. Erection of 4 AMERICAN MUSEUM NOVITATES NO. 2618 a new species for AMNH 55349 at the moment Age. Late Eocene (Ludian) or early Oligocene seems unncessary. (Sannoisian). Diagnosis. Revilliod, 1920, p. 105; but see Vespertiliavus cf. schlosseri Revilliod, 1920, Discussion below. p. 105, pl. I, figs. 9-12. Measurements. See table 1. Discussion. Specific assignment of this ma- Referred Specimen. PU 11573 skull, complete terial is left open pending revision of the genus; except for right canine, right and left Pl, zygo- on the basis of size this skull appears closer to V. matic arches, ear ossicles, and most cranial roof- schlosseri than to any other species. ing bones. Brain endocast, somewhat distorted, visible on left side. Right tympanic preserved, displaced against pterygoid flange. Right premax- Key to measurements in Table 1 illary bone and incisor present. (Revilliod, 1920, pp. 94-95) Locality and Horizon. Bech-barrond, near St. 1. Linear distance P1-M3. Antonine, Quercy Phosphorites, south central 2. Maximum cranial length-back of sagittal France. crest to anterior border nasals.

TABLE 1 Skull Measurements (in Millimeters) for Vespertiliavus and Examples of Closely Related Genera Saccolaimus Taphozous Vespertiliavus Vespertiliavus peli nudiventris Taphozous Measurement bourguignati schlosseri AMNH AMNH melanopogon Number AMNH (Revilliod, (Revilliod, (Mam) (Mam) AMNH (Mam) (see Key) 55349 PU 11573 1920) 1920) 48757 27391 107656

1. 11.5 8.6 8.0-8.5 6.8-7.3 9.8 8.3 6.9 2. 27.1 21.3 26.5a 22.7 25.8 21.7 18.9 3. 12.4 b c 11.4 12.7 13.6 12.9 4. 20.3d 17.7 22.5 ?18.3e 21.0 18.4 15.5 5. 9.5 8.5 9.8 9.5 9.6 6.8 6.0 6. 14.3 b c 11.3 13.0 8.1 7.5 7. 7.0 5.7 c 6.3 7.0 6.4 6.4 8. 27.8 21.5 c c 25.7 22.8 19.4 9. 17.2 b c c 20.8 14.9 12.5 10. 12.Of b c c 13.2 11.0 10.0 11. 14.9f 11.5 c c 16.5 13.0 11.1 12. ?6.9f 4.9 c c 6.5 4.5 5.2 13. 12.3 b c c 11.3 8.8 8.0 14. 6.1 5.1 c c 7.1 6.1 4.0 15. 14.4g 10.5 c c 11.5 10.1 8.4 16. 4.0 2.5 c c 4.0 2.4 2.6 17. b b c c 10.9 7.9 8.3

a"Longeur totale de l'occiput au bord anterieur des alveoles des C" (Revilliod, 1920, p. 95) not clearly homologous with skull length measured for V schlosseri: "Longeur de l'extremite posterieure de la crete sagittale 'a rex- tremite anterieure des naseaux" (p. 94). bNot obtainable accurately from specimen. CNot recorded by Revilliod (1920). dObscured by remaining matrix. e"Longeur basilaire" (Revilliod, 1920, p. 94): definition unclear. fEvidently affected by distortion as preserved. gLeft side; right side distorted. 1977 BARGHOORN: VESPER TILIAVUS SCHLOSSER 5

3. Length sagittal crest. skull only where it differs significantly from 4. Basal length-median length from posterior AMNH 55349 and where it exhibits morphology edge basioccipital to anterior border max- not available from the latter. illary. Preservation. AMNH 55349 (figs. 1-5) is least 5. Median length of bony palate. in 6. Snout length-front sagittal crest to anterior deformed ventral aspect, although the fol- border of nasals. lowing features are distorted oF lost. There is no 7. Basicranial length-posterior border of basi- sign of premaxillaries or incisors. The left canine occipital to level of anterior border of and all premolars except the last on each side basisphenal pits. have been broken off at the neck. The palate, pterygoid flanges, condyles, basicranium, and ear (Smith, 1972, p. 6) 8. Condylobasal length-back of condyle to regions are all present, although the roof of the front of maxillary. basisphenal pits may have collapsed dorsally. The 9. Zygomatic breadth-greatest breadth across palate is cracked, mostly lengthwise, but with no zygomatic arches. important displacement of parts. Both pterygoid 10. Braincase breadth-maximum breadth. flanges are cracked and displaced such that what 11. Mastoid breadth-maximum breadth across are presumably simple pterygoids appear ambig- back of skull. uously as compound. A sliver of bone is missing 12. Postorbital breadth-minimum breadth in from the ventralmost crest of the right pterygoid; postorbital constriction. the left has been cracked and slightly displaced at 13. Rostral breadth-maximum breadth between its juncture with the palate. Mostly transverse front of circumorbital rims (between cracking and stepwise displacement have oc- lacrimal foramina). 14. Snout breadth-maximum breadth between curred in the bones (basisphenoid and vomer) canines. roofing the internal nares, making interpretation 15. Alveolar length of tooth row-front canine of openings in this area somewhat difficult. The to M3. basisphenoid is also cracked in two places be- 16. Breadth of postpalatal extension-minimum tween the ear regions, with some displacement; breadth between bases of pterygoid possibly this has been accompanied by slight flanges at level of palate. rotation and settling of the petrosals but both of 17. Depth of braincase-from glenoid fossa ver- them are identically placed and no disturbances tically to highest point on braincase. are apparent in the mastoid regions (see below). Crushing and breakage have obscured more DESCRIPTION features in dorsal and lateral aspects (figs. 2, 3). The following description centers primarily on The entire cranium has been flattened, especially AMNH 55349 and is given to (1) describe the posteriorly, though all bone appears to be pres- state of preservation of this new fossil material; ent. Irregular pieces of bone have settled in (2) present in detail certain aspects of chirop- places and are covered with thin veneers of ma- teran cranial morphology as exemplified by this trix in the prepared specimen. On both sides the skull; and (3) compare this form with two closely sphenorbital regions are shattered and dorso- related emballonurids (Taphozous and Sac- ventrally flattened so badly that foramina and colaimust) and occasionally with more distantly other morphology can hardly be made out. It is related forms for purposes of phylogenetic recon- unlikely that cranial flattening has resulted di- struction. Because of the less well-preserved con- rectly in the broken basisphenoid. The angle be- dition of PU 11573, reference is made to this tween this part of the basicranium and the palate is approximately flat; this is also the case in living Taphozous and the condition in the fossil may be 'Saccolaimus is currently recognized as a subgenus of Taphozous. Its skull proportions and ossification, and an accurate preservation. The septum between especially auditory bulla morphology render it a distinct the basisphenal pits (see below) appears to have and valid taxonomic unit within the Family Embal- been warped dorsally under pressure applied at lonuridae, which may be reflected by according it ge- the ?hyoid fragment still remaining in the left neric status (Koopman, personal commun.). It is ranked pit. (The reconstruction of the hyoid elements as a genus in this paper. seems unlikely because of their distorted posi- 6 AMERICAN MUSEUM NOVITATES NO. 2618

5mm FIG. 1i. AMNH 55349. Vespertiliavus cf. bourguignati. Stereophotographs of skull, ventral view. tion, and in view of uncertainties in identifying dorsoposterior margin of the palate and, in lesser them; much of the matrix of the Phosphorites amounts, on the pterygoid flanges. Neither the consists of innumerable unidentifiable bone chips foramen magnum nor the external narial opening and splinters.) has been excavated. Despite the remaining matrix Both postorbital bars are fully present, pre- the morphology has been exposed to be ade- sumably in near natural shape; broken at their quately described within the limits of the state of bases, they are disoriented in the general flat- preservation. tening of the skull. Both zygomatic arches are in Preservation of PU 11573 shows almost no complete condition, cracked through at least cracking of bone and infilling by matrix. The once each, but with minimal displacement. The skull has not been flattened as has the larger fos- right paroccipital process is not in place; a ma- sil, but the cranium and occipital areas show trix-veneered flange of bone next to the base of evidence of overall distortion. Much of the roof- it is most likely the broken piece. ing bone of the left frontal, parietal, and right In preparation the left ear and mastoid region occipital areas has been removed and, due to the have been exposed more fully than the right; the inflated nature of the cranium, a faithful endo- right orbit more fully than the left. For struc- cast has been exposed over this area (cf. Des- tural support matrix has been left surrounding cheseaux, 1956, fig. 1, p. 121). The basisphenoid the (left) canine, both zygomatic arches, the is largely obscured by the displaced tympanic, 1 977 BARGHOORN: VESPER TILIAVUS SCHLOSSER 7 but basisphenal pits of characteristic shape and the maxillary, lacrimal, and frontal bones. Seen spacing are definitely present. The right ear re- ventrally the line of the cheek in front of the gion shows some morphology of the anterior zygomatic arch angles inward more sharply to process of the petrosal not available on AMNH the preorbital constriction in Taphozous, while 55349. The sphenorbital region is preserved on running nearly straight forward in AMNH 55349. both sides. Most significantly, the right premaxil- The lambdoid crest runs straighter from right to lary bone and incisor are preserved. left side than that in Taphozous, in which it General Outline. The general outline, indeed merges backward to a common point with the the whole morphology of the skull of AMNH sagittal crest above and behind the foramen 55349, is similar to Taphozous (especially T. magnum. In ventral outline AMNH 55349 and nudiventris), as discussed by Revilliod (1920). Taphozous nudiventris are nearly identical ex- Contrasts noted are: size larger by about one- cept that the skull in AMNH 55349 is less eighth (see table 1) compared with largest notched in the ear region behind the glenoid. The Taphozous (Taphozous) species; "snout," that is, mastoid region, formed differently (see below), is the area anterior to the abrupt broadening of the slightly more squared in forming the back corner skull for orbits and cheek teeth, relatively longer of the skull. These last two differences are con- and more curved on its dorsal surface; orbits sur- sidered independent of crushing in the fossil and rounded by sharper, more pronounced rims of represent valid distinctions in form. Unfor-

5mm FIG. 2. AMNH 55349. Vespertiliavus cf. bourguignati. Stereophotographs of skull, dorsal view. 8 AMERICAN MUSEUM NOVITATES NO. 2618 tunately, PU 11573 is poorly preserved in this 55349, though not entirely clear of matrix (fig. area but does tend to be intermediate between 4), seems complete except for the premaxillaries, AMNH 55349 and species of Taphozous. incisors, and a small part of the maxillary bone Skull Face. The nasal opening in AMNH above the right canine. The nasal bones extend

II 5mm FIG. 3. AMNH 55349. Vespertiliavus cf. bourguignati. Stereophotographs of skull, lateral view.

H 15mm FIG. 4. AMNH 55349. Vespertiliavus cf. bourguignati. Stereophotographs of skull, anterior view. 1 977 BARGHOORN: VESPER TILIAVUS SCHLOSSER 9

Ii I 5mm FIG. 5. AMNH 55349. Vespertiliavus cf. bourguignati. Stereophotographs of skull, occipital plane. far forward on the snout to a line over the front long. Saccolaimus resembles PU 11573 in having half of the canines. As a result the nasal opening the premaxillaries widely and laterally placed as lies in a nearly vertical plane, a condition ac- seen in anterior view. However, the incisors are centuated by the curvature of this part of the reduced to small nubbins. Taphozous tends to snout. As in most modern specimens, sutures in resemble PU 11573 in having larger incisors, this area cannot be made out and it is difficult to which are still recognizably long and conical. The see from this material, or from his plates, the posterior of the two upper incisors in Embal- basis for Revilliod's (1920) description of the lonura seems to indicate that the lateral one has shape of the nasal bones. The angle of the nasal been retained in Vespertiliavus, Taphozous, and opening (exclusive of premaxillary bones) is Saccolaimus, whereas all other emballonurids oblique in emballonurids where it is not formed have the medial one, rooted directly at the end by deep notching of the snout and palate be- of the tubular portion of the premaxillary. How- tween the canines. The nasal grooves described ever, the situation in the smaller forms is not by Revilliod (1920, p. 92) are prominent here entirely clear, for the entire remaining structure, and run straight forward from the so-called bone and tooth, is very reduced in many cases; supraorbital foramina to the edge of the nasal the morphology may be homologous to that in opening (fig. 2). Here they are associated with a the larger forms with additional loss of the slight notching of the anterior border of the medial alveolar structure. (The Diclidurinae, in- bone. The posterior part of the premaxillary terestingly, have unique complex incisors, gen- bone in PU 11573 rests seemingly loosely on the erally similar to the canines in cusp morphology.) maxillary beside this groove. The bone is flat at Nasal grooves are present in Taphozous its posterior end, but becomes tubular anteriorly nudiventris (but not in other species of the as it lines the sides of the nasal opening. The genus) as shorter, shallower depressions that nest front tubular portion, which is hollow, is pointed the dorsal leaves of the premaxillary bones and medioventrally and extends about halfway to the extend but a short distance behind them. They midline of the narial opening as seen in direct end abruptly not in foramina, or if so the foram- anterior view. The single incisor takes root not ina are minute. The supraorbital foramina in out of the end of the premaxillary, but from the AMNH 55349 are small, located immediately in anteroventral side of the tubular portion. The front of the postorbital ridges (which join pos- tooth has a round neck, long, simple, conical teriorly to form the sagittal crest) and close to form with some curvature, and is oriented the posterior edge of the postorbital bars. The slightly anterolaterally,l much as in Rhinopoma. infraorbital foramen is large in both Taphozous It has roughly the diameter of Pl to judge from and Vespertiliavus (fig. 4). In Taphozous it is the alveolus of the latter, and is about 3/4 mm. actually the confluence of two to four openings 'As disclosed during preparation; subsequently from the nasal region dorso-anteriorly and from broken off at the neck, the incisor is preserved, although the orbit posteriorly. In various species of the break was not clean enough to enable exact replace- Taphozous observed one or more of these may ment. open separately onto the surface; the condition 10 AMERICAN MUSEUM NOVITATES NO. 2618 can even vary from side to side. The internal com- thin wall of bone separates the'se in passing. In plexity has not been prepared out fully in the other emballonurids the lacrimal foramen opens fossils, but in AMNH 55349 the left opening ap- in front of (or just on the edge of) the circum- pears to be paired, the right single. In both the orbital rim. Opening into the orbit next ventrally fossil and Recent genus the opening(s) is (are) al- and posteriorly to the lacrimal is the small ways above plast sphenopalatine foramen, whose canal courses As in the nasal region sutures cannot be made through to the internal nasal passage in an an- out in the orbital region, and no defining state- terior direction. Just ventral and posterior to this ments on extents of bones are possible. The cir- the maxillary foramen marks the entrance of the cumorbital rim appears very pronounced in infraorbital canal into the orbit. It may be larger AMNH 55349, more so than in Taphozous and or smaller than the sphenopalatine among micro- Saccolaimus; this is considered not to be a matter chiropterans generally; it is roughly the same size of skull size. The maxillary bone over the roots in most emballonurids, larger in Taphozous. Sev- of the cheek teeth is so thin as to barely cover eral alveolar foramina pierce the thin bone cover- them and the entire cheek area is deeply concave. ing the molar roots in this region. Medially, on In living bats this area marks the origin of the the dorsal surface of the posterior palatine bor- buccinator musculature, but in no other embal- der, the posterior palatine foramen leads quickly lonurid, including the comparably sized Sac- through to the palate. It is the fourth major colaimus, is it quite so extensive or deeply ex- foramen in the orbital region; it lies distinctly cavated as in AMNH 55349. The postorbital bar separate from the sphenopalatine and maxillary is perhaps less slender than in Taphozous, but (contrast Canis in which this and the sphenopala- seems otherwise comparable in form. tine are nearly confluent, but compare Pteropus, The following foramina open into the Saccolaimus, and Didelphis). Another foramen orbitosphenoid region (includes here orbit opens dorsally and somewhat medially to the proper, dorsal surface of maxillary over the teeth posterior palatine; like the sphenopalatine it too alveoli, palatine, orbitosphenoid, and alisphenoid leads into the nasal cavity. In Taphozous it is bones). See also Pterygoid Region below. The smaller than the sphenopalatine and posterior lacrimal foramen opens into the lacrimal bone palatine, and is situated at the anterior end of the inside the orbit (that is, posterior to the cir- recess in the side of the skull resulting from rela- cumorbital rim, fig. 6). In Taphozous, with tive expansion of the cranium. (Pteropus has sim- nearly identical morphology, the canal from the ilar additional openings from the orbital region lacrimal foramen passes close behind, above, and to the nasal passages.) at right angles to, the infraorbital canal; only a Skull Roof Condition of the dorsal cranium in AMNH 55349 (fig. 2) of course depends on reconstruction, but the following features are of pob interest. The sagittal crest was a low but distinct 0I+of ridge not produced posteriorly as in Saccolaimus; prem it is closest to some species of Taphozous. The pf m. spf roofing bones give no indication that cranial inflations were present. PU 11573 differs markedly pp from AMNH 55349 in these features. Cranial

r inflations were well developed and apparently no

p2 sagittal crest was present. This condition is ap- proached by some of the smaller species of

p Taphozous. M2 MI c Both Taphozous and Saccolaimus exhibit a for the mandibular- FIG. 6. Right orbital region of Taphozous large foramen (possibly nudiventris, AMNH (Mam) 27391, to show fo- alveolar vein) which communicates with the ramina discussed in text. View is lateral and sinus in the postglenoid process and opens in slightly posterodorsal. Camera lucida drawing. the squamosal just dorsal to the glenoid surface Approx. x 4. Abbreviations, pp. 2-3. and at a level anterior to the base of the zygo- 1977 BARGHOORN: VESPER TILIAVUS SCHLOSSER I1I matic arch. No other chiropteran examined has and the last very large and triple-rooted. All pre- this foramen so large and anteriorly placed. molars are present in the type of the type species AMNH 55349 lacks any indication of such an and the same premolar count is indicated by all opening. PU 11573 posesses two moderately referred material. (Schlosser 1877, p. 72 incor- sized foramina opening from the squamosal rectly considered the middle premolar as sin- dorsal to the glenoid, but they are level with and gle-rooted.) Taphozous and other living genera posterior to the base of the zygomatic arch. have retained P1 and plst (Pl being the first of Palate. Preservation of the palate does not the premolar count primitive to bats, a moot obscure the fact that it was long and arched point-see Miller, 1907, p. 24). plast for all the dorsally between the tooth rows. The anterior family is molariform in being as high and nearly border is not prepared in AMNH 55349; in PU as wide as Ml and in having a lingual shelf behind 11573 the notching under the nasal opening the protocone. In crown view the tooth is irreg- comes back no farther than C. The posterior lat- ularly triangular in outline. The paracone is the eral border of the palate in AMNH 55349 (only), dominant cusp, tallest in fact of all the upper rather than skirting closely the neck of M3 (com- cheek teeth cusps (worn apically in AMNH pare Taphozous and Saccolaimus), passes straight 55349). Its surface is convex, conical labially, flat medially to form a shelf behind the protocone of to concave posterolingually, and continuous with M3 before curving to form a recess beside the the protocone basin. A posterolingual cingulum pterygoid. Two small foramina open in the (terminology of Phillips, 1971, pp. 7-8) sur- palate, one medial to the front edge of M', the rounds the basin and continues in a more or less other medial to the protocone of M2. The posterior one is presumably the palatine; the anterior one an additional foramen from the nasal cavity. In PU 11573 both foramina are relatively larger. Dentition. Formula: 1. 1. 3. 3. All other emballonurids except Emballonura: 1. 1. 2. 3. (figs. 1 and 7). The right canine in AMNH 55349 is prepared only on the front and side. The left, broken off irregularly at the base, shows a slightly sub- divided pulp cavity and a posteromedial thicken- ing of dentine in section, indicating perhaps that the canines here, as in the rest of the family, had subsidiary posterior cusps. PU 1 1573 has the left canine fully preserved, broken off at its base and replaced; the right is missing except for the base of the crown. No posterior subsidiary cusp is in- Ms prca st dicated for either tooth. Only one other upper ptca paca canine is figured for Vespertiliavus (Revilliod, 1920, p. 88, fig. 28). The diagram is unclear in its pas perspective, but it shows the tooth broken near mnca pa the tip, the pulp cavity simple in cross section, me a ac and no posteromedial subsidiary cusp, only a pr cingulum. In the modern genera the subsidiary hb IC cusp is at the end of an internal cingulum, and in some of the smaller forms may be little more FIG. 7. Upper right dentition of AMNH than a point on the crest as its skirts the pos- 55349. Vespertiliavus cf. bourguignati, P4-M3. terior border of the tooth. View is oblique: anterolabial and ventral. Vespertiliavus had three premolars: one small Approx. X 11. Stipple: matrix. Abbreviations, pp. and single-rooted, one larger and double-rooted, 2-3. 12 AMERICAN MUSEUM NOVITATES NO. 2618 straight course anterolabially until it curves (highest) mesostyle to metastyle, but is either around the front and close to the base of the worn off or not developed as a sharp ridge be- paracone; it is not continuous on the labial side hind the mesostyle. The ectoloph in both fossil of the tooth. The protocone is a small cusp skulls at hand is developed such that it forms a situated on the cingulum roughly at the level fairly high labial border for both elements in where the lingual slope of the paracone becomes each molar; in all living forms the surfaces of the convex. No anterior cingular style (cuspule of elements are more deeply basined and the ecto- Phillips, 1971) is evident on the anterior cingu- loph on their labial sides more deeply notched. lum. The only other distinctive cusp on plast is In the fossils an anterior cingulum present on what, for the sake of its position with respect to both molars, but particularly on M2, connects the the paracone and for convenience, may be stylocone to protocone. Precentrocrista and post- termed a "mesostyle." In form it is more a short centrocrista do not intersect each other, rather cingulum on the posterolabial slope of the para- they join the mesostyle separately (probably a cone (or more exactly, the precentrocrista). An- consequence of wear). The protocone is at the teriorly it reaches to the level of the base of the junction of the anterior and lingual cingula. paracone proper; posteriorly it nearly touches Ml Where these crests do not affect its form it can parastylar area. PU 11573 differs in plast mor- be seen that the cusp is conical in shape. Seen in phology in the following: protocone better de- direct lingual view, its apex forms the highest veloped on lingual cingulum, with apex quite point of the inner edge of each of Ml and M2 distinct; anterior cingular style slightly better (not entirely evident from fig. 6 due to perspec- developed, apex distinct; "mesostyle" less well tive). Posterior to the protocone the lingual developed. Among living genera the features cingulum (not postprotocrista of Phillips, 1971, described here are somewhat variable, but gen- p. 8; see below) deflects sharply dorsally (as seen erally (1) protocone is distinct and convex lin- in direct lingual view) and lingually (as seen in gually; (2) anterior cingular style is distinct, direct crown view) at a level somewhat anterior sometimes subequal to or even larger than the to the base of the metacone. The point of this protocone in size, such that the two cuspules deflection has no established name. Homology make the anterolingual outline of the tooth, as with a metaconule seems unconvincing because seen in crown view, bilobate instead of straight (1) the crest posterior to this point is the internal to apical; (3) "mesostyle" is less well developed, cingulum continuing to the hypocone, not to any if at all; (4) precentrocrista slopes more sharply part of the metacone; (2) the morphology in dorsally away from the apex of the paracone, question is neither crescentic nor connate-not a and in some cases (Saccopteryx bilineata) a cusp distinct cusp; (3) in some of the modern forms is present at its posterior end, abutting Ml (?a (e.g., Saccopteryx) the postprotocrista running true mesostyle). In features (1) to (3) PU 11573 from the base of the metacone is distinct and is closer to most of the modern forms than to does not involve this part of the tooth. AMNH 55349. Labial to the protocone, the lower shelf of the M1 and M2 are similar in morphology and molars consists of the protocone basin that ex- nearly the same size (maximum crown dimen- tends between the paracone and metacone. Its sions, AMNH 55349, 2.5 by 2.5 mm.). Both are surface is simple except for the slight convexity dilambdodont, strongly divided into a high labial of the posterolabial slope of the protocone. Pos- surface consisting of two triangular subdivisions teriorly the surface of the tooth deflects dorsally ("elements" of Phillips, 1971) formed of the to form the hypoconal basin; thus the boundary paracone and metacone, and a lower lingual sur- between the basins is only roughly homologous face comprised of the protocone and hypoconal with the postprotocrista. In AMNH 55349 this basins. The metacone is characteristically higher boundary is a smooth convexity in Ml and M2; in than the paracone (especially on Ml), nearly PU 11573 it is a smooth convexity in Ml and equal in height with the mesostyle. The ectoloph more a sharp crest in M2. In some Taphozous runs from an anterior parastyle through stylo- species it is similar to AMNH 55349. In other cone (terminology of Van Valen, 1966, p. 8) and species and in Saccolaimus it is barely discernible 1 977 BARGHOORN: VESPER TILIAVUS SCHLOSSER 13 at all; here the two basins together form a simple the parastylar area is lost and the tooth is sur- posterodorsally sloping surface. Among the rounded merely by a continuation of the low smaller emballonurids most genera have some ex- lingual cingulum (see table 2). pression of a true postprotocrista running di- M3 in Vespertiliavus is roughly half as long as rectly from the base of the metacone toward the Ml and M2 and, at its widest extent between protocone, and thus dividing the two deeply ex- parastylar area and protocone, is equally as wide cavated basins. This is considered to be a primi- as these teeth. The ectoloph, in crown view, runs tive character (see also Slaughter, 1970, p. 64). in an extended S-shaped course from parastyle In some genera (Chilonycteris, Furipterus, through stylocone before joining the mesostyle. Mormoops, Mystacina) where the hypocone is Throughout its course it is a distinctive crest with developed as in emballonurids, the postproto- a strongly convex labial surface, as is the anterior crista is also preserved. Diclidurines have a ectoloph on Ml and M2. Parastyle and stylocone smoother convexity dividing the basins as in cusps are as separated from each other as in Ml AMNH 55349. Among other genera Cormura and M2; the highest part of the ectoloph runs brevirostris is closest to Saccolaimus in this between them. The anterior cingulum on M3 feature. is complete. The protocone is distinct but The hypocone has two distinct and recogniz- not as well developed as on Ml and M2. Anterior able forms in emballonurids. In the smaller and posterior lingual cingula curve symmetrically genera it forms a distinct cusp on the postero- from the protocone (as seen in direct crown lingual cingulum, always medial to the base of view); the posterior one leads no farther than to the metacone, that is, on a level anterior (as seen the base of the metacone. The posterior slope of in direct lingual view) to most of the hypoconal the tooth crown is formed by two convex sur- basin. Taphozous, Saccolaimus, and Vespertili- faces, the mesostyle and the metacone. The avus are interpreted as having lost the distinctive mesostyle is relatively smaller and lower on the apex of the hypocone. In these forms the hypo- ectoloph than in Ml and M2; the metacone (as conal basin is rimmed by an even cingulum (PU seen in direct lingual view) is lower than the para- 11573 shows a deflection in the course of the cone, unlike Ml and M2. The lengths of the crests cingulum but no well-defined cuspule). bordering the elements are, in sharply decreasing One difference between Ml and M2 is the rela- order: paracrista, precentrocrista, and postcen- tive extent of the parastylar area. In both teeth trocrista. the parastyle when fully present abuts the pre- Characteristically Taphozous and Saccolaimus ceding tooth and is situated on a level in front of have lost the metacone on M3. The ectoloph is as the anterior cingulum (as seen in direct lingual in Ml and M2 deeply notched in the labial border view), whereas the stylocone is connected to the of the remaining single element in these forms. parastyle by a short curved portion of the The stylocone is the highest point on the tooth ectoloph and is higher than the paracrista. Para- crown, the parastyle having been reduced. The style and stylocone are strongly convex labially anterior cingulum is not continuous about the and even ventrolabially. In Vespertiliavus M1 rela- base of the paracone; as seen in direct lingual tive to M2 has a lower parastyle and stylocone view, it runs down from the parastyle into the relative to the surrounding crown and a narrower base of the paracrista, at about the level of the anterior cingulum along the paracrista face. In base of the apex of the protocone. No distinct Taphozous a complete anterior cingulum on Ml anterior or posterior lingual cingula flank the has been lost; it runs into the face of the para- paracone. The cusp has a simple, medially canted, crista from the parastyle. In some species the conical form, and is situated closer to the base same is true of M2. In Saccolaimus the complete of the paracone than in Vespertiliavus. The devel- anterior cingulum has been lost on both M' and opment of a protocone basin is variable in M2, and the ectoloph does not connect smoothly Taphozous species but everywhere much less the parastyle and stylocone but is notched be- than in Vespertiliavus. All the smaller embal- tween them. In all the smaller genera the para- lonurids have a metacone on M3, although as in stylar area is greatly reduced in M1. In some cases Vespertiliavus the tooth is not fully dilamb- 14 AMERICAN MUSEUM NOVITATES NO. 2618

dodont. Ectoloph, parastylar area, anterior cingu- mc lum, protocone, and posterolingual cingulum are generally as in the fossil forms, except: the anterior cingulum may be more pronounced and the ectoloph more deeply notched as in all the SKk4I I modern anterior molars. The principal difference lies in the robustness of the metacone which is S(S)~~~~~~~~I always strongly conical (except for the postcentrocrista) and higher than the paracone. In addi- tion the protocone basin medial to the base of A the metacone is more fully developed than in the larger living forms. Variably there is a very faint mc, preprotocrista running lingually or even antero- Ibp -, lingually toward the protocone. Pterygoid Region. The only suture visible in the pterygoid region in AMNH 55349 is that be- B tween the palatine and pterygoid bones on the pterygoid flanges (fig. 1). The region otherwise consists presumably of vomer, posterior palatine, presphenoid, ventral frontal, alisphenoid, and anterior basisphenoid. The glenoid surface of the squamosal may also be considered here. The an- C terior side walls of the braincase (ventral facing surfaces of the frontal and alisphenoid) must be reconstructed from their configuration in PU FIG. 8. Basicranial region of three embal- 11573 and living forms. (The skull of V. bour- lonurid bats to show different states of character guignati figured by Revilliod, 1920, plate I, fig. 4, table 2; see text. A. Vespertiliavus cf. bour- 13, shows mostly matrix in this guignati, AMNH 55349. B. Peronymus leu- region.) coptera, AMNH (Mam) 94474. C. Balantiopteryx The internal nares are separated by a septum plicata, AMNH (Mam) 208865. Ventral views. of the vomer as they enter the mouth cavity. Camera lucida drawings, all approx. X 3. Ab- This septum itself is perforated by a canal which breviations, pp. 2-3. leads anteriorly to the nasal cavity (fig. 8A). Pos- teriorly the canal continues as faint, parallel grooves in a single trough in the presphenoid and the anterior lateral edge of the basisphenal pits. basisphenoid as far back as the median border of In bats generally the side wall of the skull is the basisphenal pits. This trough is variously de- pinched in between the pterygoid flanges and the veloped in species of Taphozous, Saccolaimus, cranium. This feature is strongly developed in and other emballonurids (fig. 8B, C), but never Saccolaimus, where (anteriorly) the optic plus so broadly as in AMNH 55349. In Taphozous anterior lacerate foramina and (posteriorly) the and Saccolaimus it is pinched laterally by slight sphenorbital fissure plus foramen rotundum open inflations of the pterygoid and basisphenoid downward from the braincase. In Taphozous, bones and issues from a minuscule opening in the characteristic of the family, these two openings vomer. In the shorter faced genera the canal are separated by a strut of the orbitosphenoid opens more posteriorly between the pterygoid bone, which sends a flattened, sliver-like process flanges in the basisphenoid area and may be projecting forward over the anterior opening. In hypertrophied in some cases (table 2). Saccolaimus these openings are widely separated The pterygoid flanges are simple; pterygoid and the recess between the pterygoids and processes may have been present, but have been braincase extends backward and medially to broken off. The posterior edges of the pterygoid form a pocket behind the sphenorbital fissure. In flanges curve down, back, and laterally to form AMNH 55349 this recess cannot be recon- 1 977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 15 structed with much confidence, but it appears basisphenal pits from the alisphenoid where it from the position of the alisphenoid bone in reaches the petrosal; this, too, is not an ali- front of the foramen ovale that no pocket was sphenoid-basisphenoid suture. developed here, rather that the recess was closer Basicranial Region. The basicranial region is in form to Taphozous. PU 11573 also appears to largely excavated by the basisphenal pits. These be closer to Taphozous. extend forward to the base of the pterygoid The zygomatic arch is straight as in the flanges, lateral almost to the foramen ovale, and Emballonuridae and many genera outside the posteriorly so far as to contour the front of the family. It ends beside the prominent glenoid sur- cochlea. The median septum of the basisphenoid face which is concave, nearly as long as wide, and is of some width all along its length, with the set dorsal to the most dorsal surfaces of the ventral surface flat and square-edged. The bone tooth crowns. In Taphozous and Saccolaimus the broadens posteriorly to form the floor of the glenoid surface extends laterally to include the skull between and behind the petrosals. Depres- back part of the zygomatic arch, making the sions in the basicranial region are present in sev- whole articulating surface a distinctly crosswise eral microchiropteran genera outside the Embal- feature. In Vespertiliavus the surface ends medial lonuridae (see p. 21) with a distribution widely to the zygomatic process of the squamosal. Back- scattered and infrequent enough to suggest that ward movement of the condyle was prevented by outside this family they are not always homol- the very prominent postglenoid process. This is ogous. The function(s) of the pits is (are) un- about half as wide as the glenoid surface, its an- clear. The feature is variable within the family, terior face part of the articulating surface, and is being least developed in Emballonura sulcata therefore concave forward. The large postglenoid (but strongly developed in other species of that foramen opens vertically on its steeply sloping same genus), paired in some genera, unpaired in back side, giving the opening an elliptical shape. others (see p. 26). Its development does not A small foramen opens on the lateral side of the seem to vary allometrically; pits are large dom- postglenoid process at its junction with the zygo- inating features in small-skulled forms (e.g., matic process of the squamosal only in some Peropteryx, Peronymus). In the size series species of Taphozous and in Saccolaimus. The Taphozous melanopogon, T. nudiventris, Vesper- glenoid surface in AMNH 55349 is set on a prom- tiliavus cf. bourguignati, Saccolaimus peli, they inent, heavy development of the squamosal in are nearly the same size throughout, despite a 25 this region. Below the articulating surface the percent range in skull length. bone slopes convexly away to contribute to- In AMNH 55349 the basioccipital medial to gether with the alisphenoid in front to the side the petrosal is contoured such that it does not wall of the cranium and posteriorly to the middle overlap the latter directly but forms a groove ear cavity (especially epitympanic recess). The bordering it that runs back as far as the condy- foramen ovale opens in the alisphenoid on the loid fossa. Near its anterior end this groove is side of this prominence and the course of the interrupted by an extension of the petrosal, a mandibular branch of V close against the bone is thin lamina drawn in curved form from the marked by a short vertical groove exactly medial otherwise smooth surface of the cochlea and ar- to the glenoid surface. The alisphenoid extends ticulating with the basisphenoid (fig. 9). The pet- posteromedially to touch the petrosal and form rosal is thus anchored at the back walls of the the posterolateral wall of the basisphenal pit. basisphenal pits, posteromedially against the Cracking and subsequent infilling by calcite give basioccipital, and medially at the lamina exten- the impression of sutures in the area. On both sion of the petrosal. If it can be inferred from the sides of the skull cracks pass through or just be- remaining morphology of this region of the side the foramen ovale and might be misinter- basicranium that Vespertiliavus, like Taphozous preted as sutures between the alisphenoid and and Saccolaimus, possessed a medial entocarotid basisphenoid, but the cracks are not symmetrical (see below), then the artery should have entered on the right and left sides. The same sort of the cranium just posterior to the meeting of the cracking has separated the deeper walls of the cochlear process of the basisphenoid and the lam- 16 AMERICAN MUSEUM NOVITATES NO. 2618

FIG. 9. Left ear region of AMNH 55349. Vespertiliavus cf. bourguignati. View is ventral and slightly posterior. Approx. x 8. Stipple: matrix. Abbreviations, pp. 2-3. ina extension of the petrosal. Although matrix condition seen in AMNH 55349. No lamina ex- obscures the details, there appears to be space tension of the petrosal is present, and the between the bones at this point for such an cochlear process of the basisphenoid is present opening, though not so foraminate in form as in only as the border of an emargination of that the two Recent genera. (PU 11573 has been dis- bone. The medial entocarotid artery, having torted in this region.) The large hypoglossal fo- passed along the groove about the petrosal, pre- ramen faces directly anteriorly into the groove sumably (in Taphozous at least) enters the cra- about the petrosal (not visible in ventral view). nium through this emargination. In Saccolaimus The foramen magnum is excavated deeply into the situation is different. The basisphenoid an- the posterior margin of the basioccipital, such terior to the cochlear process extends up over the that it opens ventroposteriorly from the skull. promontorium and is appressed to the fully os- The articulating surface of the condyle is likewise sified entotympanic in a vertically projecting directed downward and backward and is not a knobby prominence (see p. 26). Between this symmetrical knob, rather a curved linear feature. and the entrance point of the medial entocarotid In Taphozous and PU 11573 the basicranial the border of the basisphenoid wraps down and bones are not quite so deeply grooved about the medially about the basal turn of the cochlea. petrosals, but T. nudiventris approaches the Ear Region. AMNH 55349 has one of the best 1 977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 17 preserved (and prepared) ear regions known for genera besides Taphozous and Vespertiliavus (not Vespertiliavus (fig. 9). The bulla and ossicles are, observable in Saccolaimus without dissection of however, missing or present only as displaced the skull). The fairly broad groove for the facial fragments. The bulla in chiropters is nearly nerve separates this opening from the base of the always attached so loosely to the skull as to leave cochlea. The foramen for VII opens laterally little sign of its presence after removal. The skull from the side of the promontorium (opening not is generally very thin in the region of the ecto- visible in ventral view). tympanic groove-no thicker, in fact, than the As far as can be discerned in PU 11573 the roofing bone (squamosal) that bounds the anterior process of the petrosal is also slender groove. Only in Saccolaimus and Vespertiliavus and elongate as in Taphozous. It appears on this (AMNH 55349 more so than PU 11573, apart specimen to be medial to the arterial foramen. In from pteropids, whose morphology is otherwise AMNH 55349 the base of the process is evident different) is this area widened to give the impres- just lateral to the arterial foramen. The process sion of an ectotympanic groove lateral to the has been broken off in preparation, but the cross posttympanic process. If the ectotympanic bone section of the base indicates that it was more is comparably placed in Saccolaimus and the fos- flangelike than round, and probably lay more or sil genus, however, this external groove is not the less flat dorsoventrally against the alisphenoid line of its attachment: in the Recent form the bone. In general the epitympanic recess and this ectotympanic rests inedially to this line on a sur- entire part of the tympanic cavity are relatively face of the squamosal just behind the postglenoid deeper in AMNH 55349 than in other specimens foramen. Such a possible true area of attachment observed. Both skulls of Vespertiliavus show nar- can be made out in "Vespertiliavus as well (fig. 9, rower epitympanic recesses than in species of egm). The more latetal groove may have served in Taphozous (Saccolaimus: not visible without dis- the attachment of the external ear. Taphozous section). approximates most other genera in lacking this Behind the anterior process the lateral edge of morphology; here the dorsal part of the ecto- the petrosal bone curves backward and dorsally tympanic ring is flush with the thin side of the and is appressed closely to the squamosal in temporal skull. AMNH 55349. Part of its edge forms a medially The relations of the pars cochlearis to the directed flange that completes the facial canal bones surrounding the middle ear, particularly and forms the medial wall of the epitympanic the alisphenoid, are not everywhere clear; but the recess. Following backward, this lateral edge ter- morphology is certainly, as might be expected, minates in the prominent posterior process of the close to that of Taphozous. The anterior part of petrosal that overhangs the stapedial fossa and the middle ear cavity remains unossified as the nearly touches the promontorium. The groove pyriform fenestra. This is a fairly restricted open- for VII is a broad shallow excavation about the ing between the cochlea and the alisphenoid di- base of the promontorium leading into the rectly medial to the epitympanic recess. Exactly stapedial fossa. The fenestra ovalis opens out how the anterior process of the petrosal (termi- over this groove. It is separated from the slightly nology of Henson, 1970) meets the alisphenoid larger fenestra rotundum by a small prominence laterally to the pyriform fenestra is not clear in on the promontorium. In Taphozous the homol- AMNH 55349, but it probably abuts closely. In ogous structure is a ridge of bone that actually Taphozous the anterior process of the petrosal is projects over the stapedial fossa (Segall, 1974); long and slender and projects anteroventrally Saccolaimus is closer to Vespertiliavus in lacking from that bone (as seen in ventral view). Near its this structure. In Taphozous the internal carotid base (whether medi1l or posterior varies among artery enters the middle ear just lateral to the species) is a foram n, probably for the ramus tubercle (see below), in a gap between the superior of the sta edial artery if this does not entotympanic and petrosal. A slight groove for enter the cranium through the pyriform fenestra. the artery is barely imprinted on the promon- This opening in the petrosal bone is not apparent torium just above the fenestra rotundum in both in a selection of specimens of other chiropteran this genus and the fossils. Above and medial to 18 AMERICAN MUSEUM NOVITATES NO. 2618 the fenestra rotundum is a small tubercle on the petrosal, the stapedial fossa, and a broad, pos- promontory surface which is perforated at its tip. teroventrally facing area behind the fossa that This terminates the curved lamina, present par- constitutes the surface expression of the semi- ticularly on AMNH 55349, which runs to the circular canals within the bone. The mastoid cochlear process of the basisphenoid. In spec- petrosal has become slightly separated from the mens of Taphozous, which have nearly identical squamosal bone and seemingly rotated forward morphology, the posteromedial corner of the relative to it, although the relations with the rest entotympanic rests on this tubercle and in some of the squamosal indicate that the latter has been specimens vessels or nerves can be seen entering displaced. A moderate-sized foramen opens be- its opening. The unossified portion of the ento- tween these two bones just lateral to the pos- tympanic fastens to the petrosal along the terior process (fig. 9, f?v); the opening is round lamina. anteriorly and tapers to a slit posteriorly, medial PU 11573 has part of the right bulla pre- to the posttympanic process. The course of the served. According to the criteria of van Kampen exiting vein (presumably) is marked by com- (1905, pp. 458459; see also van der Klaauw, plementary grooves in the mastoid and squa- 1931, pp. 268-269), the portion consists of the mosal. No such foramen is present in Taphozous ectotympanic and parts of the entotympanic. at the same point, but there is a possibly homol- (Henson, 1961, p. 157, and fig. 1, however, re- ogous opening between the two bones behind fers to comparable structures as ectotympanic the posterior process. However, it is difficult to only.) The anterior and posterior legs (termi- assess the size or significance of this opening nology of Henson, 1961) of the ectotympanic since in this genus it is situated between bones nearly meet, as in Taphozous. Only part of the that are not closely appressed, and possible post medial edge of the entotympanic is preserved in mortem distortion in the Recent specimens ex- the fossil, but to judge from the molding on the amined makes its original form unclear. In Sac- associated matrix, the bulla in this specimen was colaimus the mastoid and squamosal are sutured narrow anteriorly, without stylar process of the in this area; the mastoid is contoured around a tympanic as in Taphozous nudiventris, and medially directed prominence of the squamosal broader posteriorly. It was not so expanded as in at the base of the posttympanic process, such other large emballonurids, but closer perhaps to that the entire joint is tightly structured. There is the condition seen in some of the smaller forms apparently a very small foramen in this suture at (Cormura, Diclidurus, Saccopteryx). Although the base of the posttympanic process; that this is the promontorium of PU 11573 shows the tu- homologous to the much larger opening in bercle described above, there is no ridge ex- AMNH 55349 is questionable. The posttympanic tending forward from it. It is possible that the process is long anteroposteriorly and heavy in bulla of AMNH 55349 would be broader and os- AMNH 55349, Saccolaimus, and Taphozous nudi- sified more extensively than that shown by PU ventris; smaller and thinner in T. melanopogon and 11573. Van Kampen (1905, p. 459) cited a re- the smaller emballonurids. The jugular foramen is mark by Otto (1826) that the bulla in bats be- proportionally larger in Taphozous, in which the comes more ossified with age. delimiting bones are loosely associated in this area, In Saccolaimus there is no tubercle and ridge small in Vespertiliavus and Saccolaimus, but of the on the promontorium, the basisphenoid rather same morphology in all. overlaps the cochlea as described on p. 16. The In Taphozous the bony ridge overhanging the medial edge of the entotympanic is draped over stapedial fossa (p. 17, and fig. 9, pop) is con- this topography, tightly enclosing the middle ear tinued posteromedially in a ridge which projects cavity. Such a fully ossified bulla (seen in all ventrally in the face of the fenestra rotundum. specimens of Saccolaimus examined) is very rare The membranous portion of the bulla stretches among bats, noted also by van Kampen (1905) behind the fenestra rotundum and attaches here. only for Plecotus auritus. In Vespertiliavus and Saccolaimus this feature is The pars mastoideus of the petrosal in AMNH present as a lower prominence of the pars mas- 55349 consists of the posterior process of the toideus, a posteromedial termination of the 1 977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 19 stapedial fossa. The ossified bulla in Saccolaimus, lateral to this is a delicate but well-developed however, attaches not to the prominence, but flange of bone with its back surface facing rather to part of the paroccipital process medi- posteroventromedially; it forms the lateralmost ally. The mastoid is notched medial to the prom- part of the exoccipital bone and articulates on its inence; a short canal thus runs under the pos- front base with the mastoid petrosal and terior edge of the entotympanic bone where it is squamosal bones. The posterior roofing of the anchored on the paroccipital, and leads over the skull is thus deeply notched between the paroc- jugular foramen.' In AMNH 55349 the prom- cipital process and the posttympanic process of inence on the mastoid and the base of the paroc- the squamosal, and the notch is filled by the mas- cipital process are separated by a matrix-filled toid petrosal containing the semicircular canals. space. Since the petrosal may have settled rela- The supramastoid foramen opens dorsal to the tive to the squamosal bone, there is no strong ventral condyloid fossa, about midway on the argument to identify this with the morphology occipital surface. The same is true for Tapho- found in Saccolaimus. It appears to be another zous, but Saccolaimus appears to lack this open- indication that Vespertiliavus lacked a fully os- ing. In Saccolaimus the nuchal and parietal crests sified bulla, that found in Saccolaimus being a are drawn into a posteriorly directed point, unique derived feature. whereas in AMNH 55349 the nuchal line con- The epitympanic recess in AMNH 55349 is tinues smoothly over the top of the skull. Both formed much as in Taphozous but is less deeply forms differ from Taphozous, in which the fora- excavated laterally into the squamosal. The mal- men magnum is relatively larger (in relation to leus (partly damaged) lies with its head, neck, occipital area) and the occipital surface, as also in and anterior process in the recess and its manu- PU 11573, is more domed to reflect this larger brium stretching across the facial canal toward size and the more highly arched cranium. the fenestra ovalis. The malleus had a well- developed spherical orbicular apophysis (termi- NOTES ON CHARACTERS AND HOMOL- nology of Henson, 1961), detached but preserved OGIES USED IN HYPOTHESIZING RELA- in the course of preparation. The muscular TIONSHIPS AMONG EMBALLONURID process for attachment of the tensor tympani has GENERA been broken off, and the manubrium is also slightly damaged at the tip. The malleus appears This section discusses the characters listed in closer to that of Taphozous than Saccolaimus in table 2 and notes briefly the distribution of some the possession of a well-developed orbicular of them outside the Emballonuridae in cases apophysis. No other ossicles are recognizable in where this distribution appears pertinent to the specimen as currently prepared. understanding the primitive or derived condition Occipital Region. The occipital region in within the family. The outgroup comparison, AMNH 55349 is formed by the occipital, cursory and by no means comprehensive, has squamosal, and mastoid petrosal bones (fig. 5). been based on a selection of genera that are The lambdoid crest is a low but distinct edge themselves not grouped here into any preferred running from one posttympanic process to the arrangement but merely listed to show how wide- other. The occipital condyle is more robust than spread certain characters may be. The survey is in either Recent genus. It is flanked by a deep based on examination of specimens in the col- ventral condyloid fossa. The paroccipital process lections of the Department of Mammalogy, AMNH, except for the following, where the lit- 'The function of this canal in Saccolaimus is unclear; erature was consulted: Cyttarops and Depanyc- perhaps it houses the internal carotid artery if this does teris (Thomas, 1913, 1920), Craseonycteris (Hill, not pass medial to the paroccipital process. The branch- ing of the medial entocarotid from the internal carotid 1974), tPalaeophyllophora and tPseudorhino- presumably occurs anterior to this canal, since the in- lophus (Revilliod, 1920), Boneia, Hipposideros, ternal carotid presumably enters the middle ear in all Pteronotus, and Glossophaga (Miller, 1907), and three genera at a point on the promontorium anterior to Cardioderma (Koopman, personal commun.). the jugular foramen. Morphological description is presented to the ex- TABLE 2 Character Distribution Chart for Genera of Emballonurid Bats (Unless otherwise noted: - character absent; + character present. For interpretation of primitive and derived conditions see text.)

Operating

Taxonomic 0 Unit (OTU) z z

1. Reduced + + + + + + + + + + + + premaxillaries 2. Basisphenal pits 2 2 2 1/2 2 2 2 1 1

3. Lateral basal pits +1- 4. Canal and grooves ++ in interpterygoid region + 5. Nasal inflations + 6. Nasal depression ++ 7. Postorbital bar + + + 8. Supraorbital + +1- foramen

+ + _ +l_ 9. Position of lacrimal + + _ _ _ _ foramen (- out; + in) 10. Length of snout ++ (- short; + long)

11. Pterygoid region + _ length (- short; + long)

_ +l_ +1- 12. Inflated cranium +/- +1- + + +l_

+ + + + + 13. Inflation and +1- + + exposure of semicircular canals 14. Upper incisor 1 1 1 2 1 1 1 1 1 1 1 count 15. Upper premolar 3 2 2 2 2 2 2 2 2 2 2 2 count _- 16. panterior +l_ (- short; + elongate) + 17. Hypocone distinct _ + + + + + + + + 18. Parastylar area on +

(+ fully developed; - reduced or absent) 19. Anterior cingulum on + +/- + + + + + + + + + M I (+ complete;

- absent)

+ _ 20. Metacone on MI + - + + + + + + + + + 1977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 21

TABLE 2 - (Continued)

Operating Taxonomic e e

21. Reflected ascending + + + + + + + + ramus and angle of jaw (-slightly reflected; + strongly reflected) 22. Lower incisor count 3 2 2 3 3 3 3 3 3 3 3 3 23. Separate opening - - - + - + + + + - +/_ _ by foramen ovale tent needed to distinguish primitive from derived emballonurids the form is more of a bisected conditions, or otherwise to indicate the variation circle (but not consistently so-e.g., species of which is encountered within the family. Emballonura). As noted on p. 15 this difference 1. Reduced premaxillaries: In all embal- may result from difference in skull size without lonurids the premaxillaries are reduced to bony concomitant difference in pit size. splints which characteristically (1) do not con- 3. Lateral basal pits (= pterygoid pits of tribute to the anterior surface of the palate (as in Sanborn, 1937): These develop associated with Rhinolophus, Pteropus, Artibeus, etc.); (2) never the median basisphenal pit, separated from it by meet medially; and (3) rim the sides of the ex- partial bony septa, and bordered anteriorly by ternal nares in a loose association (not tight the pterygoid flanges. They vary in extent of de- suture) with the maxillary bones. The upper velopment from barely discernible (Emballonura incisor(s) is (are) essentially vestigial. This com- alecto) to major depressions involving lateral bination of premaxillary characteristics appears bulging of the pterygoid flanges. to be restricted to the Emballonuridae. Similar 4. Canal and grooves in interpterygoid region: premaxillary morphology is found in the genera See p. 14 for morphology in Vespertiliavus Tadarida, Lasiurus, and Eptesicus, but in all the and related genera. Canals in the internasal premaxillaries are fused to the maxillaries, and septum of the vomer occur in: Chilonycteris, the incisor(s) (is) are more prominent. Micronycteris, Eptesicus, Myotis, Pizonyx, 2. Basisphenal pits: These are characteris- Mystacina, tPalaeophyllophora, and other gen- tically developed in emballonurids, but basicra- era. Their size varies and they may or may not be nial depressions also occur in: Nycteris, Mor- associated with grooves in the presphenoid and moops, Phyllostomus, Tadarida. In Lavia, basisphenoid. The canal in the vomer is relatively Desmodus, Lasiurus, Myotis, and several other small where found in emballonurids in com- genera, possibly homologous structures are barely parison with most of these genera. In Peronymus discernible; the condition in emballonurids and Balantiopteryx the canal opens more pos- appears to be the hypertrophy of a widespread teriorly between the pterygoid flanges rather feature. Depressions more anteriorly situated are than at or anterior to the intemal nares. The found in Rhinolophus (median), Natalus, and canal is paired in Balantiopteryx and the associ- Thyroptera (paired). In the larger emballonurids ated groove is more a wide, local depression in the ventral plane outline of the pits takes roughly the basisphenoid. the form of a bisected diamond; in the smaller 5. Nasal inflations: Characteristically in the 22 AMERICAN MUSEUM NOVITATES NO. 2618 smaller emballonurids the nasal and maxillary the side of the face (out). The primitive condi- bones are inflated and thinly ossified on the sides tion for bats is unclear. of the snout between the orbits. Generally the 10. Length of snout: Meaning here the gen- inflation leaves the snout laterally expanded and eral region of the skull in front of P4 to the nasal dorsally flat (except Emballonura in which the openings. median area has a furrowed form, and Balanti- 11. Pterygoid region length: In long-skulled opteryx io in which the inflations are two forms the pterygoid flanges lie even or anterior spaced hemispherical protuberances on the to the front of the glenoid surfaces; in short- anterior part of the snout as in Rhinopoma and skulled forms they lie posterior to them. Craseonycteris). Nasal inflations, of various 12. Inflated cranium: The inflation of partic- form, also occur in: Rhinolophus, tPseudorhino- ular areas of the cranium over the primitive lophus, Mormoops, and possibly Lasiurus. condition. The resulting morphology is cor- 6. Nasal depression: In most emballonurids related with reduced ossification of the skull the surface of the skull is somewhat concave whereby the bone is more closely molded over anterior to the bifurcation of the sagittal crest. In the brain. Inflation, when present, appears no- Taphozous (some species more than others) and where distinctive enough to be used to form pos- Peropteryx this feature is accentuated to give a sible sister groups. medial depression between the postorbital bars. 13. Inflation and exposure of semicircular In diclidurines the skull anterior to the sagittal canals: See p. 19. Primitively the occipital crest is inflated (convex) on this primitive pat- and squamosal wrap around the mastoid petrosal tern, and, additionally, lateral nasal depressions leaving the semicircular canals inside the skull are developed in the maxillary bone near the cavity. The mastoid petrosal contributes to the postorbital bar. Nasal depressions are also en- surface of the skull to differing degrees in various countered in: Lavia, Nycteris, Rhinopoma (com- genera of both microchiropterans and mega- pare Taphozous nudiventris primitive pattem), chiropterans. It is also inflated as in the smaller Craseonyctenis, Rhinolophus. emballonurids in several species of Taphozous, 7. Postorbital bar: The relative length of the and more or less so in Nycteris, Rhinolophus, postorbital bar varies somewhat in the Embal- and Tadarida. lonuridae but the structure is always present. 14. Upper incisor count: The third, I3 (of the Also present in: Pteropidae, Nycteris, Lavia, eutherian count), is lost for the whole family, Cardioderma. Weber (1927, p. 133) argued that according to Miller (1907, pp. 27, 88, 93) and the postorbital bar develops in connection with a others. Loss also occurs in: Boneia, Rhinolophus, more powerful bite, seeming to imply that it is Lasiurus, Tadarida, Hipposideros, Rhinopoma, independently derived in those forms which have Noctilio, Craseonycteris, and many others. See p. it. The functional significance of the bar is not 9 on further loss of an incisor. clear, nevertheless. It is tentatively accepted, 15. Upper premolar count: It is probably the however, as an apomorphous feature for the middle P which is lost from the primitive family family. count, since it is double-rooted. Reduction of 8. Supraorbital foramen: This leads through premolar count also shown by: Furipterus, to the orbit and is on the base of the postorbital Pteronotus, Mormoops, Glossophaga, Nycteris, bar or homologous area where this is absent. As Eptesicus, Vespertilio, Artibeus, Rhinopoma, such it is well developed in Pteropus, Lavia. It is Noctilio, Lavia, Craseonycteris, and others. present in modified form in Cyttarops (Starrett 16. panterior: The vestigial premolar in Re- and de la Torre, 1964, p. 55, figs 8, 9), and is cent emballonurids is relatively larger in Tapho- also present in Cynocephalus. With additional zous and Saccolaimus and relatively smaller in widespread occurrences outside the bats it can be other genera though variable in form and detail. considered a primitive mammalian character and 17. Hypocone distinct: See p. 13 for descrip- its loss derived. tion. The primitive emballonurid condition is 9. Position of lacrimal foramen: This may also seen in Noctilio, Lavia, Chilonycteris, open in the orbit (in) or anterior to the orbit on Thyroptera (M1 only), among others. The con- 1 977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 23 fluence of the protocone and hypoconal basins 22. Lower incisor count: The third, I3, is lost, to the extent seen in the large emballonurids according to Miller (1907, pp. 24, 93). Loss of a seems to be uncommon. In several genera (e.g., lower incisor also occurs in: Pteropidae, Rhi- Nycteris, Rhinopoma) where the hypocone is not nolophus, Phyllostomus, Hipposideros, Rhi- a distinct cusp, the hypoconal basin is much nopoma, Desmodus, Craseonycteris, and others. lower than the protocone and the postprotocrista 23. Separate opening associated with foramen may be distinct. ovale: In the primitive condition there is a single, 18. Parastylar area on M1: See p. 13 for de- simple, large opening from the cranium-the fo- scription. The parastyle and stylocone are ramen ovale. In several genera within the family, considered well developed when they are as high however, there is a second opening; this is pre- or higher than the paracrista (as seen in direct sumably the derived condition, but several mor- lingual view); reduced when they are lower than phologies are present (fig. 10). It is not clear this. The well-developed condition is very wide- which of these derived conditions are homol- spread among chiropterans with dilambdodont ogous, if any; and the distribution of possible molars, but reduction also occurs in Mi- homologies conflicts with that of other char- cronycteris. acters, making this an unsatisfactory character on 19. Anterior cingulum on M': The primitive the whole. condition is cingulum present. However, a variety of degrees of development are observed both within and outside the Emballonuridae. In gen- RELATIONSHIPS OF EMBALLONURID eral the smaller emballonurids have a sharp, com- GENERA: DISCUSSION plete, well-defined anterior cingulum, although this may be accentuated by a wear groove be- Figure 11 presents a hypothesis of relation- tween it and the face of the paracrista. In the ships for the genera of fossil and Recent embal- larger forms it is a finer structure but distinctly lonurid bats as represented by material investi- present (Saccolaimus excepted). Absence of the gated in this study. This hypothesis is suggested cingulum also occurs in Nycteris, Noctilio. Non- as an improvement over previous statements on emballonurid genera having the cingulum show less well-developed states than that seen in the smaller emballonurids. 20. Metacone on M3: See p. 13 for description. The widespread primitive condition is metacone present; derived condition, absent. The metacone on M3 is also lost in: Nycteris, Lavia, Micronycteris, Lasiurus, Craseonycteris, and A others. It is nearly lost in Rhinopoma, but a small cusp is present connected to the mesostyle by a reduced ectoloph and to the protocone by a fmma posterolingual cingulum. (See also Slaughter, 1970.) 21. Reflected ascending ramus and angle of jaw: Megachiropterans seem to exhibit the prim- fo itive pattern of having these parts of the jaw C D more or less in a single plane with the dentary FIG. 10. Variation in the opening of the fo- ramus (compare carnivores, lipotyphlan in- ramen ovale. A. Taphozous nudiventris, AMNH sectivores, dermopterans). In microchiropterans (Mam) 27391. B. Peropteryx kappleri, AMNH both ascending ramus and angle are slightly re- (Mam) 239076. C. Emballonura sulcata, AMNH flected laterally (= moderate depth of massoteric (Mam) 87188. D. Saccopteryx bilineata, AMNH fossa). This pattern is further accentuated in the (Mam) 210488. Ventral views. Camera lucida smaller emballonurids, and also in: Mystacina, drawings. A approx. X3; B-D approx. X5. Ab- Nycteris, Rhinolophus, Noctilio. breviations, pp. 2-3. 24 AMERICAN MUSEUM NOVITATES NO. 2618

A Ci4 Z4A4;t(¶) |

(Rhinopoma)

FIG. 11. Hypothesis of relationships among emballonurid bats based on characters in table 2 and the discussion, p. 23. Numbers refer to shared derived characters, table 3. emballonurid taxonomy (e.g., Miller, 1907; consists of the genera Peropteryx, Coleura, Sanborn, 1937; Koopman and Jones, 1970) in Peronymus and Balantiopteryx-those embal- that it attempts (1) to use as many characters as lonurids characteristically with single median can be discerned in the morphology of the skull, basisphenal pits. The genera Cormura, Centro- and (2) to specify as exactly as possible degrees nycteris, Saccopteryx, and the subfamily of relationship among the genera or groups recog- Diclidurinae (includes genera Diclidurus, Cyt- nized as OTUs. Previous work has relied on tarops, and Depanycteris) are, collectively, more standard taxonomic working methods of using closely related to the single pit forms than to any characters for diagnostic purposes only without other emballonurid. Emballonura alone com- always assessing their phylogenetic significance prises the third monophyletic clade. It and, col- (i.e., degrees of apomorphy and plesiomorphy); lectively, the other smaller emballonurids are and of not assessing the significance of, or pos- equally related to the fourth clade-the large Old sible solution for, conflicting character distri- World genera. (The entire family may have its butions. closest relative in the genus Rhinopoma; this is The cladogram may be summarized as follows. merely suggested without thorough inspection of As regards major relationships within the family, the character distributions.) the genera group into four monophyletic clades, The hypothesis of figure 11 fails to satisfy in two of which there are problem areas ex- fully on two counts. First, neither this hypoth- pressed as trichotomies. The first of these clades esis nor any alternative seems to be free of mul- TABLE 3 Explanation of Characters Used in Figure 11 1. Reduced premaxillaries, absence of palatine portion, etc. (see p. 21). Presence of basisphenal pits (extent and form variable). 2. Basisphenal pits of characteristic shape and size. Loss of distinct hypocone on posterolingual cingulum of M' and M2. Confluence of protoconal and hypoconal basins. Large size. 3. Loss of second premolar (of primitive bat formula). Shortening of snout. Loss of one lower incisor. Loss of one upper incisor. Hypertrophy of foramen for ? mandibular-alveolar vein. 4. Loss of supraorbital foramen. Shortening of pterygoid region of skull and of face (distance between C and plast). Inflation and exposure of semicircular canals and pars mastoideus on posterolateral surface of skull. Accentuation of reflection of ascending ramus and angle ofjaw. Loss of second premolar (of primitive bat formula). Loss of parastylar area on Ml. Presence of nasomaxillary inflations. 5. Loss of one upper incisor. 6. Hypertrophy of cochlear process of basisphenoid (see p. 16). 7. Single median basisphenal pit (note: slight indications of lateral pterygoid pits often associated;whether these two features are always correlated is unclear). 8. Hypertrophy of canal opening in roof of interpterygoid region near border of basisphenal pit. 9. Wide separation of basisphenal pits. Hypertrophy of grooves on snout communicating from supraorbital foramina (p. 9). Absence of recessed dorsal narial border (p. 8). Extensive cheek area [for origin of buccinator musculature]. 10. Nasal depression of characteristic form anterior to bifurcation of sagittal crest. Loss of supraorbital foramen. 11. Accentuated breadth of skull, especially across zygomatic arches. Completely ossified auditory bulla. Loss of nasal depression, flat to convex snout. Posterior extension of sagittal crest beyond posterior cranial wall. Hypertrophy of cochlear process of basisphenoid (see p. 16). 12. Nasomaxillary inflations leaving median nasal area furrowed. 13. [None]. 14. Separate opening close to foramen ovale (but see table 2 for distribution). 15. Very large supramastoid foramen between mastoid and exoccipital bones. 16. Paired nasal depressions and associated bony protuberances in circumorbital region, consisting of modified premaxillaries. 17. Accentuated nasal inflations in maxillary bones. 18. [None]. 19. Dorsal narial border: straight and not indented. 20. Hypertrophy of lateral pits associated with median one, separated by partial bony septum and bordered by bases of pterygoid flanges (also occurs on some species of Emballonura). 26 AMERICAN MUSEUM NOVITATES NO. 2618 tiple evolution of features or complicated evolu- groupings. There appears to be no way to arrive tionary histories. The emballonurids fall at a at a least objectionable synthesis; all four groups glance into two distinct groups or morphotypes, remain, therefore, as separate OTUs in uncertain namely (1) large, mostly long-faced, heavy built relationship. Several possible derived characters forms; (2) small, short-faced, lightly built forms. are shared by one or another of these with the Although the fossil Vespertiliavus belongs to the clade of single pit forms. But again, any one ar- former group, it is not clear which morphotype is rangement involves multiple evolution of some the more primitive. The large forms have here characters. This difficulty is not new in the tax- been defended as the more primitive largely on onomy of the family; these genera, with some the basis of the dental formula of Vespertiliavus. others, have been described as close cousin gen- That is to say, a premolar count of three is taken era ("Genres cousins" Brosset, 1966, p. 170), to be primitive for the family; all the smaller and have even been synonymized (Simpson, forms are therefore derived with respect to Ves- 1945, p. 55) despite their earlier recognition as pertiliavus (fig. 11, character 3 in part). However, separate taxonomic entities (Sanborn, 1937). In- this character may not correlate with the large, deed they all but one have the same geographical long-faced morphotype. As with many other distribution; they are all very similar in mor- features, comparison outside the family (e.g., phology generally. Only the diclidurines are dis- Noctilio, Rhinopoma, and others) is ambiguous: tinctly different. They possess deep nasal depres- both long-faced and short-faced morphotypes are sions [accentuated more in Diclidurus and widespread among Mega- and Microchiroptera. Cyttarops than in Depanycteris (Thomas, 1913, There is also the problem that Emballonura, of 1921)] and associated unique morphology. Not the short-faced, light-skulled morphotype, has a surprisingly the three genera have been accorded primitive dental formula (table 2, no. 12). As- subfamily rank within the Emballonuridae suming that both (i) 2 upper incisors, and (ii) 3 (Koopman and Jones, 1970; and others). As in- upper premolars are primitive conditions, the fol- terpreted here, the nasal depressions are merely a lowing hypotheses may be advanced: striking autapomorphy; other cranial features in 1. Emballonurids primitively had short-faced, these three genera are either primitive for the lightly built skulls, as in Emballonura. This family or shared with perhaps some, perhaps implies three independent losses of P3 and a sin- other genera. gle loss of a lower incisor (fig. 1 2A). The presence of a single median basisphenal 2. Emballonurids primitively had long-faced, pit is taken to be homologous among the genera heavily built skulls, as in Vespertiliavus This Peropteryx, Coleura, Peronymus, and Balan- implies two independent losses of P3 and two tiopteryx. Development of pterygoid pits to independent losses of a lower incisor (fig. 12B). some degree is present in all these genera as well, The second hypothesis is preferable because it and their hypertrophy can be used to relate two involves less parallelism in the evolution of P3 of them to the exclusion of the other two. How- and because there is the possibility (contrary to ever, no consistent character can be found to re- Miller, 1907, however) that the incisors lost in late three forms to the exclusion of the fourth or the two clades may not be the same. to relate as sister groups the two forms not ex- A second problem involves Centronycteris, hibiting well-defined pterygoid pits. Some species Saccopteryx, and Cormura. These and the Dicli- of Emballonura (notably E. nigriscens, E. raf- durinae appear to share a derived character franyana cor) exhibit pit morphology quite sim- (albeit independently acquired inSaccolaimus and ilar to that seen in Coleura and Peronymus. some Taphozous species) in the hypertrophy of Emballonura beccarrii possesses a broad, shallow the cochlear process of the basisphenoid. How- basicranial depression of trapezoidal shape, un- ever, as seen in table 2 every possible syna- like any other form in the family. Emballonura pomorphy used to pair two of these forms to the sulcata has very reduced paired pits. (Variation in exclusion of the others, or to combine three to pit morphology in the genus is figured by Tate the exclusion of one other, conflicts with other and Archbold, 1939, p. 3, fig. 2.) On the basis of possible synapomorphies that imply alternate this character alone it might be suspected that 1 977 BARGHOO RN: VESPER TILIA VUS SCHLOSSER 27

jk? .0~~~ Ng ao S 4- aS rSS 4. S

\lose P3 (some) lose P3 lose 13 PIlose P3 (some) lose 13 lose P3 lose 13

lose P3 (shorten skull) A B FIG. 12. Alternative hypotheses for the evolution of the dental formula in emballonurids. See p. 26 for discussion.

Emballonura is an unnatural group, that some characters do appear to be useful in this way. It species might be reassigned among other genera. is, however, clear from table 2 and the discussion It is retained here because (1) such revision on pp. 23-26 that such morphology has not en- would entail multiple loss of the upper incisor tirely clarified relationships within the family (i.e., character 5, fig. 11 would be invalidated), Emballonuridae. It has, nonetheless, led to a and (2) all species of Emballonura examined specific hypothesis on those relationships and characteristically exhibit a median furrow on the pointed to several areas in particular which re- snout between naso-maxillary inflations (see also quire some refinement. Obviously reference to Miller, 1907, p. 87). This character is taken to be associated postcranial morphology should be a valid autapomorphy for the genus. made. 3. The discussion on pp. 19-23 has shown also SUMMARY AND CONCLUSIONS that skull morphology among chiropterans generally will yield many cases of conflicting char- 1. New material of the early Tertiary genus acter distributions, that is, will require pos- Vespertiliavus Schlosser has been described to tulating numerous cases of independent ac- show the extended range of morphology, and quisition of features. This may be true among more complete picture, of this form which has OTUs at both the genus and family levels. Under been brought to light. Vespertiliavus is on the the circumstances, cladistic analysis may be the basis of shared derived characters more closely most objective approach to sorting out these con- related to the Taphozous-Saccolaimus group of flicting character distributions and to wielding living emballonurids than to other members of the large number of named taxa. In dealing with the family and is not regarded as ancestral to the these problems the traditional higher level taxon- family as a whole (as was correctly noted by omy of bats has seemingly wavered between two Revilliod, 1920, pp. 100-101). Indeed it appears equally unsatisfying solutions. In one instance that the smaller emballonurids may exhibit a workers have recognized an excessive number of majority of the -features primitive for the family distinct but unclearly related taxa, often of fairly (contrary to Revilliod, 1920, p. 100). high rank (monospecific families) to reflect the 2. Skull characters have deliberately been inconsistent appearances or irregular combina- emphasized not only because of the nature of the tions of characters (e.g., Hill, 1974, p. 305). In fossil material but also to ascertain the extent of response, others have synonymized extensively their usefulness for bat systematics both below to maintain a workable number of OTUs in an and at the family level. A limited number of attempt to see broader relationships. (See Simp- 28 AMERICAN MUSEUM NOVITATES NO. 2618 son, 1945, p. 180 for criticism of the former and Mag. Nat. Hist., ser. 4, vol. 16, pp. advocation of the latter case.) While making pre- 345-357. liminary assertions about the primitive or derived Filhol, H. nature of certain characters, such traditional 1877. Recherches sur les Phosphorites du methods tend not to conclude in an accessible Quercy. Ann. Soc. Geol., vol. 7, pp. 1-561. manner which groups among the bats are natural Hennig, W. monophyletic assemblages and how much re- 1966. Phylogenetic systematics. Urbana, Univ. lationships among others has been obscured by Illinois Press, 263 pp. convergent evolution. The search for groups Henson, 0. W., Jr. which are wholly primitive has not been suc- 1961. Some morphological and functional cessful, for few exist which do not exhibit some aspects of certain structures of the mid- derived features or autapomorphies [e.g., rela- dle ear in bats and insectivores. Univ. tions of the Pteropodidae or Rhinopoma (Koop- Kansas Sci. Bull., vol. 42, pp. 151-255. man, 1967, pp. 116, 117); of Vespertiliavus it- 1970. The ear and audition. In William A. self]. If indeed bats have been notoriously paral- Wimsatt and D. W. Walton (eds.), Biol- ogy of bats, New York, Academic lelistic in their evolution, cladistic analysis will Press, vol. 2, pp. 1-64. provide explicit statements as to which char- Hill, J. E. acters may have evolved more than once and are 1974. A new family, genus, and species of bat not homologous where they are seen, which may (Mammalia: Chiroptera) from Thailand. be defended as validly relating groups of closest Bull. Brit. Mus. (Nat. Hist.), vol. 27, pp. common ancestry. Competing hypotheses may 303-336. then be assessed with the assurance that they Jepsen, G. L. have been arrived at through comparable 1970. Origins and evolution. In Wimsatt, methodology. The present study has indicated William A., and D. W. Walton (eds.), that the situation is challenging but potentially Biology of bats, New York, Academic workable by this approach. Press, vol. 1, pp. 1-64. Klaauw, C. J. van der 1931. The auditory bulla in some fossil mammals. Bull. Amer. Mus. Nat. Hist., vol. LITERATURE CITED 62, pp. 1-352. Kampen, P. N. van Brosset, A. 1905. Das Tympanalgegend des Saugetier- 1966. La biologie des chiropteres. In Grasse, schadels. Morph. Jahrb., vol. 34, pp. Pierre-P. (ed.), Les Grands problems de 321-722. la biologie. Paris, Masson and Co., pp. Koopman, K. 1-240. 1967. Bats. In Anderson, Sidney A., and J. Brundin, L. Knox Jones, Jr. (eds.), Recent mam- 1966. Transatlantic relationships and their sig- mals of the world: a synopsis of fam- nificance as evidenced by chironomid ilies, New York, Ronald Press Co., pp. midges. I. Principles of phylogenetic 109-150. systematics and phylogenetic reasoning. Koopman, K., and J. Knox Jones, Jr. Stockholm Kungl. Svenska Vetensk. 1970. Classification of bats. In Slaughter, Holl., vol. I1, pp. 1-472. Bob H., and D. W. Walton (eds.), About 1972. Evolution, causal biology, and classi- bats, Dallas, Texas, Southern Methodist fication. Zool. Scripta, vol. 1, pp. University Press, pp. 22-28. 107-120. Miller, G. S. Descheseaux, C. 1907. The families and genera of bats. Bull. 1956. Moulages endocrains naturels des chi- U. S. Natl. Mus., vol. 57, pp. 1-282. ropteres fossiles. Ann. de Paleont., vol. Nelson, G. 42, pp. 119-137. 1973. Classification as an expression of Dobson, G. E. phylogenetic relationship. Syst. Zool., 1875. Conspectus of the suborders, families, vol 22, pp. 344-359. and genera of Chiroptera arranged ac- Phillips, C. J. cording to their natural affinities. Ann. 1971. The dentition of glossophagine bats: 1977 BARGHOORN: VESPER TILIA VUS SCHLOSSER 29

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