Results of the Archbold Expeditions

Home , Corynorhinus, Dasypterus, Glauconycteris, Glischropus, Histiotus, Laephotis

RESULTS OF THE ARCHBOLD EXPEDITIONS. NO. 47 REVIEW OF THE VESPERTILIONINE BATS, WITH SPECIAL ATTENTION TO GENERE AND SPECIES OF THE ARCHBOLD COLLECTIONS

G. HH. ATE

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-"x'~N'' '9~' 99 /9 99,, 9 ~ '9-""' ¼ Article VII.-RESULTS OF THE ARCHBOLD EXPEDITIONS. NO. 47 REVIEW OF THE VESPERTILIONINE BATS, WITH SPECIAL ATTENTION TO GENERA AND SPECIES OF THE ARCHBOLD COLLECTIONS BY G. H. H. TATE

FIGURES 1 TO 5

CONTENTS PAGE INTRODUCTION ...... 221 THE PRIMITIVE VESPERTILIONID...... 222 PRINCIPAL ADAPTIVE CHARACTERS OF THE VESPERTILION1NAE...... 223 EXTRA-FAMILIAL RELATIONSHIPS AND THE AFFINITIES OF VESPERTILIONID SUBFAMILIES...... 226 SUGGESTED PRIMARY GENERIC GROUPS AND TREATMENT OF GENERA AND SPECIES...... 227 Myotini, incliuding "Plecotin i"...... 229 Pipistrellini, including Eptesicus and Histiotus...... 232 Nyeticeini ...... 280 Lasiurini ...... 290 TABLES OF MEASUREMENTS OF VESPERTILIONINE BATS...... 290 SYNOPTIC LIST OF VESPERTILIONINAE (EXCEPT Myotis) IN THE ARCHBOLD COLLECTIONS...... 297

INTRODUCTION This paper is the eighth and last of a tanical material assembled have also ap- series of publications devoted to the study peared. and determination of a large number of The present work on the Vespertilioninae Microchiroptera assembled by Mr. Richard has been facilitated by the use of specimens Archbold, which are deposited in The generously lent by the curators at the mu- American Museum of Natural History, seums at Chicago, Cambridge and Wash- New York. ington. In addition, numerous photo- The bats in question form a part of a graphs of types, made by the kind per- large collection of mammals assembled by mission of the authorities at various Eu- Mr. Archbold during the decade 1930-1940, ropean and Australian museums, have mainly by means of his own expeditions aided materially in establishing the exact to Madagascar, New Guinea and Australia, characteristics of species inadequately de- but also through the work of collectors scribed. whom he sent from time to time to Celebes, One new species, Pipistrellus anthonyi, Borneo, Java and Sumatra. is described. Our interest in the oriental region has Preliminary papers relating to the Ves- centered upon the mammal fauna of the pertilionidae have been published in which island of New Guinea. But in order prop- the subfamilies Miniopterinae, Murininae, erly to study the mammals of that island Kerivoulinae and Nyctophilinae were suc- it has been necessary to work intensively cessively dealt with.1 Myotis, the most with those of Australia on the one hand and primitive genus of the remaining subfamily, of southeastern Asia and the East Indian the Vespertilioninae, had been previously islands on the other. reviewed.2 But it was found in practice Reports based upon the fruits of Mr. that the remaining vespertilionine genera Archbold's tireless collecting have been were often too closely integrated one with published with considerable regularity, the another to permit satisfactory treatment field of research extending far beyond mam- 11941, Bull. Amer. Mus. Nat. Hist., LXXVIII, and other Art.9. malogy to ornithology zoological 2 1941, Bull. Amer. Mus. Nat. Hist., LXXVIII, departments. Numerous papers on the bo- Art.8. 221 222 Bulletin American Muse'um of Natural History [Vol. LXXX one by one. Instead, a procedure is here species," based upon populations relatively adopted under which the subfamily is re- homogeneous both morphologically and viewed as a whole, and the interrelation- functionally, whose individuals normally ships of the genera are pointed out, as far interbreed freely and breed rarely or never as understood. In many instances the data with other species even when they inhabit upon which certain species or races have the same environment-despite this knowl- been founded are so vague that it has not edge and theory and the understanding that been possible to reach a conclusion regarding one should deal with their status. I do not hesitate to assert large samples statisti- that many of the names will go into syn- cally, in practice the museum student is still onymy. This action should occur, how- compelled to found his assumptions or ever, only after ample facts have been "conclusions" upon quite small samples or marshaled in proof. As much evidence is often upon random individuals taken from needed to prove synonymy of a species as different localities. To a great extent also to demonstrate the distinctness of a new he must rely upon other men's observations. one. It was with the above mentioned idea Thus he still builds "species" upon mor- of an inclusive review of this very complex phological distinctions seen in single speci- subfamily that the work following was mens (types). But he is nevertheless aware undertaken. of his handicaps and awake to the short- Despite the modern concept of "the comings of methods which he must use.'

ALPHABETICAL LIST OF GENERA DISCUSSED Particular Attention Has Been Paid Those Printed in Bold-faced Type Baeodon.. 283 Mimetillus.. 265 Barbastella.. 263 Myotis. 229

Chalinolobus. 260 Nyctalus...... 254 Corynorhinus.231 Nycticeius ...... 283 Dasypterus. 290 Otonycteris.. 289 Discopus...... 233 Philetor ...... 265 Eptesicus. 271 Pipistrellus...... 233 Euderma...... 232 Plecotus ...... 229 Glauconycteris. 263 Rhinopterus... 279 Glischropus... 253 Rhogeessa..... 283 Hesperoptenus...... 268 Scoteinus... 280 Hi,stiotus.279 Scotoecus. 283 Ia...... 259 Scotomanes... 288 Idionycteris..... 231 Scotophilus...... 283 Laephoti&.279 Scotozous. 259 Lasionycteris.. 229 Tylonycteris...... 266 Lasiurus.... 290 Vespertilio.... 270

THE PRIMITIVE VESPERTILIONID Cases are numerous in the Vespertilioni- domed, with postpalatal spine; basicranial dae of parallel or convergent development region with cochleae and bullae only from relatively independent genetic lines. moderately large, without basial pits. The primitive, unspecialized vespertilionid Dentition: upper incisors, two each side, skull may be considered to have possessed bifid; lower incisors, three each side, trifid, the following characteristics: moderately not at all or weakly imbricated; canines full braincase; zygomatic arches strongly not approximated, without accessory cusps; built and bearing small postorbital proc- upper and lower premolars three in each esses; rostrum fairly slender, elongate, not jaw, subequal in height and in cingulum especially broadened at lacrimal level and 1 Reference: The symposium on species held at the tubercles. joint meeting of the Amer. Societies of Ichthyologists, without pronounced supraorbital Herpetologists, Vertebrate Paleontologists, April, Palate rather narrow, weakly arched or 1942. 1942] Tate, Results of the Archbold Expeditions. 47 223 lengths, except p4 which were less blade- tragus was simple, unnotched and prob- like and more molariform; m3 with Z-pat- ably much like that of simple types of tern complete; m3 with posterior triangle Pipistrellus. The wing was probably at- scarcely reduced. tached to the terminal part of the meta- In the skin one may expect none of the tarsus. This type of attachment is com- special adaptations of ears, feet and wings mon throughout the Chiroptera. Attach- which have been developed by certain of the ment to the wrist (some Myotis) or to one genera and subfamilies. The ears were of the phalanges of the fifth digit (Murina) simple, separate and of moderate size. The may be considered new modifications.

PRINCIPAL ADAPTIVE CHARACTERS OF THE VESPERTILIONINAE Upon the basic pattern above described bat otherwise considerably specialized, certain modifications have been imposed, e.g., in Histiotus. not once only, but so often that it is difficult to avoid the idea that hidden trends PREMOLARS or tendencies are present. The customary change in the premolars One of the earliest, most widespread of happens through the sharp reduction in these modifications came in connection with size of the third and second teeth; here seizing and masticating food: most vesper- the teeth of the upper jaw precede those of tilionid bats tend to shorten the face, jaws, the lower. The upper third premolar dis- palate and toothrows. This process comes appears, then the lower third; next the about by shortening of the bones of the upper second tooth goes. Although the face with the result that the fronts of the lower second premolar remains in many orbits continually approach the roots of genera fairly large, in the Nycticeini and the canines. In extreme cases the anteor- others it may become exceedingly small. bital foramen may reach a position almost over the canine. At the same time shortening of the INCISORS toothrow takes place byreductionand elimi- The incisor teeth undergo modifications nation of p3- and p2, the corresponding also. In part at least the changes in the teeth of the upper jaw being lost before incisors are connected with approximation those of the lower jaw. A further point of the canines. Thus imbrication of the where shortening of the toothrow takes lower incisor teeth, often accompanied by place is at m3. This is accomplished by thickening of the crowns, is seen when the reduction of the posterior limb of the Z-pat- lower canines stand close together. tern and by shortening of the body of the In the upper incisor teeth reduction takes tooth longitudinally in the toothrow. place in the secondary, or lateral (sometimes posterior), cusp of each tooth, leav- THIRD MOLARS ing a peg-like unicuspid tooth. It also Modification of the regions of the pre- occurs by displacement outward (Ia, some molars and of m3 may take place inde- Pipistrellus, etc.), inward (Hesperoptenus) pendently. The alteration of m3 appears or by obsolescence (many genera) of the to be a more advanced, more recent speciali- outer incisor (i2), permitting il ultimately zation and one occurring at random in to come in contact with the cingulum of the many genera. For example, in Eptesicus, canine (Scotophilus). nilssonii m3 is entire, while in E. serotinus The lower incisors, primitively tricuspid m3 is markedly shortened. This fact is ap- (as are the milk incisors of the upper jaw), parent too in Pipistrellus and other genera. show slight changes in addition to imbrica- But it must not be thought that m3 in un- tion. The outermost lobe of the three is modified condition necessarily indicates separated by a deeper sulcus than is the in- that the bats showing it are primitive. It side one. In Rhogeessa -this outer, lobe, be- may be merely one unmodified feature in a comes obsolescent, and i3 is reduoed.- 2.. 224 Bulletin American Museum of Natural History [Vol. LXXX

CANINES RosTRum The chief changes in the canines, apart The height and breadth of the rostrum from size, relate to development of acces- may go with that of the braincase, as in sory cusps (a) on the posterior cutting Nyctalus and Histiotus; also in species with edge, (b) on the cingulum. broad, flat braincases the rostrum may be An accessory cusp is present in Tylonyc- broad and flat, as in Scoteinus, Tylonycteris teris and sometimes in Pipistrellus, but and Scotophilus. In the case of the primi- only in certain groups of species. It is ab- tive, little modified rostrum (unbroadened) sent in Ia, Nyctalus, Chalinolobus, Hesper- there is often a weak median sulcus, as in optenus, Lasiurus, Scotophilus, Scotomanes, Myotis, some Pipistretlus and many other Otonycteris, Nycticeius and allies, Eptesi- genera. Quite marked depressed areas cus, Scoteinus (Australia). A strongly de- may appear on either side of the muzzle veloped posterior cingulum cusp is seen in just anterior to and behind the canine and Philetor and Mimetillus. in front of the anteorbital foramina (Epte- sicus fuscus group, etc.). In Barbastella a markedly depressed dorsal area is seen on BRAINCASE the rostrum and a median spine in the an- The braincase and intertemporal region terior narial sinus. in primitive forms were described earlier Coupled with such expansion of the ros- as "moderately full." Broadening of the trum and in certain cases possibly with a braincase had been indicated in the Nycti- supplementary process comes the enlarge- ceini, Tylonycteris and allies. It appears ment laterally of the supraorbital tubercles. also in Hesperoptenus and in less pro- This condition occurs in the joffrei group of nounced form in Nyctalus, Ia and some Pipistrellus and is found at its optimum in groups of Pipistrellus. Extreme flattening Mimetillus and Philetor. In Tylonycteris in combination with such broadening is seen it is less well developed. In Scotomanes, only in Tylonycteris and in Discopus. although the lacrimal region is much The reverse condition, excessive fullness widened, the tubercles are limited to nar- of the braincase, occurs twice, namely, in row bony ridges placed directly above the the Lasiurini and in Chalinolobus and Glau- eyes. conycteris, in each case combined with ex- The anteorbital foramen normally is a treme shortening of the face. small, pore-like opening placed just in In some bats the intertemporal area ap- front of the orbital fossa. In Scotophilus pears much constricted by contrast with (less in Scotomanes and not at all in Otonyc- the secondarily broadened rostrum. In the teris) the pore becomes considerably en- case of Histiotus the intertemporal region larged, so that its externally bounding wall appears definitely expanded both laterally is reduced to a slender bar separating the and upward. foramen from the orbital fossa. Consider- Osseous crests, the sagittal and lamb- able enlargement of the anteorbital fora- doid crests, are obsolescent in the majority men may be perceived also in Histiotus, an of the vespertilionine genera. They are offshoot from Eptesicus, and in Scotoecus. present in advanced Nycticeini (especially In the Lasiurini, on the other hand, the in Scotophilus kluhlii, in which a "helmet" foramen is represented by minute pores on is developed), also in Australian Scoteinus, each side, which because of extreme short- the serotinmus group of Eptesicus, in Ia, ening of that part of the maxilla lie almost Hesperoptenus and Nyctalus. It will be directly above the canine. recalled that weak crests are present in the myotis section of the genus Myotis, in cer- ZYGOMATA tain Nyctophilinae and Miniopterinae. In Perhaps as a secondary effect of the general, crests are present in large bats, modifying of biting, the zygomata become absent in small ones. They are correlated altered. In a great many genera the post- with strong temporal muscles and heavy orbital processes of the zygomata have mandibles. disappeared, following which the zygomata 1942] Tate, Results of the Archbold Expeditions. 47 225 themselves have become so weak and Nycticeini; the heavily built skull (less slender as to appear almost functionless. broadened, however, than in Scotophilus and Scotomanes), the well developed supra- PALATE orbital tubercles, the massive character of The anterior palatal sinus is normally the teeth, with reduction of P1 to peg-like about half as wide as the canine width form, the elimination of i2 and p2, the nar- (outer). In Nyctalus, Vespertilio and Lasi- rowing of the canine width and small size urus this sinual width is greatly increased. of P2 emphasize the relationship. 'Usually it is smoothly rounded, or slightly Emphasis was laid by early writers upon squarish in outline. In Tylonycteris a trace the shapes of the tragus. Dobson in par- of a median projection appears, which in ticular based much of his specific classifica- Philetor may become a spine definitely pro- tion upon minute differences of degree of jecting forward. The posterior palatal curvature of the margins of the tragi. spine is normally either finely pointed or triangular, projecting backward beneath THUMB-PADS AND FOOT-PADS the narial canal. In Scotophilus, especially Just as enlargement of the ear appears S. kuhlii, the spine projects downward to have come about more than once, so from the palatal surface. development of thickened pads or adhesive disks on thumbs or feet in the Vespertili- BASISPHENOID PITS oninae has taken place repeatedly. The "Basial pits" (Thomas) must be regarded two best known genera in this category are as indications of specialization, although Glischropus and Tylonycteris. Glischropus I do not know their function. They usually is virtually a PipistreUlus in which the pad adjoin the inner sides of the cochleae and of the metacarpal of the thumb is much may be observed in a number of genera. enlarged, though by no means a peduncu- Possibly those of Eptesicus dimissus are late disk as in Thyroptera (tropical Amer- not homologous with the pits customarily ica). The feet also are broadened and seen (e.g., in Hesperoptenus tickelli). thickened. The relation of Glischropus to Pipistrellus is best seen in the P. tenuis AUDITAL ORGANS group, in which the features mentioned Extreme enlargement of the pinnae of are weakly developed. The premaxillae the ears has appeared among the Vesper- of Glischropus are shortened much as in P. tilioninae three times. The plecotine bats, tenuis. Plecotus, Corynorhinus, Idionycteris and The pads of Tylonycteris, on the other Euderma, represent a compact group de- hand, are strongly divergent. Tylonycteris rived, if one may judge from skull charac- possesses wing- and foot-pads, similar ters and dentition, from the parent stem to, but better developed than, those of leading to Myotis and Lasionycteris. Re- Glischropus. But the extremely specialized duce the size of the ears and audital bullae flattened skull permits no close affiliation of Plecotus, eliminate the myotine p3, and of the two. Taking into account the widen- what is left is almost identical to Lasionyc- ing of the lacrimal area, the loss of p2 and teris. The second large-eared group, the the reduction of the crown length of P4 Histiotini, including Histiotus (tropical in the toothrow, Tylonycteris appears dis- America) and Laephotis (Africa), derived tantly associated with Mimetillus (Africa) probably from Eptesicus, obviously has and Philetor (New Guinea). In these progressed farther from the ancient line. latter genera the dentitions, as well as the In both of these genera an Eptesicus stage broadening of the skulls and palates and of dentition has been achieved. Both development of supraorbital tubercles, braincase and rostrum are high and full in show close correspondence to Tylonycteris. Histiotus, rather less so in Laephotis. The The flattening of the skull so conspicuous third group of big-earedvespertilionine bats- in Tylonycteris is far. less obvious-in Mime- is represented by Otonycteris. The affini- tillus and is not found in Philetor. ties of this genus appear to be with the Discopus, recently described from Indo- 226 Bulletin American Museum of Natural History [Vol. LXXX

China by Osgood, evidently became sepa- bears a small, lobe-like, flattened projec- rated from the pipistrelline stem far back tion, the calcareal lobe. This structure in time, because, like Lasionyteris, it re- may be present or absent in the same tains p3 and has the rostrum relatively un- genus (example, Myotis). It seems to have shortened. In it the braincase is depressed no systematic value when groups of higher nearly as in Tylonycteris but shows no or- order than species are under consideration. bital processes. This bat, with ear rather The baculum has been used extensively long and narrow, has pads developed on the as a basis of classification in the Rodentia. hind feet "even more extreme than in It seems probable that study of this struc- Tylonycteris and Glischropus." Pads were ture in the Microchiroptera would yield also described in the poorly known Hesper- valuable results. Beginnings have been optenus blanfordi. made in certain groups: Thomas studied the OTHER STRUCTURES bacula of the genus Nyctophilus; the late The interfemoral membrane is constantly Dr. G. M. Allen had gathered together a extended between the tuber calcanei and the number of bacula representing many of the cartilaginous spur which is attached thereto, species of Pipistrellus which he intended and the tip of the tail. In certain species to employ in revising the genus. In general, of Myotis and other genera the last caudal very few of the bacula of the Vespertili- vertebra may project beyond the mem- onidae are known, so it will be necessary, brane. Again, in Lasiurus and in the sub- before any extensive work based on them family Murininae the dorsal surface of the can be attempted, to assemble adequate membrane may be quite densely covered representation from the collections of bats with hair. In certain groups the calcar in alcohol and perhaps anew from the field.

EXTRA-FAMILIAL RELATIONSHIPS AND THE AFFINITIES OF VESPERTILIONID SUBFAMILIES Vespertilionid bats with premolar den- evolved yet a third pattern of pharyngeal tition of least specialized type occur in anatomy while also eliminating completely the New World among the Thyropteridae one of the two anterior upper premolars- and Natalidael (part). The Old World probably p2, since p2 iS considerably reduced Kerivoulinae, in respect of their premolar in size. arrangement, show indications of reducing In the Natalidae a special type of basis P2-3 and thus suggest the method of ap- cranii is present whereby that part of the proach to the present condition in Myotis. mesopterygoid fossa which lies between In most other respects those families of the wings of the pterygoids is separated ancient lineage have pursued independent from the posterior area, in which basisphe- pathways. The Kerivoulinae, Furipteri- noid pits are present in certain chiropteran dae and Thyropteridae have developed genera, by a strongly developed transverse greatly inflated braincases and low, tip- ridge aligned with the two glenoids. The tilted rostra, and the Thyropteridae have anterior portion is the more pitted, and the produced the well known modifications of pterygoid palate extends far backward, the pads of thumbs and feet. But the two partly covering it. Mesially it is divided families have diverged markedly in the in two by a thin wall of bone (the vomer?). structure of the pterygoid region and The greatly enlarged Eustachian tubes, basis cranii. ossified only externally, open on this The Furipteridae (Furipterus and Amor- pharyngeal region. phochilus) have pursued the above men- Natalus, Nyctiellus and Chilonatalus all tioned course of cranial and rostral speciali- show this character of the pterygoid area, zation to an even greater extreme but have Phodotes to a less extent. A homologous, though in detail slightly different, develop- 1 Separated by H. Allen, 1893, Bull. U. S. Nat. Mus., XLIII, p. 55. ment of the bony framework of the pharynx 19421 Tate, Results of the Archbold Expeditions. 47 227 is seen in Thyroptera. The structure taken tween the Vespertilioninae and those other as a wbole appears to be a specialization groups currently considered subfamilies, peculiar to American bats.' the Murininae, Miniopterinae, Nycto- Natalus and Phodotes are more closely philinae and Kerivoulinae,2 only the last allied to each other than to either of the can be regarded as at all nearly related to other two genera, Chilonatalus and Nyctiel- the Vespertilioninae. The dentition of the lus, in which smaller size and reduction in first three subfamilies mentioned is already size of p2 are apparent. In Nyctiellus p2 so specialized that no obvious clue to the is already minute, reaching only to the path of divergence followed by them re- depth of the cingulum of the canine. This mains. In the cases of the Murininae and type of premolar modification is wholly Miniopterinae there is no way of knowing different from that pursued in the vesper- whether the missing premolar is p2 or p3, ex- tilionine bats in which, though both p2 cept doubtfully in Miniopterus by analogy and p3 are obsolescent, p3 is always the with the lower premolars. first to grow smaller and disappear. The Nyetophilinae are, however, a little In all of the Natalidae exceptional length- different. They have reached the Nycti- ening of the low, flattened rostrum has ceius stage of dental reduction, though taken place. This is somewhat analogous they may have arrived there by a path to what has occurred in the subfamily different from that taken by the Nycticeini. Glossophaginae of the family Phyllosto- In addition they have developed their midae. Such lengthening of the jaws, with characteristic nyctophiline modifications accompanying spacing of the teeth and (noseleaves, etc.). No matter whether they modification of the premolars into com- are regarded as Vespertilionids separated pressed elongate blades, is the antithesis of in remote times from the vespertilionine, the tendency to shorten the face and which independently achieved nyctophiline mandible, visible throughout the Vesper- dentition, or whether they are ronsidered as tilioninae. hyperspecialized Nyeticeini, they can have It can be concluded that, although the played no part in the ancestry of those Vespertilionidae, Thyropteridae, Furip- lower Vespertilioninae with relatively com- teridae and Natalidae have had a common plete dentition, the Myotini. origin, they have since diverged steadily. It is also apparent that the Thyropteridae That leaves only the Kerivoulinae, alone among the last three families show which, as I stated in my earlier paper, some slight resemblance to the vespertilio- show incipient reduction of p2- 3, as pos- nid subfamily Kerivoulinae, though they sible relatives of the less progressive genera differ so greatly in the pterygoid and rostral of Vespertilioninae. The relationship can- regions of the skull and in the ears, wings not be very close, for the Kerivoulinae and feet that the similarity between the demonstrate skeletal peculiarities not two is better regarded as fortuitous. shared with the Myotini. Nevertheless Considering next the relationships be- the affinity seems to be real.

SUGGESTED PRIMARY GENERIC GROUPS AND TREATMENT OF GENERA AND SPECIES The more than thirty vespertilionine have undergone no important modifica- genera of bats fall chiefly into four main tion; p!-3 are reduced in size; m3 is gen- divisions which may be named Myotini, erally complete. Pipistrellini, Nycticeini and Lasiurini: 2.-Genera coderived with Pipistrellus. 1.-The first are long-faced genera co- In this division, too, the upper incisors are derived with Myotis, in which pi are both little or not at all altered, but p3 are absent present, or p3 is absent. The upper incisors (except in Discopus), and to a greater or 1 A slightly similar development can be seen in 2 Reviewed in Bull. Amer. Mus. Nat. Hist., Nyctalus velutinus. LXXVIII, Art. 9, 1941. 228 Bulletin American Museum of Natural History [Vol. LXXX

Pipistrelloid Eptesicoid genera, p2 genera, p2 usually present absent

Pipistrellini Nycticeini i2 present i2 absent

Disco us lYotini asn us| Llly present pabsent p3 present, redtxced in size discs on thumbs \ / / Lasiurus Dasypterus 2 absent _ _ 1F;. Relatively unspecialized Mammae 1 - 1 = 2 \/ Lasiurini highly specialized, with greatly shortened face and reduced dentition. Mammae 2 - 2 = 4

5 Kerivoulinae Vespertilioninae rarely reduced p , when present, reduced in size in size; or absent

[ Vespertilionidae I

Fig. 1. Suggested phylogeny of the Vespertilioninae. See also Figs. 2, 3, 4, 5. less degree the face has undergone short- these divisions quite profound specializa- ening; in the Eptesicus subdivision p2 has tions of various sorts have obscured the also disappeared, and in many genera re- basic pattern first indicated. In many lated to Pipistrellus p2 is very greatly re- instances the same kind of specialization duced and often displaced inward from the occurs in two or even three of the divisions. toothrow; m3 is complete or incomplete. One example is the independent develop- 3.-Genera of the general type of Nycti- ment of large ears in Plecotus, Histiotus and ceius, Scoteinus and Scotophilus, in which Otonycteris. the incisors as well as the premolars are 4.-The highly specialized American much modified, and m3 is frequently re- Lasiurus and Dasypterus stand apart from duced. In this division i2 and the acces- the divisions just defined. These remark- sory cusp of il are almost always obsolete, able bats, confused by some early authors il becoming a peg-like tooth often closely in with the Murininae in which also the uropa- contact with the canine. In all three of tagium is densely pilose, diverged unques- 19421 Tate, Results of the Archbold Expeditions. 47 229 tionably very far down the vespertilionine 2-2 may indicate primitiveness, but even trunk. The incisors, perhaps independ- that is open to question. ently, and m3 have reached the Nycticeius The Lasiurini may be regarded as having stage of reduction; the premolars are in the diverged farthest of all from the early ves- Pipistrellus stage. In the skull, extreme pertilionine bats. It is indeed possible that shortening of the face is evident, coupled they should be accorded equal rank with with enlargement of the braincase, broad- the subfamilies Kerivoulinae, Miniopteri- ening of the rostrum, broadening of the nae, etc. nasal and anterior palatal sinuses (as in The tree shown above serves to outline Nyctalus) and weakness of the zygomata. roughly the phyletic relationships of the Except the trifid nature of the lower in- bats treated in this paper and to indicate cisors almost no primitive characters re- their relationships to related subfamilies main. Possibly the mammary formula (Fig. 1). Myotini In regard to the extent of their dental MYOTIS KAIJP evolution and shortening of the face the A preliminary study of Myotis has al- bats of this division are less progressive ready been made.' than any others discussed hereafter. The basic types of the division are represented LASIONYCTERIS PETERS by Myotis and its near relatives, Pizonyx, Lasionycteris PETERS, 1865, Monatsber. Akad. Wiss. Berlin, p. 648. Cistugo and Rickettia. The American genus GENOTYPE.-Vespertilio noctivagans Le Conte. Lasionycteris, essentially a Myotis, has This monotypic genus has been compared become specialized through the loss of p3, without yet greatly shortening the face and above to Myotis. Its characters, apart jaws. The corresponding lower tooth from the loss of p3, are essentially primitive myotine. In the teeth i2, though it and P2 still agree closely with the lower premolars of less specialized species of Myotis, lacks a posterior cusp, is not reduced in not only in relation to each other but in crown height; c is simple; p2, completely their proportional sizes relative to P4. in the toothrow, and m3, unreduced. The There is no indication of displacement of lower incisors are incipiently imbricated; P2-3, as in Myotis, much smaller than p3 in Lasionycteris. One hesitates to assume derivation of P4; P3 iS completely within the toothrow. Plecotus and Corynorhinus from the Myotis In the skull the zygomata are fairly deep, processes; stock, principally because of the enormous with well developed postorbital degree of specialization of the audital re- the rostral area is somewhat broadened, gion in those genera of bats. Yet their with lateral depressions similar to those in some and There are no dental pattern, showing absence of p3 com- Myotis Eptesicus. bined with retention of Myotis-like incisors, peculiarities. is almost precisely equivalent to the tooth PLECOTUS GEOFFROY arrangement of Lasionycteris. If the four Plecotus GEOFFROY, 1818, Descirip. de 1'Egypte, plecotine genera are gathered together II, p. 112. under the collective term Plecotini, with GENOTYPE.-Vespertilio auritus Linnaeus. rank equal to the Myotini, undue notice Geoffroy's original concept was broader is accorded an admittedly important modi- than the present plecotine section, "les fication which affects only the auditory trois especes de ce genre sont, l'oreillard areas. It seems preferable that the Pleco- de Daubenton [Plecotus], la barbastelle tini be regarded instead as a specialized [Barbastella] et une nouvelle esp6ce de offshoot of the Myotis line. The fact that Timor [Nyctophilus]." Euderma has gone a stage farther than A highly specialized genus closely re- Pkecotus and has lost P3 as well as p3 does 1 1941, Bull. Amer. Mus. Nat. Hist., LXXVIII, not alter this view (Fig. 2). Art. 8, pp. 537-565. 230 Bulletin American Museum of Natural History [Vol. LXXX lated to the American Corynorhinus, Idio- Himalaya region, but members of the nycteris and Euderma. Formerly Plecotus genus extend to western Europe and east- was considered to be also a near relative of ward to Japan. Barbastella. But though it and allies may A brief comparative study of the genus be closer to Barbastella than to most other was made by Thomas,3 which I quote, vespertilionine genera and betray certain "Of the genus Plecotus, P. homochrous characters in common, they show, accord- (Nepal) and puck (Murree), doubtfully ing to Miller,' numerous important dif- distinct from each other, stand apart from ferences in the ear, rostrum, anterior part the rest owing to the narrowness of the of the skull, zygomata, bullae and teeth, braincase; auritus (Europe) has rounder

subgenera Cistugo Rickettia

Plecotus Myotis Pizonyx Corynorhinus Idionycteris

Lasionyeteris \_ / Euderma |p obsolete p3 present |p3 present | p3 absent \ ~/ Myotini (s.s,.) Plecotini Ears and bullae Ears and bullae not enlarged much enlarged

Myotini (8.1.) p2- reduced in size; p3 2-5 usually present

Fig. 2. Suggested phylogeny of the myotine and plecotine genera of Vespertilioninae. See also Fig. 1.

and are related only distantly to the South skull, small bullae, and short thumbs; American big-eared bats of the genus christiei (Egypt) a large braincase and Histiotus. very large bullae; wardi (Ladak and Less than a dozen forms of Plecotus have Kashmir) a large skull, rather small received names. G. M. Allen2, in his bullae, long thumbs, and very pale color; African list, shows only two races of P. sacrimontis (Japan) a large skull, rather auritus, reaching across northern Africa small bullae and long thumbs; and finally to Teneriffe. The Eurasian forms come the present species [ariel] has a large chiefly from the uplands adjoining the rounded skull, large bullae, long thumbs 1 1907, Bull. U. S. Nat. Mus., LVII, p. 224. 2 1939, Bull. Mus. Comp. Zool., LXXXIII, p. 96. 3 1911, Proc. Zool. Soc. London, p. 160. 1942] Tate, Results of the Archbold Expeditions. 47 231 and dark color. No doubt it is nearly allied FORE- ARM to wardi but the colors of the two are nearly FORM LOCALITY IN MM. at the opposite ends of the scale." 1. auritus auritus Europe 38 Ognevl reviewed the northern forms, auritus homochrous Himalaya 38 making wardi and sacrimontis subspecies of = puck? Punjab 38 auritus. It is quite possible that all are auritus sacrimontis Japan 39 = ognevi Japan 39 subspecies, though one gathers from 2. ariel ariel Szechwan 44 Thomas's description of mordax that it is ariel wardi Ladak 45 a well marked species. = kozlovi? Tibet 45 Recently Kishida (1927) has renamed 3. mordax Kashgar 44 sacrimontis Ognev (not sacrimontis Allen) = leucophaeus? Tashkent P. ognevi. It is unlikely that this can stand since Ognev himself (1938) places sacri- CORYNORHINUS H. ALLEN montis Ognev in the synonymy of sacri- Corynorhinus H. ALLEN, 1865, Proc. Acad. montis Allen. Nat. Sci. Philadelphia, p. 173. I have photographs of the type skulls of GENOTYPE.-PleCOtuS macrotis Le Conte. homochrous, puck, wardi and mordax and Corynorhinus is slightly more specialized specimens of auritus auritus from England. than Plecotus. The braincase is higher and An outstandingly different species is indi- fuller; the strong zygoma of Plecotus shows cated by the skull of mordax. In it the indications of reduction, and the postorbital posterior part of the mastoid area is broad- process is placed nearer the glenoid; the ened and prominent, and the relatively postpalatal spine is more accentuated; and great size of the molars, mentioned by p2 iS more reduced. On the other hand p4 Thomas, can be discerned. in Plecotus is the more specialized, being The following named forms of Plecotus narrowed, and compressed and modified have been proposed: by obsolescence of its hypocone region into a shearing blade. In the lower jaw a corresponding change of shape is seen in P4, FORE- whose crown area in Corynorhinus is wider ARM than long and in Plecotus is longer than NAMED FORM LOCALITY IN MM. wide. ariel Thomas Szechwan 44 Three full species of Corynorhinus have auritus Geoffroy Egypt(England)38 been named, all from the United States or homochrous Horsfield (Hodgson) India 38 Mexico. kozlovi Bobrinskoj Eastern Tibet 44-46 leucophaeus Swertzoff Tashkent mordax Thomas Kashgar 44 IDIONYCTERIS ANTHONY ognevi Kishida Japan Idionycteris ANTHONY, 1923, Amer. Mus. puck Barrett- Novitates, No. 54, p. 1. Hamilton Punjab 38 GENOTYPE.-Idionycteris mexicanus Anthony. sacrimontis Allen --l'Tonan 2R-Anl 7 teneriffae Barrett- This monotypic genus was distinguished Hamilton Teneriffe 44 by two fleshy lappets rising from the band wardi Thomas Ladak 45.5 uniting the ears. No generic description of the skull was made. Under the specific description the small anterior premolar It is suggested thaIt the Eurasian Pleco- was said to be "crowded" between c and p4. tus, which in this p.aper are primarily of The type is not easily available now for interest, comprise actually only three study, for it has been put away for safety species with races ancI synonyms as follows: during the war. Idionycteris was regarded as more nearly related to Old World Plecotus than to New 1 1928, Mammals of Eaistern Europe and Northern World Corynorhinus. Asia. I, pp. 593-607. 232 Bulletin American Museum of Natural IHistory [Vol. LXXX EUDERMA H. ALLEN pattern in combination with wide spacing Euderma H. ALLEN, 1892, Proc. Acad. Nat. of c and p4 in front of and behind the mi- Sci. Philadelphia, p. 467. GENOTYPE.-Histiotus maculatus J. A. Allen. nute p2, but especially by the obsolescence of Related to Plecotus and Corynorhinus. p3 (present in Plecotus, Corynorhinus and Distinguished by its remarkable color Idionycteris).

Pipistrellini In this division the anterior upper pre- were given off. In one group or another trends leading to many of the recognized molar (p2) is unstable. Reduced in size genera can be observed. Inclusive and and in process of obsolescence, it is present exclusive definition is difficult, perhaps in some genera, absent in others. In Sco- not possible. p2 present .... Pipistrellus. toecus it may be present in some species, Non-pipistrelles ...... 3. in and 3.-Pipistrelle-like bats with pipistrelle skull absent others, rarely single speci- and dentition, but with the pads of mens occur with p2 present on one side the feet and the metacarpal pad of the only. thumb greatly thickened and enlarged. Although the distinction based on the (Unlike Tylonycteris in which the skull presence or absence of this disappearing is flat and broadened. See also Hesper- optenus blanfordi.) ...... Glischropus. structure seems of doubtful validity in 4.-Pipistrelle-like bats of unusually large size distinguishing Pipistrellus-like genera from with i2-2 carried forward, rostrum not Eptesicus-like genera, I have nevertheless broadened or raised ...... Ia. adhered to the accepted practice. It will 5.-Medium to large bats, the more specialized and best known species of which have be seen that two major sections are thus high, broad rostra. Dentition as pipi- treated. The two sections, in contrast to strelles, wings generally narrow. Smaller the Nycticeini and Lasiurini, have com- species scarcely separable from pipi- plete incisive dentition, subject only to strelles ...... Nyetalus. 6.-Medium to small bats with very high, full minor modifications in a few genera. In all braincases and short, low rostra, the of them the face is considerably shortened, African members (Glauconycteris) lack- in comparison with the Myotini, and all ing p2, which is retained in Australian have lost p-. species (except C. rogersi) Chalinolobus. 7.-Medium sized bats with ear unusually The division Pipistrellini is by far the short, rostrum with markedly depressed largest and most complex in the Vespertili- median area, p2 present. Barbastella. oninae. Below I have traced out the 8.-Bats with moderately broad rostra, p2 ob- scheme of classification of its many solete, strongly developed supraorbital it is will assist in tubercles...... 9. branches, which, hoped, Very small bats with broad, very flat skulls explaining the order in which the genera are and weak tubercles, but p2 present.... 10. arranged. 9.-Braincase full, rounded ...... Philetor. 1.-Middle lower premolar retained, showing Braincase depressed (African).. Mimetillus. primitive origin. Genus otherwise with 10.-Enlarged pads on feet and base of thumbs. pronounced specializations of wings and .....Ty..lo.n.y.c..te.r..is...... feet. Face elongate...... Discopus.1 11.-Braincase full as in Philetor; no supraorbi- Middle lower premolar obsolete, a speciali- tal tubercles, i2 placed directly behind zation. All species showing greater or i...... Hesperoptenus. less shortening of the face. Various 12.-Skull with full rostrum and wide anterior special adaptations in certain of the palatal sinus very much as in Nyctalus, genera ...... 2. but p2 obsolete. Otherwise much as 2.-The large assemblage Pipistrellus, com- Eptesicus fuscus ...... Vespertilio. prising a dozen major species groups, of 13.-Small species, Pipistrellus-like but lacking quite wide morphological variability, is p2; and large species with skulls some- assumed to be the complex from which what resembling Myotis myotis in outline most, perhaps all, succeeding genera but with reduced dentition (premolars, 1 One may ask why Discopus is not placed with ... Eptesicus. . Lasionycteris in the Myotini. I have been influenced 14.-Small African bats with somewhat elevated by the fact that wing and foot disks have hitherto skulls and ears.. been found only in the Pipistrellini, except, of course, Eptesicus-like enlarged the family Thyropteridae...... Laephotis. 19421 Tate, Results of the Archbold Expeditions. 47 233

15.-Larger South American bats with enormous ment of wing and foot disks, leads me to ears, skulls with high rostrum and braincase, and Eptesicus-like dentition...... place Discopus far back along the phyletic ...... Histiotus. line leading to Pipistrellus and allies. Its dental formula equals that of Lasionycteris DiscoPus OSGOOD which I regard as derived from Myotis, Discopus OSGOOD, 1932, Publ. Field Mus. Nat. Hist., Zool. Ser., XVIII, p. 236. specialized through the loss of p3. GENOTYPE.-Discopus denticulus Osgood. Discopus denticulus Osgood Hind feet with highly developed Discopus denticulus OSGOOD, 1932, Publ. Field disk-like pads even more extreme than in Mus. Nat. Hist., Zool. Ser., XVIII, p. 236.

Fig. 3. Suggested phylogeny of the pipistrelloid genera. See also Fig. 1. Tylonycteris and Glischropus. . . . Skull TYPE LOCALITY.-Phong Saly, Laos, Indo- with broad, greatly flattened braincase, China. somewhat as in Tylonycteris but with a Osgood described the color as "Cinnamon longer, narrower rostrum; dentition with Brown, the hairs unicolored from base to two upper and three lower premolars on tip; underparts near Amber Brown, the each side, the middle lower pair small and hairs with darker bases." Forearm, 37.8; somewhat internal to the toothrow." c-im3, 5.4 mm. A single primitive character, retention PIPISTRELLUS KAUP of P3, in combination with marked speciali- Pipistrellus KAUP, 1829, Skizzirte Entwickl.- zations, flattening of the skull and develop- Gesch. u. natuirl. Syst. d. europ. Thierw., I, p. 98. 234 Bulletin American Museum of Natural History [Vol. LXXX

Romicia GRAY, 1838, Mag. Zool. Bot., II, p. canines should lack accessory cusps; p2 495. Vesperugo KEYSERLING AND BLASIUS, 1839, should not be smaller than i2, and it should Archiv. f. Naturg., V, Part 1, p. 312 (part, with be slightly or not at all, displaced inward Eptesicus). from the tooth row, leaving an obvious Hypsugo KOLENATI, 1856, Allg. deutsch. diastema between c and p4; P2 should be naturh. Zeitung, (n.f.) II, p. 131. Nannugo KOLENATI, 1856, loc. cit. only slightly smaller than P4 and retained Megapipistrellus BIANCHI, 1916, Annuaire Mus. in the toothrow. In all species of Pipi- Zool. Acad. Sci. Petrograd, XXI, pp. lxxiii- strellus, m3 remain unmodified. In reality lxxxii. one or more of these characters is changed. GENOTYPES OR REPRESENTATIVE SPECIES.- Pipistrellus, Vespertilio pipistrellus Schreber. Various partial treatments of the pipi- Romicia, calcarata Gray = kuhlii Kuhl. Ve8- strelles have been published. The follow- perugo, several species of Pipistrellus and Eptesi- ing is an incomplete list of papers in which cus were included. Hypsugo, maurus (= savii) Pipistrellus has been treated in some de- and kracheninikowii listed. Nannugo, nathusii, tail: pipistrellus, kuhlii included. 1866, Gray, Ann. Mag. Nat. Hist., (3) XVII, p. Kaup's brief characterization of Pipi- 90. Diagnosis of "Vesperugo." strellus reads as follows, "Bats with short, 1878, Dobson, Cat. Chiropt. Brit. Mus., pp. ears with bases far short 183-246. Under "Vesperugo" Dobson in- simple apart, cluded both Pipistrellus and its close allies, tragus arched inwards, and 34 teeth." and under his subgenus Vesperugo came When Kaup set up Pipistrellus, only two Nyctalus, Pipistrellus and Glischropus. He species, pipistrellus and Vespertilio kuhlii dealt with about twenty-five species. had been described. The two 1897, Miller, North American Fauna, XIII, pp. Kuhl, 1819, 87-95. Revision of North American Pipi- are very easily distinguished, as shown strellus. by Miller's (1912) key and figures of denti- 1907, Miller, Families and Genera of Bats, pp. tions. Subsequently savii Bonaparte and 204-205. Pipistrellus was characterized in de- nathusii Keyserling and Blasius were tail, and a few of the species were mentioned. 1912, Miller, Mammals of Western Europe, pp. described from Europe, a few species from 202-224. The genus Pipistrellus and its four America, and a wealth of species and races European species were treated at length. from the Old World tropics and subtropics. 1916, Bianchi, Annuaire Mus. Zool. Acad. Sci. Pipistrellus is a genus of short-faced Petrograd, XXI, pp. lxxiii-lxxxii. Mega- pipistrellus, with type V. annectans Dobson, Vespertilioninae in which the dental for- was proposed. mula is i3, p2. This number of incisors and 1917, Wroughton, J. Bombay Nat. Hist. Soc., premolars it shares in common with Ia, XXV, pp. 588-592. A synoptic list of the Nyctalus and Glischropus; also Barba- Indian pipistrelles. 1928, Ognev, Mammals of Eastern Europe and stella, Euderma and Chalinolobus. The Northern Asia, I, pp. 480-502. Much de- first three mentioned are probably detailed information concerning the boreal scended from common ancestry with species of Pipistrellus. Pipistrellus; the last three are different in 1928, Thomas, Proc. Zool. Soc. London, p. 144. Discussion of "Pipistrellus" tralatitius and P. various ways. The African forms of imbricatus. "Scotozous" have been treated by Hollister 1934, Iredale and Troughton, Mem. Australian as a group of Pipistrellus. Glischropus Mus., VI, p. 96. Check list of Australian is separable chiefly through its specialized Pipistrellus. Ia and 1934, Taylor, Philippine Land Mammals, pp. thumbs and foot-pads. Nyctalus 294-301. Four Philippine pipistrelles are evince minor peculiarities of size, teeth, recognized in this manual. rostrum and ears. 1935, Phillips, Manual of Mammals of Ceylon, No uniformly unspecialized pipistrelle pp. 109-118. A manual which includes four incisors or the canines or species of Pipistrellus occurring on the island of exists. Either the Ceylon. the premolars have in every species ac- 1938, Allen, G. M., Mammals of China and quired peculiarities. Ideally the upper in- Mongolia, Part 1, pp. 226-231. cisors should be subequal in size, and each 1940, Chasen, Bull. Raffles Mus., XV, pp. 50-51. should have its accessory cusps well set Synoptic list of twelve Malayan Pipi8trellus. off; the lower incisors should be scarcely Somewhat arbitrarily I have condensed thickened and in linear arrangement; the the many "species" of Pipistrellus into 19421 Tate, Results of the Archbold Expeditions. 47 235 fourteen species groups.' These groups Pipistrellus tasmaniensis group. Forearm, 47; braincase and skull low; zygomata weak; p2 cannot be arranged in simple gradient from much reduced; il unicuspid. One species. primitive to modified, but instead present Pipistrellus annectens group. A single large a complex of archaic and specialized struc- species, characters unknown. Forearm, 45. tures which makes their arrangement at Subgeneric name, Megapipistrellus Bianchi. Pipistrellus minahassae group. Forearm, 35; best unsatisfactory. Thus species with braincase full, with weak sagittal crest; P2 strong zygomata and with traces of post- as large as p4. One species. orbital processes on the zygomata, and with Pipi8trellus riippellii group. Chiefly African. not specially reduced, have been re- Brainease very full; zygomata weak; under- P. parts white. These are the African "Scoto- garded as relatively primitive. But other ZOUs." species occur in which one or the other of Pipistrellus joffrei group. Specialized pipistrelles those characters has remained primitive, with markedly developed supraorbital tuber- while the other has been strongly modified. cles and very minute p2. Formerly referred to For example, P. macrotis with the zygoma Nyctalus. Improbably truly annectant be- deep, strong, and provided with a strong tween Pipistrellus and Nyctalus. postorbital process, yet with p2 reduced to Under each group I have gathered vari- a mere spicule, or P. abramus in which ous named forms which I believe are refer- the zygoma is almost obsolete, but p2 iS able to it. In some, perhaps many, in- still fully as large as i2. Other such inde- stances the names should go into synonymy. pendently fluctuating characters are numerous. Therefore, though it has been hoped that the species groups might be Pipistrellus abramus Group arranged progressively from unspecialized The bats of this group have the skull to specialized, in practice exceptions will with profile sloping uniformly from brain- be numerous. The groups dealt with are case to rostrum; wide, smooth, flat ros- shown synoptically: trum; very weak zygomata; braincase wide Pipistrellus abramus group. Forearm, 32-35; and smooth; no basial pits; unshortened skull broad; p2 unreduced; zygoma weak. premaxillae. Anterior upper premolar, Pipiatreflu8 pipidtrellus group. Forearm, 30-32; though displaced inward, not reduced in skull slender; p2 unreduced; frons depressed; zygoma weak. size, its area about equal to that of i2; il Pipi8trellu8 coromandra group. Forearm, 30-32; distinctly bifid; canine with posterior skull slender; p2 unreduced but displaced in- cusp; c and p4 not in contact; lower inward; zygoma weak. cisors somewhat imbricated. Baculum Pipistrellus tenuis group. Forearm, 27-30; skull broadened; face short; premaxilla short; p2 very large. Forearm usually 33-35 mm. unreduced; zygoma weak. Color of dorsal pelage grayish brown. Pipistrellus affinis group. Forearm, 35-38; The group extends from Japan and skull long, low; p2unreduced; zygomastrong, with postorbital process present. China south to Malay, Banka, Java, Pipistrellus ceylonicus group. Bats chiefly of Sumatra, the Nicobar Islands, and thence the Indian peninsula, much like the afftnis west to Burma and (?) northern India. group but with weak zygomata which lack the Taylor (1934) recorded it from the Philip- postorbital processes. Pipi8trellus kuhlii group. Forearm, 33-35; pines. skull long, low; p2 slightly reduced; zygoma Forms referred to the abramus group aiO: weak; wings often pale. abramus, akokomuli ( = abramus ?) ir- Pipistrellus savii group. Forearm, 32-34; skull typically long; p2 greatly reduced; zygomata retitus, pumiloides, bancanus, camortae. strong, with processes. Also oriental species P. abramus, with type locality at the in which braincase is rounded and face shorter. northern edge of the geographical range of Pipistrellus circumdatus group. Forearm, 40; a whole, may not be braincase rounded; p2 reduced; zygomata the group taken as strong but without process. One species. wholly representative. It seems probable that ultimately the 1 This excludes African groups, unstudied. One forms now named as species of this group of them is probably represented by Eptesicops Rob- will be in part synonymized and in part erts (type, rusticus), another by Scotozous (African only). reduced to subspecies of abramus. 236 Bulletin American Museum of Natural History [Vol. LXXX

Pipistrellus abramus (Temminck) "o" to "s" of the abramus series' as "un des Vespertilio abramus TEMMINCK, 1835, Monogr. types du Vespertilio akokomuli." Speci- Mamm., II, p. 232. mens "o" to "r" of the same series were TYPE LoCALITY.-Nagasaki, Japan. entire in alcohol. Temminck gave the size of abramus as Of specimen "b" I noted, "Ears rather "a little smaller than the pipistrelle . . . " narrow; tragus round-tipped; skull mod- and the forearm length as 1 inch 2 lines erately deep; profile almost flat from ros- ( = 29.6 mm.). On the other hand Dob- trum to occiput; interorbital region son recorded the forearm as 1.35 inches broad...." But subsequently study of (34.5 mm.). The animals we treat today the skull reputed to belong with the skin and as abramus have dimensions much as Dob- of measurements made of it shows that it son stated. is not a Pipistrellus but a Nyctalus skull. In Leyden in 1937, I examined and The size of skull "b" is very nearly equal photographed the cotype, specimen "d" to that of N. velutinus of China, whose of the Jentink catalogue, an adult male forearm attains a length of 52 mm. It is whose skull Dr. Junge had extracted for obvious therefore that the skull in question me. From notes made then, "Profile of had nothing to do with akokomuli Tem- skull nearly flat; palate strongly domed; minck. The skin, on the contrary, may rostrum broad and flattened on top; i2 well be correctly marked, in which case three-quarters of height of il; p2 at inner akokomuli should still come within the edge of toothrow between canine and p4, synonymy of abramus. its height about one-half that of p3; canine with posterior cusp; P2 three-quarters of Pipistrellus irretitus (Cantor) height of P42" The forearm of that speci- Vespertilio irretitu8 CANTOR, 1842, Ann. Mag. men measured 33 mm. For skull measure- Nat. Hist., IX, p. 481. ments see table. TYPE LOCALITY.-Chusan, eastern China. Two topotypes (U.S.N.M. 140923, Cantor gave very little data. Color 140935) agree substantially with the type, "gray-brown, beneath dust-colored . . . Leyden "d," in all characters, as do the forearm 1.3 inches" (= 33 mm.). considerable series of specimens from many The type was a male. Our photograph of the eastern and southern Chinese prov- of the type skull shows the rather large inces which were listed by G. M. Allen in bullae in place and the characteristically "The Mammals of China and Mongolia"; broad rostrum and braincase of the abramus also three specimens from Chiunglung, group. The approximation of canines (see upper Salween River. The specimen from table) and of il-' may be due to crushing, the Philippines referred to by Taylor is although no sign of bad treatment shows still in our collections. in the photograph. The animals from North China (Wei- Irretitus is apparently the mainland rep- sien), the Salween River and the Philippine resentative of the Japanese abramus. The Islands have very slightly larger skulls than type, as shown in the table of measure- have those from Japan. Otherwise no ap- ments, is slightly smaller than the type preciable difference can be detected. of abramus. Pipistrellus pumiloides (Tomes) [Pipistrellus akokomuli (Temminck) ] Scotophilus pumiloides TOMES, 1857, Proc. Vespertilio akokomuli TEMMINCK, 1835, Zool. Soc. London, p. 51. Monogr. Mamm., II, p. 233. TYPE REGION.-China. TYPE LOCALITY.-Japan. Dobson made this form from "China" Temminck's description hardly suffices to a synonym of abramus, but the skull, separate akokomuli from abramus, and by though closely related, is uniformly smaller. Dobson and others it has been treated as A considerable series from Hainan Island, a synonym. Jentink so it and regarded I 1888, Cat. Systematique des Mammiferes, Mus. catalogued each of specimens "b," and d'Hist. Nat. des Pays-Bas, XII, p. 180. 1942] Tate, Results of the Archbold Expeditions. 47 237 identified as "tralatitius pumiloides," is It is not possible to learn from Chasen's without much question referable to the Handlist' which of his pipistrelles be con- abramus group. Forearm of type, 34 mm. siders the representative of P. abramus in The skull measurements of the type, Java, but we may limit the possibilities to shown in the table, suggest that pumiloides three: brachypterus, "imbricatus" and was slightly smaller than the type of "tralatitius." I have pointed out beyond irretitus and abramus. But on the other that due to a mix-up of the type skull at hand the animal is virtually inseparable Leyden the status of brachypterus is un- from topotypes of abramus. In A.M.N.H. satisfactory; imbricatus, based on measure- 84842 of the type series, the large size of ments of the type skull, I believe allied to the baculum, which seems to characterize coromandra, subulidens, vordermanni, etc.; the entire group, can be observed easily. and I believe tralatitius, with its "sooty" pelage and "six" lower molars (Horsfield) Pipistrellus paterculus Thomas to be a Myotis. There remains then ac- Pipistrellus paterculus THOMAS, 1915, J. Bom- tually no named form of abramus with type bay Nat. Hist. Soc., XXIV, p. 32. locality in Java, the nearest being bancanus TYPE LOCALITY.-Mt. Popa, Upper Burma. from Banka, whose type I have studied A small representative of the abramus and photographed and which I am thor- group, with the baculum proportionally oughly satisfied represents abramus. (It as long. The rostral width broad, as with may be synonymous.) A series of twelve abramus. The canine width, however, is skins and skulls in the Archbold Collections less, reminding one of the Chinese irretitus. from Cheribon, Java, is therefore referred p2 equals i2 in area, is half internal and to P. abramus bancanus (Sody). visible externally. Possibly the specimens Sody gave the length of upper toothrow from the Salween River, mentioned under in his series as 4.4-4.7 mm. abramus, belong here. Pipistrellus camortae Miller Pipistrellus pipistrellus Group Pipistrellus camortae MILLER, 1902, Proc. U. Relatively unspecialized as to dentition; S. Nat. Mus., XXIV, p. 779. zygoma weak and without indication of TYPE LoCALITY.-Nicobar Islands. postorbital process. Braincase full, ros- Miller contrasted this species with trum but little shortened as shown by the "abramus" of Java, showing that it had a considerable diastema between canine and broader rostrum. It may be a relatively P4; posterior canine cuspule present but unspecialized member (as suggested by weak; p2 equal in area to i2, placed inward the size of the baculum-very large in true from diastema but in contact with canine abramus) of the abramus group. and p4. Lower incisors very little imbricated; P2 three-quarters of height of P4. Pipistrellus bancanus Sody Here belong the common pipistrelle of Pipistrellus tralatitius bancanus SODY, 1937, Europe and its subspecies; also P. nathusii, Temminckia, II, p. 233. distinguished by its longer canines and TYPE LoCALITY.-Banka, Dutch East Indies. slightly compressed P2 The photographs before me of Sody's Compared with abramus, the type species type skull marked "39 BK" leave no room of the genus, P. pipistrellus of Europe is for doubt that it belongs with the abramus generally smaller (forearm, 31 mm.), and group. It has the same uniformly shelving has the basal phalanx of the thumb shorter, profile, the same breadth of rostrum and 2.2 mm., and the calcareal lobe better de- of braincase, with substantially wide veloped. The dentition differs slightly in mastoids and weak zygomata. that the canines are less widespread, p2 iS [Horsfield's dental formula for tralatitius, considerably smaller and its cusp only two- molars 6-6 (P3 doubtless minute and con- thirds the height of i2, and the toothrows cealed), seems to indicate that tralatitius was a Myotis.] 1 1940, Bull. Raffles Mus., XV, pp. 50-51. 238 Bulletin American Museum of Natural History [Vol. LXXX

arm, 33-35 mm.). Skull with frontal region are convergent in c-c= 3 cm3 front; m 3 -3 5.1'cr, slightly depressed. Dentition very similar 4.4. to that of pipistrellus but with height of lower canine to height of P4 about as 5:4; P2 Pipistrellus pipistrellus (Schreber) compressed and slightly displaced. Vespertilio pipistrellus SCHREBER, 1774, Shuge- This species appears to be restricted to thiere, I, PI. LIV; description, 1775, I, p. 167, central under name Zwergfledermaus. and southern Europe (Miller, TYPE REGION.-France. 1912). Ognev (1928) showed that it ex- A rather small species, with forearm, tended across Russia to the Ural Moun- 29-32 mm. Color brown, near Snuff tains. Whether records from Japan and Brown (bases fuscous), underparts slightly Baluchistan are valid is uncertain. paler. The European races P. n. unicolor Fatio Skull with narrow but rather high brain- and mediterraneus Cabrera have been de- case and low rostrum, a distinct frontal scribed. depression, in profile. Zygomata weak Pipistrellus coromandra Group and without traces of postorbital proc- This group comprises a number of small esses; rostrum unwidened; no basial pits. species, perhaps nearest to the P. pipistrel- Dentition: iP with distinct posterior cusp; lus group of the west but with less depressed i2 as long as posterior cusp of il; canine rostrum and less full braincase. The mod- with trace of posterior cuspule; area of p2 erately expanded zygomata are weak as in equal to that of i2; p2 partly displaced in- the pipistrellus and abramus groups. Den- ward but diastema between c and p4 not tition virtually as in pipistrellus, i.e., with closed. Lower incisors but slightly im- p2 little reduced and the lower incisors bricated; lower canine slightly exceeding scarcely imbricated. The approximation P4 in height; P2 three-quarters the height of the canines and the small expanse of the of P4 and almost in toothrow. lacrimal region give these bats somewhat Various European races have been set pointed muzzles (compare with tenuis or up, but Miller (1912) placed all in the syn- abramus group). A posterior canine cusp onymy of pipistrellus. is usually present. These bats are distinguished from char- Pipistrellus pipistrellus bactrianus acteristic members of the tenuis group by Satunin their unshortened premaxillae, relatively Pipistrellus bactrianus SATUNIN, 1905, Mitt. Kaukasus Mus., II, pp. 67, 85. narrower palates and less globular brain- TYPE LOCALITY.-Oasis of Tedzen, Trans- cases. But the New Guinea forms papu- caspia. anus, angulatus and collinus bridge the gap. Satunin regarded this bat as closely re- They are apparently annectant between lated to P. pipistrellus, but his statement the coromandra and tenuis groups of Pipi- that the margin of the wing and parts of the strellus. phalanges are whitish causes one to remem- Typically coromandra and allies are quite ber that in the kuhlii group whitening of small bats (forearm, 30-32 mm.) with un- the wings is prevalent. Ognev (1928), reduced p-, premaxillae not especially however, who reported obtaining more shortened, weak zygomata without post- material from Turkestan, retains bactrianus orbital processes. The skull, which shows with pipistrellus. Forearm, 34 mm. the frontal step from rostrum to braincase We have no specimens. less abrupt than in the P. pipistrellus group, is thinly ossified as in pipistrellus. On the Pipistrellus nathusii (Keyserling and other hand the bats of this group show Blasius) some relationship to the abramus group. Vespertilio nathusii KEYSERLING AND BLASIUS, The latter has the skull much more 1839, Archiv. f. Naturg., V, Part 1, p. 320. heavily constructed, the rostrum broader TYPE LoCALITY.-Berlin, Germany. and flatter and the zygomata more ex- Slightly larger than P. pipistrellus (fore- panded. 1942] Tate, Results of the Archbold Expeditions. 47 239

P. regulus, in which the braincase is ex- the abramus group extends as far as west- traordinarily flattened and low, is a special- ern India. ized Australian offshoot of the coromandra group. Perhaps it should be separated as Pipistrellus coromandra (Gray) a wholly distinct group. Vespertilio de Coromandel F. CUVIER, 1832, The coromandra group has a very ex- Nouv. Ann. Mus. Paris, I, p. 21. tensive distribution, reaching from Persia Scotophilus coromandra GRAY, 1838, Mag. Zool. Bot., II, p. 498. (and perhaps farther west), through India TYPE LoCALITY.-Pondicherry, Coromandel to South China and east to the Bonin group Coast, South India. of islands, the East Indies, Philippines and Cuvier described a "noctuloid" skull New Guinea. (with full braincase and lower rostrum), A number of species have received names, ears "6chancr6es," tragi "en couteau" (as many of them perhaps being synonyms. in Pipistrellus); upperparts "brown-gray- Arranged from west to east and north to yellowish," lower parts whitish, the hairs south they are listed as follows: dark for three-quarters of their length, and aladdin Thomas, 1905 Persia pale yellow at the extremities. Body micropus Peters, 1872 Northwest India coromandra Gray, 1838 South India length, 34 mm., tail length, 27 mm. tramatus Thomas, 1928 Tonkin Gray merely replaced Cuvier's Vesper- portensis J. A. Allen, 1906 Hainan tilio de Coromandel with the name coro- sturdeei Thomas, 1915 Bonin, southeast mandra. of Japan ridleyi Thomas, 1898 Selangor, Malaya Wroughton' set weak definitions for the subulidens Miller, 1901 Natuna Islands species. According to him the forearm is murrayi Andrews, 1900 Christmas Island 29-33 mm., tragus less than 4 mm. broad, imbricatus Horsfield, 1824 Java outer margin of ear below tip straight, p2 meyeni Waterhouse, 1845 Philippines collinus Thomas, 1920 New Guinea entirely inside toothrow, outer incisor as angulatus Peters, 1880 Duke of York Is- high as outer cusp of inner incisor and skull land relatively long (12.5 mm.). The last two ponceleti Troughton, 1936 Bougainville Is- characters were used to separate coro- land mandra from members of the short-faced Pipistrellus aladdin Thomas tenuis group. Pipistrellus aladdin THOMAS, 1905, Proc. Zool. By elimination, coromandra can be ap- Soc. London, p. 521. plied only to the species for which it is now TYPE LoCALITY.-Derbeuch, Persia. -used or to mimus. Wroughton has already Thomas compared aladdin with mimus of made the selection. Phillips2 has followed the tenuis group and with the African Wroughton's lead and lists only two small nanus. The skull is delicate, with the species, coromandra and mimus, from Cey- premaxillae normal as in others of the lon. coromandra group, not shortened as in I cannot agree with Wroughton (ibid., p. mimus. p2, though small, remains in the 588) that "the Indian representative of toothrow. Forearm, 31 mm. abramus Temminck (from Japan) is coromandra." As shown under the abramus Pipistrellus micropus (Peters) group those bats are larger and stronger, Vesperugo micropus PETERS, 1872, Proc. Zool. with the top of the rostrum broad and flat. Soc. London, pp. 707-708. TYPE LoCALITY.-Dehra Doon, near Simla, northwest India. Pipistrellus tramatus Thomas Peters gave very little on Pipistrellus coromandra tramatus THOMAS, information 1928, Proc. Zool. Soc. London, p. 144. this bat. "Brown above, gray-brown TYPE LOCALITY.-Tonkin, French Indo-China. beneath; tibia 7/16 (14 mm.)"; forearm It is very possible that tramatus, and length not indicated.- portensis from Hainan, are identical. Dobson treated it as a synonym of abramus. It is now placed provisionally 1 1918, J. Bombay Nat. Hist. Soc., XXV, pp. 589, 592. next to aladdin, as I am not convinced that 2 1935, Manual of Mammals of Ceylon, p. 113. 240 Bulletin American Museum of Natural History [Vol. LXXX

Tramatus is now treated as a species merely 'a.,' irretitus Cantor." However, irretitus because of my uncertainty about it. Per- really belongs with abramus and is de- haps all of the forms in this group should cidedly larger than meyeni. Taylor treats be regarded as subspecies of coromandra. "irretitus" (= meyeni ?) as a full species Forearm, 29.5 mm. with forearm 27.5 mm. and places meyeni in its synonymy. I measured the skull of Pipistrellus portensis J. A. Allen a cotype of irretitus (see table) which is a Pipistrellus portensis J. A. ALLEN, 1906, Bull. much larger animal than is shown in Amer. Mus. Nat. Hist., XXII, p. 487. Taylor's measurements of the type of TYPE LoCALITY.-Porten, Hainan, China. meyeni. Allen stated that portensis was "similar in form of ear and structure of incisors and Pipistrellus subulidens Miller premolars" to tenuis Temminck, as de- Pipistreltus subulidens MILLER, 1901, Proc. scribed by Dobson, but much larger. Wash. Acad. Sci., III, p. 134. "Quite different from ridleyi Thomas, TYPE LoCALITY.-Sirhassen, Natuna Islands. which ... is much smaller ... and has Miller wrote "inner upper incisor with- quite different ears and dentition .... It out supplemental cusp." He compared differs from P. abramus in smaller size ...." subulidens chiefly with P. pipistrellus. We have a good series of paratypes as Forearm of type, 32.4 mm. well as the type specimen. Forearm, 32 Mr. Miller kindly lent me paratypes of mm. subulidens to study. In one of them (U.S.N.M. 104756) the accessory cusp of Pipistrellus sturdeei Thomas il is present. In every other way the skull Pipistrellus sturdeei THOMAS, 1915, Ann. Mag. to me characteristic of the Nat. Hist., (8) XV, p. 230. and teeth appear TYPE LoCALITY.-Hillsboro Island, Bonin concept of the coromandra group which I Islands, southeast of Japan. have tried to express. The zygomata are Skull narrow and delicate. Teeth "as in proportionately a little deeper than in abramus but smaller"; p2 equal in area to pipistrellus and abramus. p2 iS unreduced. i2; P2 three-quarters height of p4; forearm, 30 mm.; c-m3, 4.2; diastema between c Pipistrellus ridleyi Thomas and p4. Pipistrellus ridleyi THOMAS, 1898, Ann. Mag. My photographs of the type of P. Nat. Hist., (7) I, p. 361. sturdeei indicate its near relationship not TYPE LOCALITY.-Selangor, Malaya. to abramus but to coromandra, portensis and Thomas compared this bat with abramus tramatus. It thus marks the extreme north- but remarked upon the markedly shorter eastern limit of range of its group. Repre- forearm (28 mm.), muzzle narrower and less sentatives of the coromandra group should flattened above, and short upper incisors. be present on the Liu Kiu Islands. P. p2 in toothrow, unreduced. meyeni probably represents it in Luzon. My photographs of the type skull clearly show the low narrow rostrum, narrow Pipistrellus meyeni (Waterhouse) canine width and narrow lacrimal width. Vespertilio meyeni WATERHOUSE, 1845, Proc. Detailed measurements are given in the Zool. Soc. London, p. 7. table. TYPE LoCALITY.-Luzon, Philippine Islands. Very little data can be drawn from Pipistrellus murrayi Andrews Waterhouse's description: forearm, 28 Pipistrellus murrayi ANDREWS, 1900, Monogr. mm.; color rufous-fuscous, hairs white at Christmas Island, I, p. 26. base, beneath grayish; foot short and TYPE LOCALITY.-Christmas Island, 250 miles broad. south of Java. The original description could serve Type specimen not seen. Smaller than equally well for a member of the tenuis abramus, near size of tenuis. Andrews group and perhaps does so. Dobson treated states that the muzzle is less obtuse than meyeni as a synonym of "abramus, var. that of abramus. The muzzle in tenuis 19421 Tate, Results of the Archbold Expeditions. 47 241 is distinctly obtuse. Forearm, 30-32.5 be races of papuanus, which appears, on mm.; p2 visible between c and p4, therefore account of its shorter rostrum and palate probably unreduced. Color brown with and smaller size, to belong in the tenuis yellowish tips. group. Pipistrellus imbricatus (Horsfield) Pipistrellus collinus Thomas Vespertilio imbricatus HORSFIELD, 1824, Zool. Pipistrellus papuanus collinus THOMAS, 1920, Researches in Java. Ann. Mag. Nat. Hist., (6) IX, p. 533. TYPE REGION.-Java. TYPE LoCALITY.-Bihagi, head of Mambari Horsfield described V. tralatitius and V. River, Papua. imbricatus, comparing them. V. tralati- This species, with forearm reaching 36 tius, with "molares :1:," appears to have mm., is the largest member of the group. been a species of Myotis'; V. imbricatus, The canine width and lacrimal width are with "molares 4-5,V to have been a Pipis- unusually great and suggest transition to trellus. There exists in the British Museum the abramus group. collections the supposed type of imbricatus, B.M. 79.11.21.108, the skull in poor condi- Pipistrellus ponceleti Troughton Pipistrellus ponceleti TROUGHTON, 1926, Rec- tion. From photographs of it the measure- ords Australian Mus., XIX, 5, p. 351. ments are: canine width, 4.0; c-iM3, 4.4; TYPE LoCALITY.-Bougainville Island, Solo- ml-3, 3.0; breadth of braincase, 6.5 can be mon Islands. taken. These measurements agree closely Troughton compared ponceleti with with those of the coromandra group but do papuanus, abramus and angulatus from not agree with either the abramus or the Duke of York Island. He gave the fore- tenuis group. It seems probable that imbri- arm length of ponceleti as 32-33.8 mm.; catus was the Javanese representative of the c-m3, 4.3-4.4. coromandra group. If so, imbricatus would have to replace coromandra as the group Pipistrellus regulus Thomas name, a step which I hesitate to take until Pipistrellus regulus THOMAS, 1906, Proc. Zool. an opportunity for further close comparison Soc. London, p. 470. of the several type specimens arises. In TYPE LOCALITY.-King River, King George's case it can be shown that imbricatus and Sound, southwest Australia. coromandra really belong in the same This strongly specialized species seems species-group, it will probably be found to have no very close allies. It is dis- that one of the species nearby geographi- tinguished by its extremely low flattened cally is a synonym of imbricatus. skull. The zygoma is weak and without Horsfield stated that the pelage of im- postorbital process. p2 about one-fourth bricatus was "brown with a fulvous luster." of the area of i2; canine with supplementary cusp. No basial pits. Pipistrellus angulatus (Peters) Vesperugo angulatus PETERS, 1880, Sitzungs- Pipistrellus tenuis Group ber. Ges. Naturf. Freunde, p. 122. TYPE LoCALITY.-Duke of York Island, be- The bats of this group include the small- tween New Britain and New Ireland. est of the species of Pipistrellus. Pre- Peters gave the pelage color as red- maxillae exceptionally short. Zygomata brown; forearm, 34; ear "rounded as in weak, the braincase full and rather short, maurus (= savii)." I have not seen the with rather steep frontal region and low type. It seems likely that the subsequently rostrum, somewhat as in pipistrellus. described collinus and ponceleti must be Some degree of broadening of the muzzle closely related, perhaps are subspecies. contributes to give the palate an appear- I do not believe that any one of them can ance of unusual wideness and shortness. Forearm length, from 27 to 30 mm.; p2 1 Thomas disregarded Horsfield's published molar formula for tralatitius and accepted the "type" as somewhat reduced, about one-half of area valid. See Proc. Zool. Soc. London, 1928, p. 144. See of i2. also this paper under P. bancanus and again under P. tenuis. It seems possible that V. tylopus Dobson, 242 Bulletin American Museum of Natural History [Vol. LXXX separated by Peters to form the genus 1937, Dr. Junge kindly had a number of Glischropus, was derived from ancestry skulls cleaned for me to study. Among common to it and to the P. tenuis group. them were the two cotypes of V. tenuis It differs chiefly from P. tenuis by the un- from Sumatra mentioned by Jentink2 as usual development of thumb-pads and specimens "a," "b," "types de l'espece." foot-pads. These specimens proved to belong to two The following named forms, arranged different genera. Specimen "a," with three geographically from west to east, are here upper and lower premolar teeth, was a referred to the P. tenuis group: Myotis, close to M. macellus, of which I glaucillus Wroughton, 1912 Northwest India also examined cotypes. Specimen "b," mimus Wroughton, 1899 Near Bombay, premolars 2 was a Pipistrellus. western India In the past V. tenuis has been regarded principulus Thomas, 1915 Assam as one of the pipistrelles. Jentink (tom. cit.) tenuis Temminck, 1835 Sumatra nitidus Tomes, 1858 Labuan, northwest placed it in Vesperugo, and Thomas has Borneo treated it as a species of Pipistrellus. It papuanus Peters and Doria, Salawatti, Nether- becomes necessary therefore to restrict the 1881 lands New Guinea basis of the specific concept of V. tenuis orientalis Meyer, 1899 Papua Temminck to specimen "b" of Leyden Museum. The more typical members of the tenuis My photograph of the type (skull "b," group (tenuis, principulus and nitidus of Leyden) is poor. The skull was in frag- the Malay-Sunda region) seem to be con- ments when cleaned for me. Nevertheless nected morphologically with the coro- the few measurements made it possible mandra group through the Papuan form for me to indicate its relationships. papuanus (= orientalis?). In my notes I find, 'p2 in contact with On the other hand the specialized (as re- inner margins of c and p4, half of height of gards shortening of face and palate) west- p4; P4 slightly higher than p2; canine ern Indian forms mimus and glaucillus scarcely higher than P42" Measurements: represent a special offshoot. forearm, 30 mm.; hind foot (s.u.), 6.1; It should be noted that the North Ameri- c-mi3, 2.5; crown areas, ml, 1.0 X 1.0; can P. hesperus group, recently revised mi2, 0.9 X 1.1; mi, 0.6 X 1.1; ln,-3, 3.0. by Hatfield,' and the African P. musciculus In addition, in the British Museum are appear to find nearest relatives in the two more specimens marked "cotypes"'3 tenuis group. P. subflavus of the eastern obtained by Tomes from Leyden Museum: United States is quite unlike any member B.M.7.1.1.407-408, the former without of this group. It has the height of crown braincase, the latter in quite good condi- of p2 one-third of height of p4, and the tion. Both are from Sumatra. Their lower incisors spaced, not imbricated. forearms measure, respectively, 26 and 27 mm. I have no further notes concerning Pipistrellus tenuis (Temminck) them. But their photographs are fairly Vespertilio tenuis TEMMINCK, 1835, Monogr. clear. Distinctive features are: shortness Mamm., II, p. 229. of the premaxillae, which places the upper TYPE LOCALITY.-Sumatra. incisors only slightly in advance of the Temminck compared tenuis with "tralati- canines; peculiarly narrowed U-shaped tius" (not with V. tralatitius of Horsfield notch, whose width is only 0.7 mm., at base which was a species of Myotis, but with a of narial sinus; canine width, c-mi = pipistrelle apparently closely related to P. 3.3:3.8 (measured on photograph, in which coromandra). Tenuis, he wrote, had millimeter scale was included); width "shorter muzzle, . . . ears shorter, less across supraorbital processes, 4.0; breadth broad, more pointed, . . . braincase smaller of braincase, 6.6. In both the Leyden and and less inflated, ... only one false molar." 2 1888, Cat. Systematique des Mammif&res, Mus. During my visit to Leyden Museum in d'Hist. Nat. des Pays-Bas, XII, p. 179. 3 But see Thomas's remark, 1898, Ann. Mag. Nat. 1 1936, J. Mamm., XVII, pp. 257-262. Hist., (7) I, p. 361. 1942] Tate, Results of the Archbold Expeditions. 47 243

London photographs a distinct canine The photograph of the type of prin- cusp can be seen, and p4 is almost in con- cipulus shows the braincase a little higher tact with canine. The infraorbital foramen and broader than in mimus. Zygomata is very close to the edge of the orbit (0.5 weak. Basial pits at least visible. mm.). In view of the close tally of the char- Pipistrellus mimus glaucillus Wroughton acters of the British Museum animals with Pipistrellus mimus glaucillus WROUGHTON, the Leyden specimens and their history, I 1912, J. Bombay Nat. Hist. Soc., XXI, p. 769. am inclined to believe that they may be TYPE LOCALITY.-Multan, Indus River, Pun- true cotypes of tenuis Temminck. jab, northwest India. Finally there is a specimen in the Arch- Apparently a "mouse-gray" race of bold Collection from Bali, not in good con- mimus, whose color Wroughton states is dition (the canines have been lost), which near "bistre." Only skin measurements I am persuaded is also a specimen of tenuis. were given. Forearm, 29 mm. The dimensions of the type and two A photograph of the type skull shows: topotypes in the table of measurements a small-sized, short, muzzled skull with give a good idea of the proportions of the steeply sloping frontal region, braincase skulls in P. tenuis. very slightly higher than in mimus. For skull measurements (made from photo- Pipistrellus nitidus (Tomes) graph) see table. Scotophilus nitidus TOMES, 1858, Proc. Zool. Soc. London, p. 538. Pipistrellus mimus mimus Wroughton TYPE LoCALITY.-Northwest Borneo. P-ipistrellus mimus mimus WROUGHTON, 1899, Tomes wrote, "head somewhat ele- J. Bombay Nat. Hist. Soc., XII, p. 722. vated . . . p2 in line with other teeth, TYPE LOCALITY.-Konkan (coast), near Bom- visible from the outside . .. ,i2 in advance bay, western India. of i." Wroughton compared mimus with his My photograph of the type skull shows "abramus" (not from its type region, Japan, the frontal region steep, the braincase quite but from near Bombay). "Inner and outer high and the rostrum relatively low, p2 incisor [of mimus] close together . . . linear; scarcely reduced, palate fairly short and forearm 27-28." broad. He gave the following skull measure- It is necessary to compare this species ments: greatest length, 11.5; greatest with Glischropus tylopus from the same breadth, 6.5-7.0. general region. They are much alike, apart Our photograph of the skull of the type from the character of the thumb. Of the indicates a palate rather broad and short thumb of nitidus Tomes wrote, "Free por- and braincase moderately full; zygomata tion of thumb longer than that enclosed in apparently weak; P2 three-quarters of the membrane." Surely he would have height of P4. Measurements made from noted the pads, had they been enlarged. photograph of skull. Pipistrellus principulus Thomas Pipistrellus papuanus (Peters and Doria) Pipistrellus principulus THOMAS, 1915, Ann. Vesperugo papuanus PETERS AND DORIA, 1881, Mag. Nat. Hist., (8) XV, p. 231. Ann. Mus. Civ. Genova, XVI, p. 696. TYPE LOCALITY.-Gauhati, Assani. TYPF LOCALITY.-Salawatti, west NewGuinea. In this bat the inflation of the brain- This little species, found all over New case, common to this group as a whole, is Guinea, is apparently annectant between stated to be accentuated. The comparison the tenuis and coromandra groups. The with "tenuis" is doubtful. Under descrip- face is slightly less shortened than in tion of P. ridleyi, Thomas wrote, "no au- tenuis, considerably less than in mimus. thentic specimens of tenuis in Museum col- A number of specimens of the species are lection." Probably, however, he had the present in the Archbold Collections from two "topotypes" with which to make com- Papua and Netherlands New Guinea. It parisons (see under tenuis). may be expected in northeast Australia. 244 Bulletin American Museum of Natural History [Vol. LXXX

Pipistrellus papuanus orientalis (Meyer) Pipistrellus pulveratus (Peters) Vesperugo papuanus orientalis MEYER, 1899, Vesperugo pulveratus PETERS, in Swinhoe, Abh. Ber. K. Zool. Anthr. Ethn. Mus., Dresden, 1870, ProC. Zool. Soc. London, pp. 618-619. VII, 7, p. 14. TYPE LOCALITY.-Amoy, Fukien, China. TYPE LOCALITY.-Bongu, Astrolabe Bay, The skull of a near topotype (from north New Guinea. Chungan Hsien, Fukien), A.M.N.H. 84844, I have been unable to distinguish valid male, shows characters which recall those differences between descriptions of P. p. of P. affinis. The skull is low, but the orientalis and true papuanus. braincase is rather fuller than in affinis; the Pipistrellus affinis Group supraorbital ridges are similarly developed; Typical P. affinis and P. pulveratus are the zygoma and its postorbital process are possibly the most primitive members of the both stoutly developed; p2, though dis- genus Pipistrellus. The second upper pre- placed inward, has approximately the same molar is still as large as i2. The zygomata area as i2; the palate is rather narrow, and are deep and strong and possess postorbital the toothrows converge slightly in front; no processes. The skulls are long and rather basial pits. low, and the molar rows are almost paral- P. pulveratus is, however, considerably lel. No basial pits are developed. smaller than affinis (forearm, 33-34.5 mm.; The species affinis, pulveratus and petersi affinis, 38) and lacks the area of dull white appear more nearly related to each other hairs on upper thighs and pubic area be- than to others. I have no facts on the size neath. The pelage is blackish, grizzled with of p2 in lophurus and kitcheneri. Both pale gray above, with buffy beneath. species have strongly developed basial Thestronglydeveloped zygomais reminis- pits just as in the case of P. circumdatus. cent of the savii group. But p2 in the pres- They are included ent species shows no sign of reduction, provisionally. whereas in savii it is markedly reduced in Pipistrellus affinis (Dobson) ize. Vesperugo affinis DOCBSON, 1871, Proc. Asiatic We have a second topotype and a speci- Soc. Bengal, p. 213. men from Yunnan Province. TYPE LOCALITY.- Bhamo, Yunnan. The present interpretation of affinis is Pipistrellus petersi (Meyer) founded upon a male specimen of P. affinis Vesperugo petersi MEYER, 1899, Abh. Ber. K. from Li Chiang, Yunnan, China, 9000 feet, Zool. Anthr. Ethn. Mus., Dresden, VII, 7, pp. A.M.N.H. 44565, formerly marked "P. 13-14. pulveratus." The identification with Dob- TYPE LOCALITY.-Minahassa, north Celebes. son's affinis is made chiefly by skin char- The type specimen of this species was acters, which are very distinctive, the hairs not seen. Two individuals lent by the U. S. fuscous, with hoary tips, and (most strik- National Museum, U.S.N.M. 219409, ing) the hairs of the pubic region dull white 219413, agree very closely with the little to the bases. Forearm, 38 mm. which has been written about the dentition, The skull of our specimen is remarkable for apparently the skull had not been for its low braincase, unbroadened rostrum, cleaned when Meyer wrote. fairly substantial zygomata bearing low The skulls of both of the Washington but distinct postorbital processes, narrow specimens are clean and in good condition. palate. In the dentition, p2 is not smaller They can be referred without hesitation than i2 but is placed internally, and p4 to the affinis group. Elongate, low, with nearly makes contact with canine. The low braincase, no basial pits, unbroadened lower incisors are moderately imbricated; rostrum, subparallel molar series, narrow canine is not large; P2 stands uncrowded in palate, zygoma deep and strong and pro- the toothrow, its crown height about one- vided with postorbital process. half of that of p4. Teeth: upper incisors heavy, not in con- The type of affinis is probably at the tact with canine; canine with heavy pos- Indian Museum, Calcutta. terior blade but without posterior cusp; 19421 Tate, Results of the Archbold Expeditions. 47 245 p2 as large in area as p4, wholly out of line, Pipistrellus ceylonicus Group and canine in contact with p4; inner face Bats of moderate size (forearm, 36-41 of molars (protocones plus hypocones) mm.), their skulls with low braincases, no longer than in true affinis (1.0 mm.: 0.8 frontal groove. The zygomata are weak mm.); lower incisors well imbricated; P2 and unprovided with postorbital processes. slightly displaced, about three-quarters of Compared with species of the affinis height of P4. group, the facial part of the skull (rostrum Meyer described the dorsal color as Van- and palate) is shorter and broader; the dyke Brown. The Washington specimens palatal length and toothrows are shorter, are in alcohol. The animals are from Cele- and the lacrimal width greater. Basial bes but without precise locality. pits weak. Dentition: the canine bears a strong supplemental cusp placed a little below the Pipistrellus lophurus Thomas cingulum; p2 equal in area to i2. Pipistrellus lophurus THOMAS, 1915, J. Bom- Forms referable to the group are ceyloni- bay Nat. Hist. Soc., XXIII, p. 413. cus, indicus, subcanus and chrysothrix, and, TYPE LOCALITY.-Maliwun, Victoria Prov- tentatively referred, shanorum and raptor. ince, Tenasserim. The first four are from the Indian peninsula; The photograph of the type shows a the others, from the Burma-Indo-China braincase fairly high at the occiput; slight region. frontal depression in profile; a deep zygoma with well developed postorbital proc- Pipistrellus ceylonicus (Kelaart) ess; the postpalatal spine obsolescent. Scotophilus ceylonicus KELAART, 1852, Prodr. Dentition: p2, twice as large as in imbrica- Fauna Zeylanica, p. 22. tus, is as great in crown area as i2. Fore- TYPE LOCALITY.-Ceylon. arm, 35 mm. I have before me specimens from Ceylon Thomas emphasized a dorsal tuft of hairs (M.C.Z. 27525-26). The color is dark at the base of the tail which he supposed brown, slightly paler beneath. No trace of marked the position of a gland. pale tips of hairs as in affinis group. Fore- The dorsal color was described as "warm arm, 37-42 mm. (Phillips). The thumb ap- bister-brown, below paler than Prout's pears large and strong; its basal phalanx, Brown." 3.5 mm. The skull is as described under the "group" heading. Dobson (1878, pp. 222-223) pointed out Pipistrellus kitcheneri Thomas that the type had been lost and that cey- Pipistrellus kitcheneri THOMAS, 1915, Ann. lonicus might, because of discrepancies in Mag. Nat. Hist., (8) XV, p. 229. Kelaart's description, remain unrecogniz- TYPE LOCALITY.-Boentok, Barito River, able. However, we may perhaps accept south central Borneo. the decisions of Wroughton, of Thomas and My photograph of the type skull shows of Phillips that the name belongs to the a deep zygoma with a small postorbital present kind of bat. ossification; the general appearance of skull very like that of lophurus. If the Pipistrellus ceylonicus indicus (Dobson) supposed caudal gland of lophurus were Vesperugo indicus DOBSON, 1878, Cat. Chiropt. shown to be individual, or in reality a Brit. Mus., p. 222. wound or ulcer peculiar to the single speci- TYPE LOCALITY.-Mangalore, Malabar Coast, men, we might well conclude that lophurus Madras. and kitcheneri were subspecies. The type of this species is probably at Forearm, 37 mm. Color, brown. the Indian Museum, Calcutta. Thomas compared kitcheneri with im- Dobson gave no data on the skull, and bricatus. Length of palate indicated by of the teeth he wrote, ". . . first upper pre- "muzzle less conspicuously shortened as molar quite internal to toothrow, not visible compared with braincase." from without; second upper premolar al- 246 Bulletin American Museum of Natural History [Vol. LXXX most touching canine; lower incisors in the Pipistrellus raptor Thomas direction of the jaws" (indicating a V- Pipistrellus raptor THOMAS, 1904, Ann. Mag. shaped symphysis). Nat. Hist., (7) XIII, p. 387. He mentioned possible relationship to TYPE LOCALITY.-Tonkin, French Indo-China. ceylonicus. Zygoma weak, without process. Skull of type essentially as in shanorum but p2 un- Pipistrellus ceylonicus subcanus Thomas reduced. Muzzle slightly broadened and Pipistrellus ceylonicus subcanus THOMAS, 1915, smooth as in ceylonicus. J. Bombay Nat. Hist. Soc., XXIV, p. 30. Measurements show the lacrimal width, TYPE LOCALITY.-Yalala, Junagadh State, zygomatic width and skull length somewhat Kathiawar, western India. greater than in other species of this group. "Grayer than ceylonicus." Forearm, 38 Our photograph of the type skull sug- mm. gests relationship with shanorum and The photograph of the skull of the type ceylonicus rather than with circumdatus or is very similar to that of our ceylonicus. mordax. The toothrow appears a little longer and the braincase slightly wider. Probably Pipistrellus kuhlii Groip .subcanus, indicus, ceylonicus and chrysothrix are races of a single species. Thomas Chiefly bats of Europe, western Asia and suggested that chrysothrix "was founded on Africa, reaching central India. Skull low a specimen of P. ceylonicus with abnormal and broad, the zygoma weak, p- small but incisors." less reduced than in savii group, and the lower incisors thickened and strongly im- Pipistrellus ceylonicus chrysothrix bricated. Wroughton Named forms besides kuhlii are: canus, Pipistrellus chrysothrix WROUGHTON, 1899, ikhwanius, lepidus, leucotis and lobatus. J. Bombay Nat. Hist. Soc., XII, p. 720. Several have been treated as synonyms of TYPE LOCALITY.-Konkan, near Bombay, western India. kuhlii; others are races. I have been able Thomas, when writing of subcanus, re- to examine only European kuhlii, and the ferred chrysothrix as a subspecies of cey- type of babu, which is referred here pro- lonicus. visionally. Our photograph of the type skull indi- The group badly needs revisional treat- cates the reasonableness of this association: ment, but many of the types must be in the rather broad rostrum, weak zygoma, India. The obsolescence of the secondary etc. cusp of il in babu is also to be noted in The dorsal color was described by kuhlii. Wroughton as "golden brown"; il with secondary cusp "posterior." Pipistrellus kuhlii (Kuhl) Vespertilio kuhlii KUHL, 1819, Ann. Wetterau. Pipistrellus shanorum Thomas Gesellsch. Naturk., IV, p. 199. Pipistrellus shanorum THOMAS, 1915, J. Bom- Romicia calcarata GRAY, 1838, Mag. Zool. Bot., bay Nat. Hist. Soc., XXIV, p. 29. II, p. 495. TYPE LOCALITY.-Pyaung-gaung, North Shan TYPE LOCALITY.-Trieste, Adriatic Sea. States, Burma. Miller separated kuhlii from savii by the The character of the zygoma, broken, less reduced p2 and by the tragus being cannot be seen in the photograph of the longer and narrower, and from pipistrellus type. Otherwise the skull agrees well with and nathusii by the shorter outer upper those of the ceylonicus group. Thomas incisor and closure of the diastema between wrote, "closely allied to ceylonicus but c and p4. smaller"; p2 smaller than in ceylonicus, its The skull of one before me (U.S.N.M. area two-thirds of that of i2. Forearm, 37 172128) shows only a slight frontal depres- mm. sion in profile; a slight tendency to an Color "dark, warm brown, tipped with occipital "helmet"; zygomata weak, basial dark buffy or cinnamon." pits almost undeveloped; no trace of pos- 1942] Tate, Results of the Archbold Expeditions. 47 247 terior cusp on canine; posterior cusp of il visible, its area about equaling that of i2; obsolete; area of p2 about one-half that of c-mi3, 5.0 mm. p4. Lower incisors very markedly imbri- I have seen only the type and photo- cated; height and cingulum length of P2 graphed its skull. about two-thirds of same measurements of Pipistrellus leucotis (Dobson) P4. The wings of kuhlii are frequently mar- with whitish. Vesperugo leucotis DOBSON, 1872, J. Asiatic gined Soc. Bengal, XLI, p. 222. TYPE LOCALITY.-Kachh, near Quetta, north- Pipistrellus kuhiii lepidus Blyth west India. Pipistrellus lepidus BLYTH, 1845, J. Asiatic Wing membrane near body white; rest Soc. Bengal, XIV, p. 340. TYPE LOCALITY.-Kandahar, Hyderabad, cen- of wing traversed by white reticulations. tral India. Dobson (1878) placed this bat in the Not mentioned in Dobson (1878). synonymy of kuhlii. Wroughton (1917) Wroughton (1917, p. 591) treated it as a followed Dobson's views. He gave the subspecies of kuhlii, but the type was number of the type: 154 p. India Museum, unknown, so it is not probable that he had Calcutta. more than descriptive information to sup- Pipistrellus canus (Blyth) port his decision. Nycticejus canus BLYTH, 1863, Cat. Mamm. "General color a light yellowish-clay, Mus. Asiatic Soc. Bengal, p. 32. pale sandy or isabella brown; underneath TYPE REGION.-India. paler . . . forearm 1 inch and three-eighths Dobson (1878, p. 230) placed canus in the or a trifle less. . ." (34 mm.). synonymy of kuhlii. Wroughton (1917, p. 591) agreed. He mentioned two cotypes, Pipistrellus kuhlii ikhwanius Cheeseman Nos. 154a, b, in the collection of the Indian and Hinton Museum, Calcutta. Pipistrellus kuhlii ikhwanius CHEESEMAN AND This would give to kuhlii a remarkably HINTON, 1924, Ann. Mag. Nat. Hist., (9) XIV, extended range, if true. Probably canus p. 549. TYPE LOCALITY.-Arabia. should be considered a geographical race. "Near kuhlii"; color pale, light buff; Pipistrellus lobatus (Jerdon) forearm, 33-34 mm.; c-m3, 4.6 mm. Scotophilus lobatus JERDON, 1867, Mammals of India, p. 35. Pipistrellus babu Thomas TYPE LOCALITY.-Madras, India. Pipistrellus babu THOMAS, 1915, J. Bombay "Ears small, ovoid, ends rounded, Nat. Hist. Soc., XXIV, p. 30. scarcely emarginate, with a lobe at the base; TYPE LOCALITY.-Murree, near upper River tragus short, of nearly uniform breadth ... Indus, northern India. muzzle very short, pointed. Fur, above, Thomas called babu the north Indian rep- black.sh-yellow, ashy, beneath. . resentative of "tralatitius." As pointed out Jerdon (1874 edition) queried whether elsewhere in this paper, he regarded trala- lobatus might not equal abramus Tem- titius Horsfield as Pipistrellus, not a Myo- minck. Forearm length was published as tis.' The species does not readily fit into 12/12 inches (30 mm.). Jerdon's broad con- one of the groups here arranged. It appears cept of Scotophilus included Eptesicus sero- to be too large for the coromandra group and tinus and Pipistrellus. a little small for the affinis group. Perhaps Dobson, however, considered lobatus a it is best regarded as a member of the synonym of kuhlii. Wroughton ignored the kuhlii group. name. Forearm, 33-35 mm.; a distinct rostral sulcus; il with only a trace of supplemental Pipistrellus minahassae Group cusp (see kuhlii); p2 wholly internal but Pipistrellus minahassae Meyer 1 Horsfield gives the molar formula of tralatitius as Pipistrellus minahassae MEYER, 1899, Abh. 5 5. With three lower premolars and three lower Ber. K. Zool. Anthr. Ethn. Mus., Dresden, VII, molars,-6ctralatitiuss could not have been a Pipistrellus. 7, pp. 14-16. 248 Bulletin American Museum of Natural History [Vol. LXXX

* TYPE LOCALITY.-Tomohon, Minahassa, north fifth of that of p4, but p2 retained either in Celebes. toothrow or but slightly moved inward; A single specimen contained in the Arch- lower incisors not imbricated; i, and i2 with bold Collections, from Roeroekan, north crowns unthickened, i3 with crown distinctly Celebes, is provisionally referred to Meyer's thickened; canine slightly higher than p4; species. Unfortunately no description of P2 crown one-half height of P4 crown. the type skull of minahassae has been pub- Wings pale gray; dorsal pelage pale gray lished, so identity cannot be guaranteed. with slightly darker bases; ventral pelage The skull of the animal from Roeroekan pure white to roots. (A.M.N.H. 102359, e) presents novel Reference of coxi to this group is pro- characters sufficient in themselves to set it visional. off from other Pipistrelli as a distinct group. The distinctive characters are: the short, high braincase with rudiments of sagittal Pipistrellus coxi Thomas Pipistrellus coxi THOMAS, 1919, J. Bombay crest; the prominent supraorbital tubercles Nat. Hist. Soc., XXVI, p. 747. suggestive of the joffrei group but less de- TYPE LOCALITY.-Amara, Mesopotamia. veloped than those of Philetor; the fact Compared with the African ri4ppellii, that p2 iS scarcely, if at all, smaller than P4. under surface whitish. This last is perhaps a primitive character "This bat ... seems to mark the northern harking back to the period before p2 be- limit of a series beginning with the Uganda came markedly reduced in size. P. fuscipes, which has a large and much Other features of the skull include: inflated braincase, etc. . . Egyptian and zygoma slender, as in abramus group; deep Sudanese P. riippellii, in which the brain- basial pits; p2 but little moved inward from case is more normal, while in coxi it is dis- the toothrow, its area slightly exceeding tinctly smaller than usual." that of i2; il long, with well developed posterior accessory cusp; canine slender without extra cusp; lower incisors scarcely Pipistrellus savii Group imbricated. Comprising rather smaller bats than This bat is wholly different from P. those of the affinis and circumdatus groups, petersi, described by Meyer from the same but the well built skull with its strong island (see under that species). Its affini- zygomata suggests relationship. There are ties appear to be with the more primitive traces of postorbital processes on the zygo- pipistrelles. mata. P. savii has, however, become specialized bythe great degree of reduction of p2 Pipistrellus riippellii Group to only about one-fourth of the area of i2. African bats, with one putative repre- The braincase is relatively low as in the sentative in Mesopotamia. These bats more typical bats of the affinis group. were referred by Dobson (1876) to Scoto- The group apparently is considerably zous, but Hollister regarded them as special- developed in Africa: ariel from Egypt, ized Pipistrellus. Read under Scotozous. darwini from Canary Islands, maderensis P. riappellii, of which we have specimens, from Madeira. Present inclusion of several is unlike any of the Asiatic Pipistrellus oriental species with savii may not be ten- groups. Braincase very full, interorbital able. It is based upon one character com- area broad, rostrum short, unwidened and mon to all, the extreme degree of reduction slightly flattened above. Zygoma very of p2. Otherwise the eastern forms curtatus weak and delicate. Palate narrow. No and cadornae (and possibly austenianus) basial pits. differ rather sharply from savii by their Dentition: il with accessory cusp almost full, rounded braincases and short muzzles. as big as primary cusp; i2 slender, small, its The braincase of savii is low and flat and tip reaching only one-half way down supple- the muzzle less shortened. The only other mentary cusp of il; canine slender, long, species known to me which has p2 much re- without cusp; p2 small, its cusp only one- duced is circumdatus, a much larger form 1942] Tate, Results of the Archbold Expeditions. 47 249 with forearm 40 mm., possibly related to pelage agrees quite closely with the descrip- mordax, also with forearm 40 mm. tion of Temminck, "brown-bister or onion- color," a light brown near Sayal Brown or Pipistrellus savii (Bonaparte) Tawny. The ears are large and the tragus Vespertilio savii BONAPARTE, 1837, Iconogr. broad, deeply notched near the base of its Fauna Ital., I, fasc. 20. posterior margin, narrowed toward its tip, Vespertilio maurus BLASIUS, 1853, Archiv. f. Naturg., XIX, Part 1, p. 35. and with the posterior margin strongly TYPE LOCALITIEs.-Pisa, Italy (savii); cen- curved while the anterior margin is slightly tral chain of Alps (maurus). excavated. The thumb is slender, its Miller separated this species from the re- basal phalanx 3.2 mm. A small pollical maining three European pipistrelles by the pad covers the joint between the meta- extremely small size of p2 measured in carpal and the phalanx. terms of the area of i2, and by the widened ear and extremely broad tragus. Pipistrellus vordermanni (Jentink) The skull of a specimen of savii (U.S. Vesperugo vordermanni JENTINK, 1890, Notes N.M. 103322) before me is low, without Leyden Museum, XII, p. 152. frontal depression between braincase and TYPE LoCALITY.-Billiton Island, Dutch East rostrum. Rostrum depressed just in front Indies. of supraorbital ridges. Zygoma strong Jentink referred to the white wing mem- but without postorbital process. Basial branes of vordermanni. Dentition: i2 equal pits almost undeveloped. Area of p2 in height to external cusp of il; p2 very about one-sixth of area of i2. Lower in- small, hardly visible without lens, placed cisors markedly imbricated, lower canines inside the toothrow. Forearm, 33 mm. short, height and cingulum length of P2 The braincase is full and rounded, ros- respectively one-half of height and cingu- trum and palate rather short, somewhat as lum length of P4. in the tenuis group, but the skull is much larger. Well marked basial pits can be seen Pipistrellus macrotis (Temminck) in the photograph of the type. The zygo- Vespertilio macrotis TEMMINCK, 1835, Monogr. mata are deepened, and traces of postor- Mamm., II, p. 218. bital processes are present. TYPE LOCALITY.-Padang, Sumatra. At Leyden I found the following cotypes: Pipistrellus cadornae Thomas mounted skins "n" and "o," both with Pipistrellus cadornae THOMAS, 1916, J. Bom- skulls inside, both from Padang, also skull bay Nat. Hist. Soc., XXIV, p. 415. "m" from Padang. TYPE LOCALITY.-Darjiling, Assam. The skull of "m" had the frontal region Forearm, 33 mm.; p2 very small; basial slightly raised; bullae rather large; p2 pits developed. Thomas compared ca- minute, included on inside between c and dornae with kitcheneri and ceylonicus. But P4; P2 about one-half of height of P4. it is even more closely related to curtatus My photograph of skull "m" shows a Miller from Engano Island and to a speci- skull very similar to that of P. vordermanni. men (IJ.S.N.M. 141097) labeled "macrotis." The forearm length of specimen "n" was These bats are distinguished by their full 34.5 mm. Temminck published the length braincase, minute p2, deepened zygomata for macrotis as 1 inch 2 lines (29.5 mm.). bearing distinct postorbital processes and A series of three specimens in the Arch- short rostra and palates. Their association bold Collection from Koeta, Bali, belongs with savii comes from the similar reduction without doubt to the assemblage which in- of p2 in that species. cluded curtatus, cadornae and macrotis. It has the characteristic deep zygoma Pipistrellus curtatus with well developed postorbital process of Miller the zygoma, full rounded short Pipistrellus curtbtus MILLER, 1911, Proc. Biol. braincase, Soc. Washington, XXIV, p. 25. rostrum and palate, deep basial pits and TYPE LOCALITY. Engano Island, west of extremely reduced p2. The color of the Sumatra. 250 Bulletin American Museum of Natural History [Vol. LXXX

Forearm, 33.6 mm. Crown area of lower trelle is characterized by the distinctly premolar much reduced. Rostrum and bronzy tips to the hairs, the basal and sub- palate short. Basial pits well defined. terminal portions being fuscous. The color scheme of the hairs is somewhat like that of Pipistrellus austenianus (Dobson) Myotis sicarius. Vesperugo austenianus DOBSON, 1871, Proc. Asiatic Soc. Bengal, p. 213. TYPE LOCALITY.-Cherrapunji, Assam. Pipistrellus mordax (Peters) In his 1878 syn- Vesperugo mordax PETERS, 1866, Monatsb. "Catalogue" Dobson Akad. Wiss. Berlin, p. 402. onymized austenianus with savii of Europe. TYPE LOCALITY.-Java. Wroughton' treated it as distinct in his "Color above red-brown ... with paler key: forearm, 33 mm. or less; tragus broad, tips . . . forearm 40 mm." No mention of above 4 mm.; color black, hoary. The cranial or dental characters. type is No. 150b of the Indian Museum at Peters compared this species with the Calcutta. European maurus (= savii), but in 1866 Dobson states: forearm, 1.4 inches (35 he was just beginning his remarkable work mm.); p2 "may be distinguished from with- on the Chiroptera and probably was unac- out." quainted with many species. Later Dob- P. austenianus is here placed in the savii son, Trouessart and others included mordax group solely because Dobson once synony- as a race of savii, which clearly it is not. mized it with savii. The skull of the type The type of mordax should be compared should be critically reexamined and de- with P. circumdatus which has a forearm scribed. of similar length. Sody2 gives measurements of five speci- Pipistrellus circumdatus Group mens, with forearms 40-42 mm., which he The only species definitely known to be- believes referable to mordax. He gives no long here, circumdatus, shows several special cranial or dental dimensions. characters: short, broad outline of skull; short permaxillae, as in tenuis group; strong Pipistrellus tasmaniensis Group zygomata without any trace of postorbital processes; distinct basial pits somewhat Represented by a single large species, as in Ia; supraorbital region short, un- tasmaniensis, of which krefftii is commonly broadened; a deep frontal depression; P held to be a synonym. Distinguished from heavy, virtually unicuspid; i2 placed well most pipistrelles by the unicuspid iP and forward; canine width markedly less than by the "helmeted" form of the back of the lacrimal width; p2 much reduced, area of skull. In it p2 are much reduced. The p2 about one-fourth of that of i2. zygoma is weak and slender as in many Inclusion of mordax with circumdatus other pipistrelles. Tasmaniensis is slightly may not be permissible. The forearm reminiscent of Nyctalus, both having p2 lengths are approximately the same. much reduced, but it differs in the relative narrowness of its rostrum. The group is Pipistrellus circumdatus (Temminck) distinctly aberrant. Vespertilio circumdatus TEMMINCK, 1835, Iredale and Troughton3 placed tasmani- Monogr. Mamm., II, p. 214. ensis in Glischropus. It is quite unrelated TYPE LoCALITY.-Tapos, Java. to that genus, lacking all trace of the en- Forearm, according to Temminck, 40 larged pad on the basal joint of the thumb. mm. (1 inch 7 lines); 38 (1.5 inches) ac- The Archbold Collection contains a large cording to Dobson. In a specimen from series of G. tylopus, the type species of north Burma (A.M.N.H. 114850) the fore- Glischropus, in which the character above arm measured 41 mm. mentioned is clearly discernible. Tasmani- The pelage of this relatively large pipis- ensis is further separable from Glischro- 1 1917, J. Bombay Nat. Hist. Soc., XXV, pp. 589, 2 1937, Temminckia, II, p. 215. 591. 3 1934, Mem. Australian Mus., VI, p. 96. 19421 Tate, Results of the Archbold Expeditions. 47 251 pus by its large size (forearm, 48 mm.), the other known species by its greater size form of il and its greatly reduced p2. (forearm, 45 mm.). It may, in fact, belong to a different genus-to Ia or Nyctalus, Pipistrellus tasmaniensis (Gould) both of which agree in dental formula with Vespertilio tasmaniensis GOULD, 18581 (1863), Pipistrellus. The type in the Indian Mu- Mammals Australia, III, P1. XLVIII (with text). seum is still the only specimen known, and TYPE, REGION.-Tasmania. knowledge of the species still depends upon The photograph of Gould's type speci- Dobson's description of it. men, B.M. 43.2.22.6, shows only the front Bianchi's name Megapipistrellus may be of the skull preserved (muzzle and tooth- invalidated by Alobus Peters,2 which was rows). The unicuspid il; i2 in virtual con- proposed to contain the small African tact with canine; no space between c and species temminckii Cretzschmar, which p4; p2 not visible (actually extremely lacks the calcareal lobe, has the ear pinna minute); palate narrow; lacrimal width much like Myotis dasycneme, and the ros- small. trum relatively low and short. Dobson The bats of the Auistralian mainland (1876) employed Alobus for annectens and (krefftii) mentioned next are in all prob- pulcher Dobson, as well as for temminckii. ability not separable from those of Tas- Thus, if temminckii and annectens are con- mania. It is customary to place them in subgeneric, Megapipistrellus becomes a the synonymy of tasmaniensis. synonym of Alobus. Pipistrellus tasmaniensis krefftii (Peters) Vesperugo krefftii PETERS, 1869, Monatsber. Pipistrellus annectens (Dobson) Akad. Wiss. Berlin, p. 404. Vesperugo annectens DOBSON, 1871, Proc. TYPE LOCALITY.-New South Wales. Asiatic Soc. Bengal, p. 213; 1876, Monogr. Asiatic Chiroptera, p. 116; 1878, Cat. Chiropt. Peters described "il large, single-cusped; Brit. Mus., p. 234. . p2 very small...." The forearm was TYPE LOCALITY.-Naga Hills, Assam. given as 48 mm. and the tibia, 19. "Upper incisors nearly equal . . . canine In two specimens from Ebor, New South small, scarcely exceeding the second pre- Wales, il is long and without trace of ac- molar in the upper jaw [p4] in vertical ex- cessory cusp; i2 is very small, its crown tent; in the lower jaw the second premolar only one-fourth of height of crown of il; [P4] slightly exceeds the canine; first canine long; without supplemental cusp; upper molar minute, placed slightly inside p2 excessively small, about one-third of the toothrow, but distinctly visible from crown area of already reduced i2; c and p4 without." contiguous; lower incisors very strongly Forearm, 45 mm. (1.8 inches). imbricated; canine strong, curved, con- No other specimen is known. siderably higher than P4; P2 small, com- In his 1876 article Dobson wrote ". pletely in toothrow, but its cusp less than this species unites the external appearance one-third of height of cusp of P4. of a Vespertilio [Myotis] to the dentition The skull is low and rather flat; the of Vesperugo [Pipistrellus]. In the form of zygoma, though weak, shows a trace of the ears, and tragus, and the elevation of postorbital process; posterior part of sagit- the roof of the skull above the face, it very tal crest developed to form slight "hel- closely resembles some species of the former met"; palate long, little domed; no basial genus." Two woodcuts of the head were pits. shown with the same text. Pipistrellus annectens Group (= Alobus Peters? = Megapipistrellus Bianchi) Pipistrellus joffrei Group A single species, annectens Dobson, for A special Burmese-Malaya group which which Bianchi has proposed the subgenus in some ways connects Pipistrellus to Megapipistrellus, differs sharply from all Nyctalus but is distinguished by peculiari- 1 Fide Iredale and Troughton, 1934, Mem. Aus- 2 1867, Monatsber. Akad. Wiss. Berlin, p. 707. But tralian Museum, VI, p. 96. see also this paper under Glischropus. 252 Bulletin American Museum of Natural History [Vol. LXXX ties. It is characterized by its strongly Nyctalus. The remarkable accessory canine specialized upper canine (see beyond), cusp is clearly visible. Rostrum much minute p2 and small i2; subequal P2 and lower than in Nyctalus. Basial pits present P4; short, broad rostrum, in which the but weak. width across prominent supraorbital tuber- The skin was described as "uniform pale cles exceeds the lacrimal width. A small brown above and below." descending process from the zygoma external to m3. Pipistrellus anthonyi, new species Included are Nyctalus joffrei Thomas, Pipistrellus affinis (?) ANTHONY, 1941, Publ. type of the group, P. anthonyi, new species, Field Mus. Nat. Hist., Zool. Ser., XXVII, p. 81. TYPE.-No. 114849, Amer. Mus. Nat. Hist., described beyond, P. stenopterus and P. ad. a', Changyinku, northern Burma; 7000 feet; brachypterus. collector, H. E. Anthony, Vernay-Cutting Burma The species joffrei, according to Thomas, Expedition; March 13, 1939. The type is a has a "build suggesting a large pipistrelle skin with skull, the latter in bad condition. GENERAL CHARACTERS.-A dark brown pipis- rather than a noctule but the proportions trelline bat structurally close to joffrei but of the digits quite as in Nyctalus .... colored very dark brown instead of "pale Skull of a somewhat different shape from brown." that of other species of the genus (Nyc- DESCRIPTION OF TYPE-.Pelage firm, glossy to the broad- and velvety. Dorsal color near Bone Brown; talus), particularly in regard hairs of underparts tipped with Wood Brown, ened rostrum and expanded supraorbital their bases fuscous. Thumb short, its basal tubercles." phalanx 3.4 mm. (heavier than in affinis) and I do not feel that it is proper to include provided with a small basal pad (smaller than in joffrei with Nyctalus as heretofore. The Glischropus); claw short and blunt. Feet heavy; a moderately developed calcareal lobe. special nature of the canine, the shortened Skull (lacking cranial portion): rather wide cingulum length of P4, the pronounced and only moderately high, with exceptionally tubercles above the eyes are repeated al- wide lacrimal region and even wider space across most exactly in Philetor. The P. joffrei well developed supraorbital tubercles, but narrower muzzle and canine width. Zygoma flaring group approaches oriental members of the from maxilla, provided with a small but distinct P. savii group. In the latter, p2 iS minute descending process, 0.5 mm. in depth, external and the palate short, but supraorbital to m3. Sagittal suture very faintly crested and tubercles are undeveloped. For the present continued as faintly defined paired crests curving outward to meet the supraorbital tubercles. the joffrei group is treated as an offshoot Face short, the narial sinus level with a line of Pipistrellus. It may later be accorded drawn through the fronts of the orbits. Palate subgeneric rank. short, domed. P. stenopterus and P. brachypterus are Dentition: upper incisors missing (alveolus of il larger than that of i2); canine with very seemingly slightly less advanced members strongly defined posterior cusp which descends of the joffrei group. Stenopterus possesses from the posteriorly extended cingulum and is the specialized canines and broad rostrum partly fused with the main cusp; p2 quite as but has less developed supraorbital tuber- small as in P. savii or P. macrotis, its crown diameter about one-fourth of crown diameter of i2, cles; in brachypterus the canine cusp is low that of its alveolus about one-third of diameter of and weak and the rostrum less broadened. alveolus of i2; p4 a large tooth, the cusp descending to midway between the main and pos- Pipistrellus joffrei (Thomas) terior cusps of canine, but the body of the tooth is much shortened (in the toothrow), and Nyctalus joffrei THOMAS, 1915, Ann. Mag. a diastema exists between the front of the pro- Nat. Hist., (8) XV, p. 225. tocone area of ml and what represents the area TYPE LOCALITY.-Kachin Hills, Upper Burma. of the hypocone of p4 (actually the hypocone is Forearm, 39 mm. Our photograph of the obsolete); lower incisors also missing; canine type skull clearly shows the great expanse about as tall as p4; P2 and P4 both rather small, P4 only slightly taller than P2 but almost as much across the supraorbital tubercles in com- compressed longitudinally (in toothrow) as it is bination with relatively narrow canine in the case of Philetor. width (a character not shown in Nyctalus). MEASUREMENTS OF THE TYPE.-Skin: fore- The anterior palatal sinus is narrow, and arm, 38 mm.; tibia, 12; total length, 961; tail i1 are more approximated than in true 1 Made in field. 1942] Tate, Results of the Archbold Expeditions. 47 253

vertebrae, 41'; hind foot (c.u.), 101. Skull: The photograph of the type skull indi- lacrimal width, 7.0; width across supraorbital cates a bat near joffrei, with the rostrum tubercles, 7.8; intertemporal width, 4.9; width across canines, 5.1; width across m3-3, 7.2; short and broad in comparison with Nyc- palate length, 4.8; c-m3, 5.3; cingulum length talus. The canines are less approximated of lower canine, 1.0; p2, 0.55; P4, 0.45. than in joffrei and anthonyi, the first upper REMARKS.-At first sight the characters incisors project farther forward, and the of this interesting bat suggested the genus supraorbital tubercles are less prominent. Philetor more than Pipistrellus, particularly in the development of supraorbital GLISCHROPUS DOBSON tubercles and the modification of P4. But Glischropus (subgenus of Vesperugo) DOBSON, in Philetor p2 is no longer present. 1875, Proc. Zool. Soc. London, p. 472. The species is named for Dr. H. E. GENOTYPE.-AS restricted by Miller,2 Ves- Anthony, Curator of the Department of perugo tylopus Dobson. Mammals, The American Museum of Na- Glischropus contained originally Ves- tural History, who collected it. perugo nanus Peters from Africa, and Ves- perugo tylopus Dobson frN North Borneo. V. nanus Peters is placed b G. M. Allen' Pipistrellus brachypterus (Temminck) under Pipistrellus. Another species for- Vespertilio brachypterus TEMMINCK, 1835, merly and erroneously referred to Glischro- Monogr. Mamm., II, p. 215. TYPE LOCALITY.-Padang, Sumatra. pus is Pipistrellus tasmaniensis (Gray, 1843). Temminck's published forearm length of Batjanus Matschie, 1901, and javanus are as- brachypterus was 1 inch 3 lines (32 mm.) Chasen, 1939, probably correctly but the cotype, Leyden "a," measured by signed. I have pointed out earlier in this paper me gave 35 mm. "Pelage short, smooth ... brown, approaching blackish, beneath the close resemblance of the skull of the umber." type species, G. tylopus, to those of the Pipistrellus tenuis and Pipistrellus coro- The type skin seen by me in Leyden was mandra groups. P. nanus, which formed a mounted skin, specimen "a" of Jentink's catalogue. The skull (only the short, the basis of Alobus Peters,4 is, I think, un- broad, high rostrum and the palate remain) questionably a Pipistrellus and lacks any had lacrimal width, 7 mm.; canine width, trace of the modifications of the thumb- 5.5; c-m3, 5.5. and foot-pads visible in G. tylopus. Glis- It is open to question whether the meas- chropus must then be restricted to the three urements given for brachypterus by Dobson named forms, tylopus Dobson, batjanus can be relied upon. They were taken not Matschie and javanus Chasen. from the type at Leyden but from a speci- The genus may be thought of as a direct men in Berlin. offshoot of Pipistrellus in which the appara- The only data on the type skin appearing tus for grasping has undergone modifica- in my notes are, "ear rather broad and tion. The enlarged pads covering the meta- rounded, with small posterior basal lobe; tarsal-phalangeal joints of the thumb are clearly to be seen, as well as thickening tragus a rather broad, short lobe ... tail, 30 mm.; forearm, 35; ear from crown, 7. and broadening of the entire plantar sur- A specimen from the Philippines, TU.S. faces of the feet. Presumably they improve N.M. 125313, though labeled stenopterus, prehension. appears to agree more with brachypterus. The African species nanus, which Dobson included in Glischropus with tylopus, has Pipistrellus stenopterus (Dobson) quite differently proportioned incisors, i2 Vesperugo stenopterus DOBSON, 1875, Proc. being elongate and its position as in normal Zool. Soc. London, p. 470. Pipistrellus. Also, enlargement of the pads TYPE LOCALITY.-Sarawak Island, Borneo. 2 1907, Families and Genera of Bats, p. 205. Forearm, 39 mm. 3 1939, Bull. Mus. Comp. Zool., LXXXIII, p. 95. 4 Alobus Peters, 1867, Monatsber. Akad. Wiss. Ber- Fur "veryshort, darkbrownthroughout." lin, p. 707, is homonym of Alobus Le Conte, 1856, J. Acad, Nat. Sci. Philadelphia, (2) III, p. 273 (Coleop- 1 Made in field. tera). 254 Bulletin American Museum of Natural History [Vol. LXXX seems to be rudimentary in comparison to it probable that batjanus will have to be those of tylopus. The zygoma of nanus, placed in synonymy. too, is a far more heavily built structure than that of tylopus and bears traces of a Glischropus javanus Chasen postorbital process. Glischropus javanus CHASEN, 1929, Treubia, XVII, p. 189. TYPE LoCALITY.-Pangeango, west Java. Glischropus tylopus (Dobson) Chasen, comparing javanus with tylopus, Vesperugo tylopus DOBSON, 1875, Proc. Zool. Soc. London, p. 473; 1876, Monogr. Asiatic. found the forearm and lower leg longer, Chiroptera, p. 114; 1878, Cat. Chiropt. Brit. the braincase flatter, p2 more displaced and Mus., p. 236. P2 smaller. Certain of the measurements TYPE LOCALITY.-North Borneo. given by Chasen for javanus, of which he The skull of the type, photographed by had apparently only the type, when com- me in 1938, fails to show the Ia-like ar- pared with the series of six of our Bornean rangement of the upper incisors, whereby specimens appear markedly larger. This i2 is thrust so far forward that a straight greater size, coupled with what must be line will pass through the bodies of the four differences of proportion in the skull, sug- teeth, i2, i, ii,ij2. In addition, the cusp of gests that javanus may well be a good i2 is very short, while the supplementary species. cusp of iP is placed posteriorly and descends almost as far as the main cusp. Canine NYCTALUS BOWDICH Nyctalus BOWDICH, 1825, Excursions in Ma- short, its cingulum extended posteriorly but deira and Porto Santo, p. 36. without distinct posterior cuspule. p2 Pterygistes KAUP, 1829, Skizzirte Entwickl.- only slightly reduced, its crown area Gesch. u. naturl. Syst. d. europ. Thierw., I, slightly less than that of j2, but its cusp P. 99. Noctulinia GRAY, 1842, Ann. Mag. Nat. Hist., only one-fifth of height of cusp of p4. X, p. 258. Lower incisors moderately imbricated; P2 Panugo KOLENATI, 1856, Allg. deutsch. naturh. partly displaced, its cingulum length equal Zeitung, (n.f.) II, p. 131. to, its cusp height one-half of, that of P4. GENOTYPE.-(NydtalUs) verrucosus; (Pterygis- tes) noctula; (Noctulinia) proterus and fulvus, Skull and rostrum somewhat lower. representatives; (Panugo) representatives, noc- Zygoma extremely weak, broken in all but tula, leisleri. two or three specimens of our large series; The genus Nyctalus has had a peculiar no basial pits. history. Bowdich wrote of N. verrucosus The pads of thumb and feet are promi- from Madeira, the type, "It forms a new nently swollen, as described by Dobson. subgenus between pharopus (Pteropus, fide The color of the pelage is dark brown above, Palmer, 1904) and cephalotes." Therefore light orange-brown beneath, with dark gray one would assume it to be one of the Mega- bases. chiroptera rather than a noctule. The Of the large series in the Archbold Collec- name Nyctalus was ignored by Dobson, tion, probably all from Perboewa, north- 1878, and by Miller, 1907. But next year west Borneo, may be considered topotypes. Andersen' analyzed Bowdich's description Those from south Borneo may grade over of verrucosus in considerable detail and into javanus. reached the conclusion that it was equal to Pterygistes madeirae Barrett-Hamilton, Glischropus batjanus Matschie 1906, and had nothing to do with the Glischropus batjanus MATSCHIE, 1901, Abh. Senck. Gesellsch., XXV, p. 277. Pteropodidae. The type may not be in TYPE LOCALITY.-Batchian Island, Moluccas. existence. Matschie had five specimens in alcohol. Pterygistes Kaup, 1829, was based upon He distinguished them from tylopus by Vespertilio noctula Schreber. This name their average slightlysmaller ear. However, was employed by Miller (1912) for bats of the forearm length given by Matschie fits the genus Nyctalus. By Dobson they were perfectly into the pattern shown by meas- considered merely a section of "Vesperugo." urements of six Bornean animals. I think 1 1908, Ann. Mag. Nat. Hist.' (8) I, pp. 434-435. 19421 Tate, Results of the Archbold Expeditions. 47 255 With the exception of N. joffrei Thomas With the removal of the short-faced and N. stenopterus Dobson, removed in Malayan stenopterus-joffrei group to form a this paper to a special section of Pipistrellus, special group of Pipistrellus, Nyctalus be- the structure of the rather numerous named comes virtually a temperate-subtropical forms of Nyctalus is remarkably uniform. zone genus. Taken from west to east there Distinctions are based chiefly upon differ- are four European forms, noctula, leisleri, ences of size. The largest known species is lasiopterus = maximus, princeps; in West N. maximus Fatio (= lasiopterus) from Africa and islands, verrucosus, azoreum = Switzerland and Italy; the smallest, N. madeirae; in Transcaspia, meklenburzevi; azoreum. in the Himalayas, montanus, labiatus (?); Miller (1912) distinguished Nyctalus in China, plancei, velutinus; in Japan occur from Pipistrellus and Barbastella by "length three names, aviator = molossus, montanus (of fifth finger) only a little more than that and motoyoshii. of metacarpal of fourth or third" compared Miller's (1912) key to the European with "length greater than metacarpal and species of Nyctalus is built almost solely first phalanx of fourth or third." In "re- upon differences of size, but he also employs marks" he wrote, "This genus is well dif- a hair character (with or without dark ferentiated from allied groups by the pecu- bases). The division with dark hair bases liar narrowing of the wing due to the short- includes only the two smallest species of ness of the fifth finger." the genus, L. leisleri and L. azoreum. Thomas had already pointed out' that The measurements of the species of the difference (breadth of wing) between Nyctalus, as far as known to me, are com- Nyctalus and Pipistrellus was more or less pared beyond. Obviously azoreum and bridged over by oriental forms of Pipis- leisleri are thoroughly distinct species, trellus like P. ceylonicus and P. chrysothrix. markedly smaller also than any others. Earlier (1907) Miller wrote, "skull with The group of very large forms, maximus, nares extending unusually far back, half molossus, aviator, is also sharply marked off. way to interorbital constriction, and with The remaining forms, though they show very large palatal emargination . . . outer considerable variability in size, conform to upper incisor very deeply concave and with a broad pattern which transgresses neither a large anterior and small posterior second- the large nor the small species mentioned ary cusp; canine and p4 always strongly in above. contact; pm3 [p2] very small, quite invisible The large forms may be grouped as two from the outside; canines without trace of geographical races: lasiopterus aviator = secondary cusps; first and second molars molossus, and its European representative, with small but distinct hypocones...." lasiopterus = maximus. To these characters may be added the A slightly smaller species, noctula, has convexity of the frontal area in profile; been made to include several races, N. the generally rather high rostrum; promi- princeps, N. meklenburzevi and possibly nence of the lambdoid crests; very weak N. motoyoshii. It appears to be primarily zygoma, narrowed anterior part of basioc- boreal in distribution. N. lasiopterus and cipital; well developed basial pits; and N. noctula are without secondary basal pits great breadth of the rostrum (almost as and lack dark-colored bases to the pelage. broad as the braincase), swollen in the Following these largest Nyctalus comes supraorbital region but not produced as an oriental group centered upon China supraorbital tubercles (as in Philetor, which includes N. plancei = sinensis and Tylonycteris, Mimetillus and the joffrei N. velutinus. As pointed out by G. M. group of Pipistrellus). The posterior cusp Allen2 these bats may be synonyms. Prob- of il is usually obsolete. The body of P2 ably at least they are conspecific. Both is much shortened in the toothrow as in have distinct secondary basial pits and dark Philetor, etc. brown pelage with lighter tips. 2 1938, Mammals of China and Mongolia, Part 1, 1 1901, Ann. Mag. Nat. Hist., (7) VIII, p. 34. p.235. 256 Bulletin American Museum of Natural History [Vol. LXXX

These special characters are present also Forearm, 40-42 mm. No specimen seen, in the supposed type specimens of labiatus but the small size, the color of the dark Hodgson, B.M. 43.1.12.146, which is rather pelage (Prout's Brown), the less imbricated larger (see under Nyctalus, incertae sedis). lower incisors, less reduced p2, the sub- Hodgson described the molar formula as equal crown-areas of the upper incisors 66.6 y i.e., as in Myotis, so there is doubt combine to set the species off sharply from whether the specimen is truly the type, the larger species of Nyctalus. though I do not question that it was sent by Hodgson from Nepal. Nyctalus montanus (Barrett-Hamilton) With one exception (that of montanus Pterygistes montanus BARRETT-HAMILTON, Barrett-Hamilton, to be mentioned later) 1906, Ann. Mag. Nat. Hist., (7) XVII, p. 99. the remaining species are markedly smaller TYPE LOCALITY.-Mussooree (= Masuri), and more Pipistrellus-like. They have northern India. dark-based pelage with paler tips, and the Size, color, characters . . . very rostra of the skulls are less inflated. They similar to leisleri . .. skull well built, mas- are leisleri, verrucosus = madeirae, and sive, lambdoid crests well developed... azoreum. Probably they differ specifically teeth very large, except small anterior from each other. Compared with the upper premolar... ." Forearm, 42.5 mm. groups previously dealt with they are less The photograph shows the type skull of specialized, or closer to the basic Pipis- montanus, B.M. 79.11.21.164, to be of the trellus-pattern. same general dimensions as that of velu- The remaining form, montanus, from the tinus but shorter and wider and wholly Himalayas, is unusual in possessing the lacking the secondary basial pits of velu- short forearm of verrucosus conjoined with tinus. On the other hand the skull of the much larger skull of velutinus. montanus exceeds considerably the meas- The measurements, chiefly made from urements given by Miller for leisleri of type specimens, shown in the table beyond, Europe. demonstrate and support the species groups just suggested. Nyctalus azoreum (Thomas) Pterygistes azoreum THOMAS, 1901, Ann. Mag. Nat. Hist., (7) VIII, p. 33. Nyctalus verrucosus Bowdich TYPE LOCALITY.-Azores. Nycatlus verruTUC8U BOWDICH, 1825, Excursions This species is the most Pipistrellus-like in Madeira and Porto Santo, p. 36, and footnote. Nyctalus madeirae BARRETT-HAMILTON, 1906, of all Nyctalus seen by me. It is unique in Ann. Mag. Nat. Hist., (7) XVII, p. 98. having preserved the posterior cusp of il; Nyctalus verrucosus ANDERSEN, 1908, Ann. and the crown area of p2, although some- Mag. Nat. Hist., (8) I, p. 434 (discussion of identity of Bowdich's bat). what reduced, is still two-thirds of the TYPE LoCALITY.-Madeira. area of i2, while the canine is less enlarged Forearm, 43 mm. Barrett-Hamilton and bears a distinct rudiment of a posterior stated that madeirae had a quite different cuspule. The C-p4 diastema is not com- skull from that of N. leisleri, "smaller, less pletely closed, and the basial pits are but massive, relatively longer and narrower weakly developed. The zygoma is weak. ... not at all like the much smaller azoreum. The profile of the dorsal surface of the skull ." This appears to be true of the zygo- at the frontal region is slightly convex. matic width but untrue of the lacrimal The rostrum is broad, without supraorbital width (see table). I have had no specimen tubercles, and is thus typical for Nyctalus. from Madeira to study. The general appearance of the skin is that of a pipistrelle rather than of a noctule. The pelage dorsally and ventrally has dark Nyctalus leisleri (Kuhl) bases. Vespertilio leiwleri KUHL, 1819, Ann. Wetterau. The may regarded as an un- Gesellsch. Naturk., IV, pp. 46-49. species be TYPE LoCALITY.-Hanau, Hessen-Nassau, progressive offshoot from the primitive Germany. line leading from Pipistrellus to Nyetalus. 1942 ] Tate, Results of the Archbold Expeditions. 47 257

Nyctalus plancei (Gerbe) doid crests, weak zygomata. The second- Vesperugo plancei GERBE, 1880, Bull. Soc. ary basial pits described in velutinus are Zool. France, V, p. 71. absent or rarely faintly suggested. Crown Vesperus sinensis PETERS, 1880, Monatsber. area of i2 larger than il; diameter of p2 Akad. Wiss. Berlin, p. 258. TYPE LOCALITY.-Peking, China (plancei and about one-half that of i2. Lower incisors sinensis). strongly imbricated. Gerbe's description is almost valueless. This is the characteristic race of Nyetalus Fortunately, however, I obtained a photo- of northwestern Europe. graph of the type skull, B.M. 82.7.29.2. Only the rostrum remains, the braincase Nyctalus noctula princeps Ognev and having been destroyed. Worobiev Color (Peters) "brown, the individual Nyctalus noctula princeps OGNEV AND WORO- BIEV, 1923, Fauna Vertebr. Mamm., Gov't hairs dark brown with their tips paler." Moscou, p. 97. Gerbe gave the forearm length as 47 mm.; TYPE LoCALITY.-Voronesh, Russia. Peters, as 49. This, combined with the A slightly larger, strongly reddish repre- characters of the teeth, palate, rostrum and sentative of the true noctule. The distri- color pattern, indicates relationship to the bution seems to be over eastern Russia medium-sized group with dark hair-bases and Transcaspia to Turkestan. to which velutinus belongs. The dimensional range shown by Ognev is extraordinarily wide, even for a species of Nyctalus velutinus Allen Nyctalus. Nyctalus velutinus G. M. ALLEN, 1923, Amer. Mus. Novitates, No. 85, p. 7. TYPE LOCALITY.-Futsin, Fukien, China. Nyctalus noctula meklenburzevi Kuz- A form about the size of noctula, but jakin Nyctalus noctula meklenburzevi KUZJAKIN, with dark bases to the ventral hairs. Fore- 1934, Bull. Soc. Nat. Moscou, XLIII, 2, pp. arm of type, 49 mm.; c-mi, 7.3. Allen had 323-329. previously referred his material to labiatus TYPE LoCALITY.-Tashkent, central Asia. Hodgson. This race was compared by Kuzjakin The skull is rather readily distinguished directly with noctula. It must, however, be from that of noctula by the secondary set even closer to the race princeps Ognev from of basial pits placed anterior to the pri- eastern Russia and Turkestan, or even mary ones. These secondary pits are placed synonymous. The measurements average virtually in a vertical plane with the glenoid very slightly smaller. fossae and are separated from the primary pits by a low ridge. Both p2 and i2 are Nyctalus noctula motoyoshii Kuroda decidedly smaller. Nyctalus noctula motoyoshii KURODA, 1934, in Closely related to or a synonym of plancei. Siebold, Fauna Japonica (Jap. ed.), 1II, p. 3. See also remarks by Osgood.' TYPE LoCALITY.-Hondo, Japan. This form was also referred to in Kuroda, Nyctalus noctula (Schreber) 1938 (A List of the Japanese Mammals, Vespertiltio noctula SCHREBER, 1775, Siuge- Tokyo, p. 99-100) in which the citation of thiere, P1. LII, pp. 166-167. Kuroda, 1934, is marked "nom. nov." The TYPE LoCALITY.-France. work was published in Japanese, and I have Forearm, 51-54 mm. (Miller). Pelage not seen a copy. "rather dark, yellowish brown near Wood Brown and Cinnamon." The hairs are Nyctalus lasiopterus aviator Thomas without dark bases. Vespertilio molossus TEMMINCK, 1835, Monogr. Skull with the distinctive character of the Mamm., II, p. 269. Vespertilio molossus TEMMINCK, 1850, Fauna noctules, with high, broad rostrum, frontal Japonica. depression absent in profile, strong lamb- Nyctalus aviator THOMAS, 1911, Ann. Mag. 1 1932, Publ. Field Mus. Nat. Hist., Zool. Ser., Nat. Hist., (8) VIII, p. 380. XVIII, pp. 229-230. TYPE LOCALITY.-Japan. 258 Bulletin American Museum of Natural History [Vol. LXXX

Thomas correctly pointed out that Ves- Italy. Ognev (1928) records it under the pertilio molossus Tenuninck, 1850 (and name "siculus" from Bessarabia, Caucasus 1835), was a homonym of Vespertilio molos- and IJral. Siculus, according to Miller sus Pallas, 1767,' a name applied to an (1912), equals Vespertilio murinus. American free-tailed bat described by Buf- fon, but he had no excuse for designating Nyctalus, incertae sedis one of the British Museum specimens, Nyctalus labiatus (Hodgson) B.M. 5.1.4.5, "type." Cotypes of molossus Vespertilio labiata HODGSON, 1835, J. Asiatic Temminck, e.g., Leyden "b," of whose Soc. Bengal, IV, p. 700. skull I have a photograph, exist. Actually TYPE LOCALITY.-Nepal. Jentink2 listed four cotypes, skins "a," "Thick-lipped bat . . . ears shorter than "b" and "c," and skeleton "a." the head, remote, erect, spheroidal; auricle The forearm lengths of skins "a" and of the same form, directed towards the "b" and of skeleton "a" are, respectively, conch of the ear; posterior margin of the 60.5, 59.5 and 61 mm. That of skin "c" is helix folded outward and carried forward 56 and may represent a smaller species, to the gape ... saturate brown throughout. princeps. The forearm of Thomas's ... Teeth262 . . . Labiatais closely "type" was 62 mm. affined to M. Geoffroy's Noctula...." The skull of Leyden "b" is very large An obvious discrepancy exists here. It is and shows some tendency to the formation hard to believe that anyone could describe of a "helmet," the posterior part of the the molar formula of the British Museum sagittal crest as well as the lambdoid crests specimen marked type as " for it is being appreciably raised. No trace ap- unmistakably a species of Nyctalus. Yet pears of secondary basial pits. Upper in- Hodgson so wrote. But he also mentioned cisors: il long and pointed, i2 much shorter its close relationship to noctula Geoffroy,4 and hollowed to receive the tip of lower which equals noctula Schreber. canine. We are faced with the alternatives of (1) an error in observation by Hodgson (the Nyctalus lasiopterus (Schreber) molar formula given by him being that of Vespertilio lasiopterus SCHREBER, 1781, Sauge- Myotis or Kerivoula), or (2) substitution of thiere, P1. LVIII B. a skull of a large Nyctalus for the real type Vesperugo noctula var. maxima FATIO, 1869, skull with six teeth in each molar-premolar Fauna Vert. Suisse, I, p. 57. series. Unfortunately I have no record of The forearm of lasiopterus in Schreber's having seen the skin of B.M. 43.1.12.146, plate measures 67 mm. The forearm length the alleged type of Vespertilio labiata. given by Miller (1912) for maximus is 64- Our photograph of the skull of the same, 68, while Thomas's dimension for the Japa- marked "type" shows not only p, ml but nese aviator is 62 mm. p2 greatly reduced, the typical broad, high The Schreber plate is a relatively excel- rostrum of N. noctula and allies, and the lent portrayal of N. lasiopterus. There same kind of secondary basial pits in line seems to be no reason to suppose it repre- with the glenoid fossae which I have de- sented the smaller Japanese N. aviator. scribed earlier from paratypes of velutinus. This identity of lasiopterus and maximus The measurements of the labiatus skull seems genuine, despite the work of "first given beyond are larger than those of velu- revisers,"3 Fischer and Wagner, who had tinus and about equal to those of noctula or erroneously placed the name lasiopterus as a princeps. The color, "saturate brown," synonym of noctula. suggests again velutinus. The area of distribution of this largest Elsewhere (V. tralatitius Horsfield) I of the noctules is Switzerland, Sicily and accepted the author's statement of "molars, as was 1 1767, Spicillegia Zool., Fasc. 3, p. 8. 6 proof that the bat in question 2 1887, Cat. Ost6ologique des Mammif5res, Mus. a Myotis, or at least not a Pipistrellus. Hist. Nat. Pays-Bas, IX, p. 276. 3 See Thomas, 1911, Ann. Mag. Nat. Hist., (8) 41809, Ann. Mus. Hist. Nat., Paris, VIII, pp. VIII, p. 379. 193-194. 1942] Tate, Results of the Archbold Expeditions. 47 259

IA THOMAS Forearm, 72 mm. Color of pelage very Ia THOMAS, 1902, Ann. Mag. Nat. Hist., (7) dark, blackish brown, with a trace of ashy X, p. 163. tips on some of the hairs; beneath Hair GENOTYPE.-Ia io Thomas. Brown, with the bases fuscous. Terminal Here is a case of relative gigantism among point of tail free. No trace of a calcareal the pipistrelline bats. The only other large- lobe. Ear subtriangular with rounded tip; sized species in that aggregate are Nyetals tragus broad, falcate, with tip rounded. and the tasmaniensis group of Pipistrellus. Photographs of the type skull, B.M. Neither of these approaches Ia in body 2.6.10.2, c, agree closely with our speci- size. men from Szechwan, A.M.N.H. 56873. In In Ia the arrangement of the upper in- the type specimen, however, the basicranial cisors surpasses that seen in Glischropus, area is destroyed, the description of the in that i2 is carried so far forward that the basial pits being drawn from our specimen. four incisive teeth are virtually in a row. Similarly, too, i2 is much reduced in size. SCOTOZous DOBSON But in the case of Ia reduction of i2 is far Scotozous DoBsoN, 1875, Proc. Zool. Soc. greater, for its cusp does not attain the London, p. 372; 1876, Monogr. Asiatic Chirop- cingulum level of i'. The first upper incisor tera, p. 118; 1878, Cat. Chiropt. Brit. Mus., p. 243. is relatively massive and unicuspid (in GENOTYPE.-Scotozous dormeri Dobson of Glischropus bicuspid, the posterior cusp India. almost as long as the anterior), in this re- Africa is obviously the geographical spect resembling Pipistrellus tasmaniensis. headquarters of this genus, provided that p2 iS greatly reduced, its area less than that the numerous described species from that of i2, in spite of the small size of the latter. continent are correctly referable to Scoto- Canine very heavy, half as long again as p4, zous. Hollister' believed the African forms and without enlargement of cingulum or to be a special group of Pipistrellus. They accessory cuspule. Diastema, C-p4 closed. have been so treated in this paper and will Molars very heavy, with hypocone weakly be found under the P. riippellii group of present (in m'-2) on commissure running Pipistrellus. backward from protocone. Lower incisors The possibility that African and Indian profoundly imbricated and canines rather species should not be placed in the same close together; p2 with large crown area genus was indicated unwittingly by Dob- but low cusp, the latter one-half as high son, 1876, when he wrote, "To the subgenus as cusp of p4, the tooth wholly within the [Scotozous] belongs Vesperugo schliefenii ... toothrow. from Egypt. In that species the upper Skull strongly formed: a suggestion of a incisors are much smaller than in V. "helmet," lacrimal width and width of dormeri, the first upper premolar is decidu- braincase moderate; zygoma moderately ous, and the lower incisors are not crowded." deep, with merely a trace of postorbital The incisors of African Scotozous are process; palate narrow, the anterior palatal modified from the Pipistrellus pattern, i2 sinus also narrow. Rostrum not shortened. being minute (by Dobson's original defini- In the basicranial area well developed basial tion of S. dormeri, i2 was absent) and, ac- pits may be observed. cording to Miller (1907), external to the In many respects Ia is merely a very large line between canine and iP (as in P. tenuis Pipistrellus. It could be considered as group, Ia, etc.). Miller wrote that il was representing one of the more specialized bicuspid in the African riuppellii, simple in species groups of that genus. the Indiandormeri (the genotype). Thomas Ia, as far as known, is monotypic. (1915) referred to the variable presence of Ia io Thomas i2 when writing of S. dormeri caurinus. Ia io THOMAS, 1902, Ann. Mag. Nat. Hist., (7) Elsewhere in this present paper an at- X, p. 164. tempt is made to show that the Nycticeini, TYPE LoCALITY.-Chung Yang, south Hupeh, 11918, Bull. U. S. Nat. Mus., XC, p. 89. Allen China. (1939) included them as Pipistrellus. 260 Bulletin American Museum of Natural History [Vol. LXXX

in which iP is unicuspid and i2 obsolete, marked that forms "in the intermediate represent a stage of incisive evolution in area will need further investigation.... advance of that signaled by the Pipistrel- lini, with iP bicuspid and i2 present. CHALINOLOBUS PETERS However, iP in the unicuspid condition Chalinolobus PETERS, 1866, Monatsber. Akad. with P2 still present has been noted occa- Wiss. Berlin, p. 680. sionally in the Pipistrellini, for example, in GENOTYPE.-Vespertilio tuberculatus Forster. P. tasmaniensis and in Ia. Discovery, since Dobson wrote,' of It is possible that Scotozous dormeri repre- Chalinolobus rogersi has weakened the main sents a stage in the transition from the less distinction made between Chalinolobus and specialized pipistrelles to the more special- Glauconycteris, namely, the presence or ized Asiatic members of the Nycticeini. absence of the small upper premolar (p2). This idea is supported by the presence of a The skull of Chalinolobus has a rather "helmet," and by the elongate, slender high, broad rostrum and braincase, weak form of the canines, almost as in Scoto- zygomata without trace of postorbital proc- philus. In Scotozous dormeri ml remains ess, supraorbital area more or less swollen, unreduced, the rostrum and mastoid re- moderately large bullae, very short ros- gions are unbroadened and the skull (rostrum and palate, and large postpalatal trum and braincase) is relatively less de- spine. pressed. p2 iS present in dormeri but may In the dentition, iP may have a posterior be absent in some African species. notch or cusp; i2 is very small, sometimes minute; p2 very small, set in angle between Scotozous dormeri Dobson c and p4, or (in rogersi) absent; m3 never Scotozous dormeri DOBSON, 1875, Proc. Zool. reduced. Soc. London, p. 373. In the skins the lobe at the angle of the TYPE LoCALITY.-Bellary Hills, Assam. mouth accounts for the popular Australian The photograph of the skull of the type name, "lobe-lipped bat"; the ear is broad specimen shows the premaxillaries broken and low, somewhat like the ear of Miniop- away, so that the incisive arrangement can- terus; the basal phalanx of the thumb not be determined directly. But in the pic- large; the attachment of wing to tip of ture of the subspecies S. d. caurinus the metacarpal. appearance of iP (i2 obsolete) conforms The forms of Chalinolobus can be grouped closely to that of Scotophilus. in four principal groups, perhaps four "Fur above brown, the extreme tips species. These are: (a) the gouldi group, ashy; beneath darker brown, the terminal containing the largest bats of the genus, third of the hairs white." Forearm, 33 mm. gouldi and its subspecies venatoris; (b) the According to Kuroda, S. dormeri is pres- tuberculatus group, of which tuberculatus, ent in Formosa. picatus and nigrogriseus are members; (c) the morio group, which includes besides Scotozous dormeri caurinus Thomas morio, signifer and microdon (and perhaps Scotozous dormeri caurinus THOMAS, 1915, australis); and (d) the rogersi group, with J. Bombay Nat. Hist. Soc., XXIV, p. 33. only the small-sized rogersi. TYPE LoCALITY.-Junagadh, Kathiawar, west- The four groups are distinguished by ern India. color: gouldi, with the anterior two-thirds Our photograph of the type skull of this of the body black, the posterior third with race shows no anatomical characters dif- the hairs brown-tipped; tuberculatus, black- ferent from those of dormeri, except slightly ish gray; morio, chocolate brown; rogersi, greater width of rostrum. As pointed out hoary (with dark gray bases). by Thomas the western race is larger, with The distinctness of the four groups of the forearm 36 mm. Chalinolobus is further supported by skull Thomas admitted that caurinus was a characters; the gouldi group has moder- name of convenience, intended to describe 1 1875, Proc. Zool. Soc. London, pp. 381-388; 1878, the larger western race of dormeri. He re- Cat. Chiropt. Brit. Mus., pp. 246-248. 1942 ] Tate, Results of the Archbold Expeditions. 47 261 ately inflated supraorbital swellings, a much larger than tuberculatus and morno trace of a posterior cusp behind i'; i2 and exhibits distinctive characters: il has about one-third height of il above cingulum, a weak posterior cusp; i2 iS proportionately an incipient anterointernal cingulum cusp (to il) smaller but actually equal in size to on p4. In the tuberculatus group the inter- i2 of tuberculatus; it is very much larger orbital area is broadened and the intertem- than i2 of morio; a weak internal cingulum poral constriction accentuated; no trace cusp appears in p4 (as in tuberculatus); the of a posterior cusp appears on il; i2 about degree of development of the supraorbital as large as in gouldi, the cingulum cusp on swellings is less than in tuberculatus, p4 accentuated. The morio group shows the greater than in morio. The subspecies C. g. interorbital width scarcely greater than the venatoris differs but very slightly from intertemporal, a well defined posterior cusp gouldi in dentition, not at all in externals. of i1, i' greatly reduced both in height and The Archbold Collections contain five diameter, no cusp on p4 cingulum. Rogersi specimens of C. gouldi venatoris from north has the intertemporal region much as in Queensland, four from Pentland, on the tuberculatus, il lacking accessory cusp, i2 railroad about 100 miles southwest of small as in morio, p4 with trace of cingulum Townville, and one from Malbon, farther cusp; but the special peculiarity of rogersi west. The table well shows the range of (at least in U.S.N.M. 237822) is the fact measurements of forearm and skull and that p2 is absent and that P2 iS SO much demonstrates that Thomas's type was a reduced that its breadth and length in the small individual of its kind.

SERIES OF (7. g. venatoris, NORTH CENTRAL QUEENSLAND, COMPARED WITH C. g. gouldi 107764 107800 107765 109275 109274 IJ.S.N.M. 221242, o 9 0 9 Gloucester, N.S. W. Forearm 41.0 43.0 43.0 39.0 38.0 44.00 Skull total length 14.6 14.3 14.4 14.0 Zygomatic breadth 10.1 10.4 10.1 10.5 10.5 11.00 Interorbital breadth 6.1 6.0 5.9 6.4 6.2 6.30 Least intertemporal breadth 4.6 4.8 4.5 4.7 4.6 4.50 Breadth braincase 7.8 8.0 7.7 7.9 7.9 8.40 Mastoid breadth 8.7 - 8.8 8.8 9.0 9.40 Diameter tympanic ring 3.4 3.5 3.5 3.6 3.4 3.70 C-m3 5.7 6.0 5.8 5.7 5.4 5.70 M' f Length of crown 1.5 1.6 1.5 1.4 1.4 1.60 Breadth of crown 1.7 1.6 1.5 1.6 1.5 1.50 M2 Length of crown 1.5 1.6 1.4 1.5 1.5 1.60 Breadth of crown 1.8 1.8 1.7 1.7 1.7 1.70 M3 Breadth 1.8 1.8 1.8 1.8 1.8 1.75 toothrow to those of p4 are as 0.5:0.8 and The only way in which the specimen from 0.4:0.6, respectively, and the height of the New South Wales differs from our venatoris two cusps above cingula, 0.4:1.0. is in p4. This tooth is decidedly smaller Neocaledonicus may represent a fifth and presents shorter internal and external group. It was compared with nigrogriseus, faces in the toothrow: external face, 1.0 but, although the forearm length is approxi- (gouldi), 1.15 (venatoris); internal face, 0.8 mately the same, the teeth are very much (gouldi), 0.9 (venatoris). The space be- larger, c-m3, 5.7, being approximately the tween p4 and ml internally is closed in same as in gouldi (in nigrogriseus, 4.9). venatoris, open in gouldi. Chalinolobus gouldi Group Chalinolobus tuberculatus Group Chalinolobus gouldi is a distinctive species This group contains the two species with blackish pelage shading rather sud- picatus (= nigrogriseus, fide Iredale and denly to brown on the rump. The skull is Troughton) and the genotype tuberculatus 262 Bulletin American Museum of Natural History [Vol. LXXX from New Zealand. Not only the scheme sidered synonymous with monrio. It also of coloring but the skulls of these animals has brown pelage. are singularly alike, particularly in regard C. australis, type locality not known to the lack of the cusp on the posterior face (Tasmania, fide Iredale and Troughton), of il and in the relatively broad interorbital is also regarded as equal to morio. and narrow intertemporal widths. Alphabetical list of named forms of C. tuberculatus is a species of medium Chalinolobus: size (forearm, 39 mm.). Contrary to the australis Gray, 1841, in Grey, Journals Two view expressed by Thomas' in 1889, tuber- Exped. Australia, II, pp. 400, 406. culatus is not precisely equal to morio. The No type locality (Australia). following comparison shows the distinction Described as Scotophilus. Regarded between the two: tuberculatus (U.S.N.M. as synonymous with morio. 38031/14003 from New Zealand) is wider gouldii Gray, 1841, in Grey, Journals Two than morio across the canines, 4.6:4.0; il Exped. Australia, II, pp. 400, 405. in tuberculatus is single, in morio with pos- TYPE LoCALITY.-Australia (restricted terior cusp; height of cusp of i2 above cin- to Launceston, Tasmania, by Thomas). gulum, 0.5:0.3; intertemporal width, 4.1: MATERIAL.-Photograph of type, B.M. 4.7; interorbital width, 5.4:5.1 (note the 41.1516 (sic). change of proportions in the last two items); Coloring almost exactly as in venatoris. diameter of tympanic ring in tuberculatus, The largest species of Chalinolobus. 3.0, of morio, 2.8; p4 in tuberculatus with microdon Tomes, 1860, Ann. Mag. Nat. anterointernal cingulum cusp, in morio Hist., (3) V, pp. 51-53. without such cusp; length of lower tooth- TYPE REGION.-Tasmania. row (c-m3), 5.4:5.0. MATERIAL.-Photograph of mandible only of type, B.M. 41.1513. Chalinolobus morio Group Described as Scotophilus. Dorsal color near brown, ventral color C. morio, U.S.N.M. 236766, from Kanga- slightly paler, with gray bases. Con- roo Island, South Australia, has fore- sidered synonym of morio. arm 38.5 mm. In the skull the interorbital morio Gray, 1841, in Grey, Journals Two width (5.2) scarcely exceeds the intertem- Exped. Australia, II, pp. 400, 405. poral width (4.7); the zygomatic breadth TYPE REGIoN.-South Australia ac- only slightly exceeds the mastoid width cording to Dobson (restricted by Iredale (8.7:8.5). The first incisor has a well de- and Troughton to Tasmania). veloped internal secondary cusp, while i2 MATERIAL.-Photograph of type skull, is exceedingly minute (height of its cusp B.M. 37.4.8.118 (90a). above cingulum, 0.3 to 0.4; in tuberculatus, Erroneously made a synonym of 0.6); c-m3, 4.6. tuberculatus (Thomas, 1889), but nearest Closely related or perhaps identical to ally appears to be picatus (see discussion morio is a specimen, M.C.Z. 14927 (in above). alcohol, skull cleaned), from Broome Hill, Dorsal color chocolate brown. A Western Australia, previously identified as strongly marked post-calcareal lobe, "C. signifer." Forearm, 38 mm.; basal which itself is supported by a minute off- phalanx of thumb, 3.8; skull, greatest shoot of the calcar. length, 12.7; zygomatic breadth, 8.8; neocaledonicus Revilliod, 1914, in Sarsain mastoid breadth, 8.1; interorbital width, and Roux, Nova Caledonica, Zoologie, 5.1; intertemporal width, 4.6; c-m3, 4.6; I, p. 355. ml-3, 3.15. Iredale and Troughton regard TYPE REGION.-New Caledonia. signifer as equal to morio. Above pelage uniform brown- C. microdon (forearm, 37 mm.), concern- black, a light brown tuft of hairs being which I have little data (photograph hind each ear ... forearm, 35.3 . . . skull, of the mandible only), is currently con- basilar length, 10.5; mastoid breadth, 1 Ann. Mag. Nat. Hist., (6) IV, p. 462. 8.8; zygomatic breadth, 10; c-m3, 5.7." 1942] Tate, Results of the Archbold Expeditions. 47 263

Compared by Revilliod principally study I have had before me a specimen with nigrogriseus. from Dunedin (U.S.N.M. 38031/14003). nigrogriseus Gould, 18521 (1863), Mammals Nearest Australian relative, picatus, Australia, III, P1. XLIV (with text). not morio. TYPE LoCALITY.-Moreton Bay, south vagans Dobson, 1879, Ann. Mag. Nat. Hist., Queensland. (5) IV, p. 135. MATERIAL.-Photograph of type skull, TYPE LOCALITY.-"Bermuda." B.M. 56.10.28.3. MATERIAL.-Photograph of type skull, Color dorsally fuscous; beneath fus- B.M. 79.1.7.1. cous, with tips of hairs gray. Forearm, Described as Vesperugo. Synonym of 36 mm. This species apparently equal to Chalinolobus tuberculatus (Thomas, 1915, picatus. Proc. Biol. Soc. Washington, XXVIII, p. picatus Gould, 18521 (1863), Mammals 69). Australia, III, PI. XLIII (with text). venatoris Thomas, 1908, Ann. Mag. Nat. TYPE REGION.-Northwestern New Hist., (8) II, p. 372. South Wales. TYPE REGION-Northern Territory, MATERIAL.-Photograph of skull of Australia. type, B.M. 53.10.22.33. MATERIAL.-Photograph of skull, Described as Scotophilus. B.M. 6.3.9.4. Color black above and beneath. Fore- Described as race of gouldi. .... Fore- arm, 32 mm. Currently believed equal arm, 40 mm., skull, greatest length, 14; to nigrogriseus. Picatus appears to be the braincase breadth, 7.6...." closest of the Australian species to tuber- Dorsal color blackish brown with culatus of New Zealand. brown rump; ventrally very dark brown rogersi Thomas, 1909, Ann. Mag. Nat. anteriorly, and light brownish buff pos- ffist., (8) III, p. 150. teriorly. External appearance exactly as TYPE LoCALITY.-Wyndham, north- gouldi. western Australia. MATERIAL.-Photograph of type skull, GLAUCONYCTERIS DOBSON Glauconycteris DOBSON, 1875, Proc. Zool. B.M. 9.4.23.1. Soc. London, p. 383. "A small species, black, with hoary GENOTYPE.-Kerivoula poensis Gray. tips to the hairs. Most nearly allied This wholly African genus was originally to C. nigrogriseus . . . a well-marked proposed as a subgenus of Chalinolobus, postcalcareal lobe . . . forearm, 34.5; now limited to the Australian region. condylobasal length, 12.2; zygomatic In 1906 Miller2 recognized that G. breadth, 8.6; c-m3, 4.5...." floweri, which he separated under the ge- signifer Dobson, 1876, Ann. Mag. Nat. neric term Rhinopterus, had nothing to do Hist., (4) XVII, p. 289. with Glauconycteris but was actually closer TYPE LOCALITY. PeakDowns, Queens- to Eptesicus. land. Seven species of Glauconycteris are given MATERIAL.-Photograph of type skull, by Allen in the "Checklist of African B.M. 76.3.29.11. Mammals" Treated by Iredale and Troughton as (1939). equal to morio. I am unable to separate BARBASTELLA GRAY a specimen from northwest Australia Barbastella GRAY, 1821, London Medical Re- (M.C.Z. 14927) from specimens from pository, XV, p. 300. South Australia and Kangaroo Island. Synotus KEYSERLING AND BLASIUS, 1839, tuberculatus Forster, 1844, Descript. Anim. Archiv. f. Naturg., V, Part 1, p. 305. Itin. ad Maris Austral. Terres, p. 62. GENOTYPE.-Vespertilio barbastellus Schreber. TYPE REGION.-New Zealand. The teeth of Barbastella are in virtually MATERIAL.-For purposes of this the same stage of modification as the teeth of Pipistrellus, i.e., il is bifurcate, i2 slightly 1 Fide Iredale and Troughton, 1934, Mem. Austra- lian Museum, VI, p. 97. 2 Proc. Biol. Soc. Washington, XIX, p. 85. 264 Bulletin American Museum of Natural History [Vol. I,XXX reduced; p2 exceedingly minute and dis- jelingensis Dobson (= blanfordi), because placed inward by the already contiguous c he believed darjelingensis Hodgson to be a and p4; m3 unreduced; lower incisors im- Plecotus. bricated, trifid, with additional internal But there is little actual reason for so cusp on i3; crown of P2 one-third of height complex an explanation of the problem. of cusp of P4. Special characters of teeth Homochrous, with which Horsfield com- and skull are seen in the anterointernal pared his darjelingensis, is a Plecotus; I cingulum cusp on p2; the broadly depressed have a good picture of the type skull. Bi- rostrum and median sinual spine; the ex- anchi's chief stumbling-block in Horsfield's treme weakness of the zygomata. The description of darjelingensis lies in the ears are proportionately shorter than those words "the dimensions are the same [as of Pipistrellus, and the bullae are unin- homochrous] in all points." Horsfield con- flated. tinued, "the color is deeper, inclining to In quite early times' Barbastella was re- blackish ['uniform obscure brown' in garded as a close relative of Plecotus. Per- homochrous]. The lobes of the ear are haps because of this confusion, Horsfield2 spreading with a small appendicule at the published his description of darjelingensis, bases ['Ears enormous, 2'/2 of head, ellip- collected by Hodgson in Nepal, under tic, nude, transverally striolate']. The Plecotus. I have a photograph of the type tragus is narrow [Inner ear narrow, pointed, skull, B.M. 54.9.1.3., which shows distinctly erect, with a small basal process for tragus, the median spine of the much broadened answering which is a small internal anti- narial sinus and thus allocates darjelingensis tragus ]. . . ." The matter in single quotes Horsfield (Hodgson) definitely within Bar- refers to Plecotus homochrous. Apart from bastella. Dobson3 placed "darjelingensis" the implication that the dimensions of the and barbastellus in Synotus Keyserling and ear of darjelingensis Horsfield are as great Blasius, 1839 (genotype V. barbastellus as those of homochrous, the description Schreber). His description of the ear quoted above of the former is perfectly states "the outer margin has no projecting suited to Barbastella. I do not think the lobe at the junction of its upper and middle "small appendicule at the base" (Hors- third...." Bianchi4 assumed, on the basis field) equivalent to the "small well-defined of the wording of Horsfield's description, circular projecting lobe at the junction of that darjelingensis Horsfield (Hodgson) was the upper and middle third" of the outer in reality a Plecotus and suggested a mix-up margin of the ear (Dobson). But if it were, of labels, whereby Dobson (in Bianchi's darjelingensis Horsfield would be a repre- opinion) was writing, as he was, of a Bar- sentative of the barbastellus group and the bastella. Just why Synotus darjelingenis lobeless specimens before me, as I write, Dobson, which Bianchi renamed Barba- from United Provinces, northern India, stella blanfordi, should be considered a would have to bear some other name. synonym of Plecotus darjelingensis Hors- Ognev5 accepted Bianchi's conclusions. field, assuming for the moment that Bianchi The following forms of Barbastella have was right in his surmise, that author did not received names: explain. barbastellus Schreber from Burgundy, France (synonyms: daubentonii Bell, communis Gray) Bianchi (tom. cit.) recognized two divi- b. caspica Satunin from Caucasia sions of Barbastella, one, typified by bar- b. walteri Bianchi from Transcaspia bastellus, with projecting lobe from the leucomelas Cretschmar from Arabia Petraea base of the ear; the other, without such darjelingensis Horsfield from Nepal (synonym: lobe, represented by "darjelingensis Hodg- blanfordi Bianchi) son." He must have meant instead dar- Ognev (tom. cit.) placed darjelingensis Dobson, caspica and walteri together under 1 1838, MacGillivray, Naturalist's Library, XVII, caspica Satunin, 1908. Even if these races pp. 83-84. 2 1855, Ann. Mag. Nat. Hist., (2) XVI, p. 103. were synonymous they should have been 3 1878, Cat. Chiropt. Brit. Mus., p. 177. 4 1916, Annuaire Mus. Zool. Acad. Sci. Petrograd, 5 1928, Mammals of Eastern Europe and North- XXI, p. lxxiii. ern Asia, I, p. 588. 19421 Tate, Results of the Archbold Expeditions. 47 265 named darjelingensis, the oldest name. form dark brown coloration, the heavy According to Ognev's key (tom. cit., p. 584) thumbs and large ears; also in the skull, none of them has the supplementary lobe the short, broad rostrum and very strongly on the outside of the ear. Leucomelas, as developed supraorbital tubercles, as well as illustrated (tom. cit., P1. XXVIII, figs. b, 3, 4), the remarkable posterior-basal cusp on the has rather larger ears than typical Bar- canine, the much shortened bodies of the bastella; it lacks the accessory lobe, and the premolars, p4, and the scarcely reduced P2, bullae are small (not large as in Plecotus). which is virtually as high as p4. Almost Height given for ear, 7 lines (= 14 mm.). the only point of difference in Philetor is the Hairs of lower body with white tips. The disappearance of p2. That tooth in all of description supports the view that Vesper- the Pipistrellus-like genera is a vanishing tilio leucomelas is a Barbastella. The ears, structure. as illustrated, are considerably larger than those of the type species. Philetor rohui Thomas Provisionally the following arrangement Philetor rohui THOMAS, 1902, Ann. Mag. of the forms of Barbastella is suggested: Nat. Hist., (7) IX, p. 220. B. barbastellus TYPE LoCALITY.-Albert Edward Range, B. darjelingensis (= caspica = walteri = blan- New Guinea, 6000 feet. fordi) B. d. leucomelas Forearm, 34.5 mm. The following comparison is drawn from I have excellent pictures of the type skull barbastella (M.C.Z. 37000) from Berlin and of this species, B.M. 1.11.24.11, 6. The darjelingensis (F.M.N.H. 48571, 48573 zygomata are weak as in Nyctalus. The [young]) from Mussoorie, TUJnitedProvinces, posterior process of the palate, in place of northern India. These species differ being spine-like, is broadly ligulate in form. chiefly, as stated earlier, by the presence or No secondary basial pits. Expanse across absence of the supplemental lobe on the supraorbital tubercles, 8.2 mm. outer margin of the ear. In addition the B.M. 3.3.5.3-4 are two specimens from pinna of darjelingensis is larger and less the Owen Stanley Range, both of which abruptly truncated or "broadly rounded have forearm 32 mm.; B.M. 8.10.8.60, in- off" (Dobson). No significant color dif- land from Port Moresby, has forearm 35 ferences appear; both are blackish gray mm. It appears that the forearm in this with the tips whitish. In the young speci- species varies at least from 32 to 35 mm. men the grizzled tips have not developed. A male specimen, A.M.N.H. 152466, In both, the membrane attaches to the end from the Idenburg River in the Archbold of the metatarsal; a calcareal lobe is pres- Collection is slightly larger than the type, ent. its forearm, 37, and c-im3, 5.0 mm. It In spite of the quite significant difference looks conspicuously different from other in the ears, the skulls show few distinguish- pipistrelloid bats recently examined, by ing characters: the premaxillae and in- possessing a prominent, pointed, palatal cisors are slightly more projecting in the spine projecting forward 1 mm. in the center case of darjelingensis and the diameter of of the anterior palatal sinus, which appears the tympanic ring is greater (3.1:2.9). to be the counterpart of the generally present posterior palatal spine (ligulate in PHILETOR THOMAS Philetor). I hesitate to make this spine the Philetor THOMAS, 1902, Ann. Mag. Nat. Hist., basis of a new race (in company with the (7) IX, p. 220. larger measurements of our specimen) be- GENOTYPE.-Philetor rohui. cause in the type the spine may have been Thomas compared Philetor with "Ptery- present and been broken off. gistes" = Nyctalus, showing that it had one less upper premolar. MIMETILLUS THOMAS In many ways Philetor resembles the Mimetillus THOMAS, 1904, Proc. Zool. Soc. Pipistrellus joffrei group (ioffrei was origi- London, II, p. 188. nally described as a Nyctalus): the uni- GENOTYPE.-Vesperugo moloneyi Thomas. 266 Bulletin American Museum of Natural History [Vol. LXXX

This peculiar African genus has been re- developed; the canine width remains un- garded as related to the oriental Tylonyc- exaggerated; the zygoma is weak. teris. It has achieved a relative degree of Perhaps we may regard Tylonycteris and flatness of the skull. It has also developed Philetor as independently derived from near large orbital swellings' and the obsolescence the Pipistre7lus joffrei group, and as having of p2. But certain characteristics about it lost p2 independently. suggest its independent origin from some The flattening of the skull is an attribute branch of Pipistrellus other than the line of Tylonycteris only. It is not to be traced which led to Philetor and Tylonycteris. In in Philetor. Mimetillus the zygoma is deep and strong A considerable number of forms of Ty- and exhibits distinct signs of a postorbital tonycteris have received names: enlargement. The canine is longand slender with no lengthening of the cingulum pos- pachypus GROUP teriorly to accommodate the posterior pachypus Java accessory cusps of Philetor and Tylonyc- fulvidus (= rubidus2) Sittang River, Burma meyeri Philippines teris; the lower incisors are scarcely im- aurex Western India bricated and are arranged in an open transverse arc around the broadened symphysial robustula GROUP portion of the mandibles; P2 and p4 are robustula Sarawak, Borneo not arranged as in the genera mentioned, malayana Perak, Malaya instead P2 becomes much reduced in size, while p4 remains unshortened in the axis of Thomas was right in part in 19153 when the jaw. he recognized three types of Tylonycteris, In the skin the large ears, less reduced but only two main divisions actually occur. thumb, the lack of tumescence in the foot- His attempt to partition this genus was and thumb-pads, and the whitening of the held later to have failed4 until Osgood5 wings are characters foreign to Tylonycteris. concluded that two distinct types existed. In this paper no attempt can be made to Even so, G. M. Allen6 again expressed place the origin of Mimetillus. No African doubt. groups of Pipistrellus have been studied. I am able, I believe, to point to two ana- It may come from a specialized African tomically proportional characters, in addi- group, or it may be derived from Eptesicus tion to absolute differences of size and color, ancestry in which p2 was already lost. which place any species of Tylonycteris definitely in one of two groups. The first of TYLONYCTERIS PETERS these characters relates to the degree of de- Tylonycteris PETERS, 1872, Monatsber. Akad. velopment of the supraorbital tubercles, Wiss. Berlin, p. 703. extreme in the larger robustula, minimal in GENOTYPE.-Vespertilio pachypus Temminek. fulvidus, aurex, etc. The character is ex- If the extraordinary flattening of the skull pressed clearly in the table beyond by set- of Tylonycteris is for the moment disre- ting "width across supraorbital tubercles" garded, its similarity to Philetor becomes against "lacrimal width," this latter being apparent. The upper incisors are much the the minimum interorbital width taken same, the canine has developed a strong across the orbits infront of the supraorbital accessory cingulum cusp; p2 is absent; the tubercles. In the larger forms with dark lower premolars have much the same char- pelage this differential appears as 0.5 mm. acter, P4 much shortened longitudinally in and upward; in the smaller fulvous forms the toothrow, and the height of p3 only 2 This name (nomen nudum) apparently used er- inferior to roneously by authors for fulvidus. It was attributed slightly that of p4. In the skull to the same type locality as fulvidus by Thomas and supraorbital tubercles are more weakly by Gyldenstolpe. 3 Ann. Mag. Nat. Hist., (8) XV, p. 226. 4 Wroughton, 1917, J. Bombay Nat. Hist. Soc., 1 These swellings are not strictly the homologues XXV, p. 586. of the supraorbital tubercles in Tylonycteris but are 5 1932, Publ. Field Mus. Nat. Hist., Zool. Ser., swellings of the area anterior to that part of the orbit XVIII, p. 235. which incloses the anteorbital foramen, but posterior 6 1938, Mammals of China and Mo.agolia, Part 1, to the foramen. pp. 247-249. 1942] Tate, Results of the Archbold Expeditions. 47 267 as 0.1-0.2 mm. In the small species the "c," 27. In the specimen in the British tubercles are minute, in the large species Museum (44.4.4.24), which Thomas stated substantial. was a cotype received from Leyden, the The second distinct difference lies in the forearm measures 26.4. degree of enlargement of the lambdoid The foregoing dimensions alone make it crests. In the small species these crests are reasonably sure that pachypus was a strongly developed and on each side just "small" Tylonycteris. The skull of speci- above the condyle are drawn strongly back- men "e," which I studied, fully confirms ward so that, seen from above, the two that view, possessing relatively slight ros- backwardly projecting lambdoid crests and tral tubercles and prominent lambdoid the bulge of the supraoccipital form almost ridges. a straight line. In the large species, on the I am unable to distinguish between Bali- contrary, the crests are poorly developed, nese and Sumatran skulls in the Archbold are scarcely drawn backward and do not Collections and skulls from Laos, Indo- form a straight line with the bulge of the China, which represent fulvidus. At best supraoccipital. fulvidus (= rubidus) can be but a slightly The type species of the genus, pachypus, more rufescent race of pachypus. is a member of the smaller group. This T. aurex, the form from western India, is shown by the small toothrow of the co- may differ very slightly; T. meyeri from the type, Leyden "e," and the narrowness of Philippines appears much smaller. the measurement across the supraorbital In the Archbold Collections are series tubercles, as well as by Temminck's state- from Palembang, Sumatra (17); Selat, ment about the color of the skin, ".... bi- Bali (5). color ... above more or less lustrous maroon ... with bases more or less golden red; Tylonycteris pachypus fulvidus (Blyth) chest red, brown at the endsofthe hairs... Scotophilus fulvidus BLYTH, 1859, J. Asiatic Specimen "e" came from Bantam, Java. Soc. Bengal, XXVIII, p. 293. To this group can be referred fulvida, Tylonycteris rubidus THOMAS, 1915, Ann. Mag. Nat. Hist., (8) XV, p. 227. aurex and probably meyeri, a considerably TYPE LOCALITY.-' 'From Schwe Gyen," Sit- smaller bat. tang River, Burma. The robustula group should probably in- I have not found an earlier usage of clude malayana, which Chasen shows to be rubidus (used also by Gyldenstolpe, 1916). ,even larger than typical robustula. The name seems to have been substituted inadvertently for fulvidus. Tylonycteris pachypus Group "A small Scotophilus ... of a pale fulvous color throughout, with black membranes. Tylonycteris pachypus (Temminck) Length, 23/4 inches, of which tail 1 inch; Vespertiliopachypus TEMMINCK, 1835, Monogr. ... forearm, 11/16 inches [27 mm.]; ... 4 Mamm., II, p. 217. Java. females." TYPE LOCALITY.-Bantam, The type is probably at Calcutta. The. only specimens at Leyden marked Thomas, Gyldenstolpe, Osgood and G. by Jentink as types were mounted speci- M. Allen have at various times referred bats mens "a-c" and alcoholic specimen "e." from Burma and Indo-China to this race. Of skeletal material, "a-h" were stated to be types. All were from Bantam, Java. Tylonycteris pachypus aurex Thomas Dr. Junge was good enough to have the Tylonycteris aurex THOMAS, 1915, Ann. Mag. skull of specimen "e" (alcohol) taken out Nat. Hist., (8) XV, p. 228. and cleaned. My notes on skull and den- TYPE LOCALITY.-Astoli, Belgaum, south of tition were derived from that skull. The Bombay, western India. following are forearm lengths of seven co- My photograph of the skull of aurex types: skin "a," 27 mm.; skin "b," 26; and the measurements taken therefrom ;skin "c," 25; alcoholic "e," 26; skeleton offer no point of differentiation from pachy- "a," 26.5; skeleton "b," 25; skeleton pus. Aurex is maintained a geographical 26S, Bulletin American Museum of Natural History [Vol. LXXX race upon Thomas's authority. We have Tylonycteris malayana Chasen no specimens. Tylonycteris malayana CHASEN, 1940, Bull. Raffles Mus., XV, p. 52. Tylonycteris pachypus meyeri Peters TYPE LOCALITY.-JOr, Batang Padang, Perak, Tylonycteris meyeri PETERS, 1872, Monatsber. Malay States. Akad. Wiss. Berlin, p. 705. There is only Dr. Chasen's description TYPE LOCALITY.-South Luzon, Philippines. of this bat to go upon. He was disinclined This little bat, forearm, 22.3-24 mm., is to regard it as a race of robustula. Forearm, retained as a distinct race of pachypus on 28.5 mm.

RATIO OF LACRIMAL WIDTH TO WIDTH ACROSS SUPRAORBITAL TUBERCLES IN Tylonycteris Lacrimal Width TuberCle Width Differential T. robustula group A.M.N.H. 109144 Peleng 6.0 6.9 0.9 A.M.N.H. 109145 " 5.9 6.8 0.9 F.M.N.H. 32158 Laos 5.7 6.2 0.5 F.M.N.H. 32157 " 5.6 6.1 0.5 M.C.Z. 27067 Tonkin 5.6 6.3 0.7 A.M.N.H. 102641 South Sumatra 5.9 6.7 0.8 A.M.N.H. 102642 It 5.6 6.8 1.2 T. pachypus group F.M.N.H. 32160 Laos 4.8 5.0 0.2 F.M.N.H. 32189 " 5.1 5.2 0.1 M.C.Z. 27066 " 5.0 5.2 0.2 account of its smaller size. Peters had HESPEROPTENUS PETERS three cotypes. I have a photograph of the Hesperoptenus (subgenus of Vesperugo) skull of B.M. 75.11.3.12, 9. As shown in PETERS, 1868, Monatsber. Akad. Wiss. Berlin, p. 626. the table, its dimensions are generally smaller than those taken from the cotype To Peters' genotype, Vesperus doriae of pachypus. from Sarawak, Dobson' added Nycticejus tickelli Blyth, 1851 (with its supposed Tylonycteris robustula Group synonym N. isabellinus Horsfield, 1851) from India and Ceylon, and Vesperugo Tylonycteris robustula Thomas (H.) blanfordi Dobson, 1877, from Tenas- serim. Tylonycteris robustula THOMAS, 1915, Ann. Mag. Nat. Hist., (8) XV, p. 227. Contrary to the conditions in Scotomanes, TYPE LoCALITY.-Sarawak, Borneo. in which the three "races" are doubtfully Thomas pointed out the larger size and separable, Hesperoptenus, if not poly- dark brown coloring by which this species phyletic, contains very strongly differ- could be distinguished from T. pachypus entiated species. Miller2 rightly suggested and allies, and earlier in this paper I have that more than one generic group might be indicated two additional cranial differences. involved. Thus in doriae, the type species, Thomas gave the range of this species as Peters makes no mention of any unusual Malay Peninsula, Borneo, Java, Celebes position of i2, merely calling it "low and and Timor. Osgood (1932) recorded it from small." Had it stood behind il as in the Indo-China, and G. M. Allen (1938) re- mainland species tickelli he would scarcely corded three specimens from Yunnan, have failed to notice the fact. Again the China. There is no indication that it relationships of the anteorbital foramina reaches India. to the supraorbital ridges in tickelli and The Archbold Collections contain good tomesi show them to be "good" species. series from Palembang, Sumatra (5); Selat, The remarkably small size and specialized Bali (8); and Peleng Island, east of Celebes '1878, Cat. Chiroptera Brit. Mus., p. 240. (13). 2 1907, Families and Genera of Bats, pp. 211-212- 19421] Tate, Results of the Archbold Expeditions. 47 269 thumb of blanfordi make one suspect that dentition ... short, flat, obtusely trilobate it, too, differs widely. or quadrilobate second pair of upper in- I have seen specimens of tickelli and cisors, situate posterior to the usual large tomesi only, but it is probable that, when pair, and immediately behind the contact of the skull of the type of doriae has been each of the latter and the canine of the same studied, these.larger bats may require re- side... consideration from the generic standpoint. Herewith a few notes made on M.C.Z. Peters gave virtually no generic char- 27514-15 from Ceylon: A "helmeted" acters with doriae, and Miller's definition skull with high mandibular coronoid; il of Hesperoptenus, based chiefly upon tickelli, without trace of secondary cusp; ji inward may have to be recast. from i', the reverse of condition in Ia; In H. tickelli an interesting condition of crown area of i2 about equal to il; p2 ab- the upper incisors exists (in addition to sent; m3, though short, with nearly com- the position of i2), namely, the unicuspid plete pattern; lower incisors much thick- condition of iP. In most Pipistrellus-like ened, strongly imbricated; P2 wider than genera il remains bicuspid, and in the long (in jaw), height of its cusp two-fifths Scotophilus-like genera obsolescence of the of P4. Supraorbital processes angular, with- posterior cusp is accompanied by loss of i2. out conjunction with anteorbital foramina In tickelli an intermediate condition is seen, as in tomesi. whereby the unicuspid state is achieved in il before obsolescence of i2. [Hesperoptenus isabellinus (Horsfield)] Nycticejus isabellinus HORSFIELD, 1851, Cat. Hesperoptenus doriae Peters Mamm. Mus. East India Comp., p. 38. Hesperoptenus doriae PETERS, 1868, Monats- " . . Size of N. temminckii [= Scoto- ber. Akad. Wiss. Berlin, p. 626. philus temminckii], clearly characterized by TYPE LocALITY.-Sarawak, Borneo. a uniform isabellina tint both above and ". . . Trilobed lower incisors stand oblique underneath." to edge of jaw ... second upper incisor low Dobson (1876) treated isabellinus as a and small .... Light brown, dorsal hairs synonym of tickelli. No locality was given. darker at bases.... Forearm, 36.5." Apparently Dobson (1876, p. 112) was responsible for the association of tickelli, Hesperoptenus tomesi Thomas and other forms in which i2 stands behind Hesperoptenus tomesi THOMAS, 1905, Ann. Mag. Nat. Hist., (7) XVI, p. 575. ii, with doriae. He wrote, "Most probably TYPE LOCALITY.-Malacca. . . .it will be found that they can both [doriae and tickelli] be included in the sub- Our picture of the type skull, B.M. 7.1.1. genus Hesperoptenus." He went so far in 428, shows a moderately short rostrum with the "Catalogue" (1878, p. 239) as to con- wide supraorbital area which is, however, trast doriae "outer incisors in the tooth- pinched in sharply at the level of the ante- row," with all other species, "outer incisors orbital foramina; back of skull raised and quite posterior to the toothrow." The first back part of sagittal crest moderately de- of those statements remains, I believe, yet veloped into a "helmet"; mastoid area to be proved. slightly expanded as in Scotomanes. Ca- nines very long; i2 strongly displaced in- Hesperoptenus tickefli (Blyth) wardly; P2 not compressed in toothrow; Nycticejus tickelli BLYTH, 1851, J. Asiatic Soc. lower incisors well imbricated. Bengal, XX, p. 157. "General color . . burnt umber but TYPE REGION.-"Central India, Ceylon, and darker . . . under surface uniform brown doubtless the intervening country." (almost matching Ridgway's Burnt Um- ". . . Forearm, 23/8 inches (61 mm.) ... ber) throughout. . . i, which is also very fur ... pale fulvescent or whitish fulvous ... large, placed quite behind il ... premolar considerable growth of hair upon basal much extended transversely . . . forearm, half of interfemoral membrane above . . . 51 mm.... only near ally H. tickelli." 270 Bulletin American Museum of Natural History [Vol. LXXX

Hesperoptenus blanfordi Dobson pertilio and Eptesicus as synonymous. Ten Vesperugo (H.) blanfordi DOBSON, 1877, J. years later, following the views of M6hely,2 Asiatic Soc. Bengal, XVI, p. 312. Miller,3 recognizing Eptesicus, restricted TYPE REGION.-Tenasserim. Vespertilio to contain only murinus and "...A broad adhesive cushion occupies superans. the base of the inferior surface of the meta- The formation of the skull, especially the carpal bone [of the thumb], and extends massive rostrum, and the dental formula outward and backward upon the base of the indicate that the bats of this genus are metacarpal of the second finger. allied on the one hand with Nyctalus and on "Forearm, 1.1 inches (27 mm.). the other with Eptesicus. The tricuspid "Fur reddish-brown above, slightly paler of il is a while the beneath . . . . Dentition almost similar to condition specialization, that of V. tickelli, the outer incisors even cusp of the outer incisor is less reduced than smaller and placed more behind the inner in Eptesicus. The canines are unusually incisors...." broad at the cingulum level. Last upper Here is a digital adaptation similar to molar unreduced. that of Glischropus. The size of the type species, murinus, is

subgenera Rhinopterus Vespertilio I Rostral and palatal sinuses Histiotus broadened Eptesicus Laephotis

Ears and bullae Ears and bullae not enlarged greatly enlarged

Eptesicoid genera p2-5 absent; i2 present

Fig. 4. Suggested phylogeny of the eptesicoid genera. See also Fig. 1. VESPERTILIO LINNAEUS about equal to that of Eptesicus nilssonii, Vespertilio LINNAEUS, 1758, Syst. Nat., I, the forearm, 43-45 mm. 10th ed., p. 31. GENOTYPE.-Vespertilio murinus Linnaeus. Vespertilio is placed next to Eptesicus Vespertilio Linnaeus, 1758, contained all only provisionally, on account of the loss of the seven bats described by him, of which of p2. However, the broadened rostrum two only, auritus [Plecotus] and murinus, and especially the width of the anterior belonged in the present family Vesper- narial and palatal sinuses make it equally tilionidae. The type was fixed in 1897 by probable that it has a co-origin with Nyc- Miller,1 who at that time regarded Ves- talus and has since lost p2. 1 1897, North American Fauna, XIII, pp. 18-19, 2 1900, Monogr. Chiropt. Hungariae, p. 219. 95-103. 3 1907, Families and Genera of Bats, pp. 207, 209. 1942] Tate, Results of the Archbold Expeditions. 47 271

Until recently only V. murinus and its genus by Miller,3 nilssonii Keyserling and geographical race superans were known. In Blasius, 1839, from Sweden, and sodalis 1920 Kuroda described namiyei. V. dis- Barrett-Hamilton from Roumania, were color is a synonym of murinus, and siculus described later. The American fuscus and has also been used for that species. European serotinus are unquestionably con- subgeneric. They are rather large, with Vespertilio murinus Linnaeus c-m3 approximately 7 mm. In both, m3 is Vespertilio murinus LINNAEUS, 1758, Syst. strongly modified, and the skull is drawn Nat., I, 10th ed., p. 31. Vespertilio discolor KUHL, 1819, Ann. Wet- back into an occipital "helmet." terau. Gesellsch. Naturk, IV, p. 187. Since Rafinesque originated the genus, Vesperus siculus DADAY, 1885, Orvos-Termes- the addition of numerous species, including zettudomanyi, X, p. 275. tropical forms from Africa, Australia and TYPE LOCALITY.-Upsala, Sweden (discolor, Vienna; siculus, Homorod-Almas Cave, Hun- South America, has necessitated such gary). broadening of the generic scope of Eptesicus Forearm, 43-45 mm.' that, apart from the similar dental formula, many species are found to have little in Vespertilio murinus superans Thomas common with melanops Rafinesque. Vespertilio murinus superans THOMAS, 1898, This dental formula differs from that of Proc. Zool. Soc. London, p. 770. Pipistrellus only by the absence of the TYPE LOCALITY.-Se-sa-lin, Ichang, Yang-tse, anterior upper premolar, an obsolescent China. tooth which varies in size in Pipistrellus but Consistently larger than the European is always reduced. Moreover, certain murinus. Forearm, 50 mm. species groups of Eptesicus resemble given species groups of Pipistrellus in the form of Vespertilio namiyei (Kuroda) their skulls, so that one asks one's self Nyctalus namiyei KURODA, 1920, Annot. Zool. Japon., IX, Pt. 5, p. 601. whether obsolescence of p2, with conse- TYPE LOCALITY.-Otsukuejima, coast of quent transformation of a pipistrelle into Chikwzen Province, Kiusiu, Japan. an Eptesicus, may not have taken place Writing in 1938,2 Kuroda shows how he more than once. In the present paper, first placed namiyei in Nyctalus, next (1934) however, the conventional assumption will in Eptesicus, and finally in Vespertilio. be upheld, that all species of Eptesicus are Forearm given as 45-48 mm. No descrip- more closely related to one another than to tion of skull (in 1938). "Color of both sides any species of Pipistrellus. of body much deeper [than superans] and Study of -a representative assemblage of without a distinct 'frosted' area on back." the species of Eptesicus has resulted in their Kuroda's reference of namiyei to Ves- provisional segregation into a number of pertilio is accepted provisionally. species groups whose number and scope are shown synoptically: EPTESICUS RAFINESQUE Eptesicus RAFINESQUE, 1820, Annals of Na- SPEcIEs GROUPS OF EURABIAN Eptesicus ture, p. 2. Eptesicus nilssonii group (subgenus Amblyotus Adelonycteris H. ALLEN, 1892, Proc. Acad. Kolenati). Nat. Sci. Philadelphia, for 1891, p. 466. New This is perhaps the least modified group. Old name for Vesperus, preoccupied. World. Numerous forms. The basic concept of the genus Eptesicus Eptesicus nasutus group (subgenus Rhyneptesicus is founded upon Rafinesque's Bianchi.) genotype, Subgeneric characters imperfectly known. E. melanops = Vespertila [sic] fuscus Beau- India. See text beyond. vois and V. mydas, both North American Eptesicus fuscu8 group (subgenus Eptesicus_ bats, and Kaup's Cnephaeus, 1829, with Rafinesque) .4 type Vespertilio serotinus Schreber. The Exceptionally large. Postorbital process de- other two European species admitted to the veloped on strong zygoma; m3 reduc3d. Cir- 3 1912, Cat. Mammals of Western Europe, pp. 1 For full description see Miller, 1912, Cat. Mam- 224-237. mals of Western Europe, pp. 238-242. 4 Pachyomus Gray, as revived by Bianchi, synonym 2 1938, A List of the Japanese Mammals. of Eptesicus subgenus. 272 Bulletin American Museum of Natural History [Vol. LXXX

cumpolar, with tropical representatives. with full braincase, deep zygoma, weak Numerous forms. Eptesicus pachyotis group (subgenus Paraep- basial pits and unreduced m3; minutus,3 a tesicus Bianchi). species nearly as small as pusillus, whose Subgeneric characters not studied. India. skull shows strong general likeness but has Eptesicus demissus group. stronger zygomatic arches and larger ante- Much resembling fuscus group, but skull with basial pits. orbital foramina; and somalicus, which is Eptesicus pumilus group. of the size of pusillus but differs in numer- A group of very small species with reduced ous small details which need not be enu- zygomata and unmodified m3. Australia. merated here. Besides the groups indicated above, In spite of the differences pointed out it others in Africa or South America may later seems very likely that the closest relatives be segregated, e.g., Neoromicia Roberts. of the Australian group of Eptesicus occur It appears that the headquarters of in Africa and not in Eurasia. Eptesicus in regard to numbers of species In South America Eptesicus is not well lies in lower temperate and subtropical represented. Apart from members of the Asia and Africa, while rather meager off- serotinus-fuscus group, the species are small shoots are present in Europe, America and to medium-sized, dark brown in color: Australia. In southern Asia only two andinus, forearm, 45 mm. (size of nilssonii), species, dimissus and verecundus, are present has the braincase high and full, zygoma in the Malay States. Taylor' shows no rec- with slight postorbital process, well defined ord of Eptesicus from the Philippines. basial pits, muzzle narrow, m3 unreduced; None is known from the East India islands the skull of cubensis is almost a replica of west or north of New Guinea and Australia. hingstoni (Persia) but has the postorbital Thus the Australian pusillus group is sepa- processes higher and bullae smaller; in rated from mainland Eptesicus by a dis- chapmani, forearm 40 mm., the zygoma is tance of some 2000 miles. strong and deep, a weak "helmet" appears, Africa is quite rich in species of Eptesicus. muzzle narrow, m3 somewhat shortened in Allen2 shows twenty-six species which be- the toothrow, anteorbital foramen unusu- tween them extend over most of the con- ally large; in doriae, forearm 37 mm., the tinent. characters of chapmani are repeated but the In Africa is found a series of species or size is in every way less, and the anteorbital races conspicuous because of their nearly foramen is not enlarged; hilarii appears vir- translucent flying membranes; also there tually equal to chapmani. are many quite small species which possibly This short survey, although incomplete, find their nearest allies not with northern suggests that no close relatives of thepusillus Eptesicus but with the Australian pumilus group will be found in the neotropical re- group. Possibly some of those, including gion. The South American forms (other the pumilus group, are referable to Neoro- than the allies of serotinus) appear on the micia Roberts (type, zuluensis). whole to be unspecialized species derived Those African species with "white" or from early Eptesicus stock. Hilarii and translucent membranes include phasma, chapmani, in which m3 is slightly shortened, rendalli, faradjius and tenuipes. They are may be distantly allied to serotinus and characterized by completeness of m3 and fuscus. extreme weakness and delicacy of the zygoma, as in the Australian pusillus group, Eptesicus nilssonii Group but they differ from pusillus and allies by (Subgenus Amblyotus) broadening of the supraorbital area and In Amblyotus,4 typified by Vespertilio development ofsmall supraorbital tubercles. nilssonii, the skull is relatively delicate and Of African dark-winged forms I have tapered, the braincase low. A definite seen flavescens, a moderate-sized species 3 Our series, identified as "minutus," may be in reality capensis. 1 1934, Philippine Land Mammals, p. 115. 4 Ognev, 1928, Mammals of Eastern Europe and 2 1939, Bull. Mus. Comp. Zool., LXXXIII, pp. Northern Asia, I, p. 543, treats Amblyotus as a full 83-87. genus. 1942] Tate, Results of the Archbold Expeditions. 47 273

postorbital deepening of the zygoma at the pears in Miller, 1912.1 Pelage dark brown jugal succeeds the anterior, slender zygo- with tips whitish; forearm, 38-40 mm. matic process of the maxilla. The supra- The details of the skull have been described orbital ridges are weak and play no part in under the group heading. Measurements forming the anteorbital foramen. Ros- are shown in the table. trum, palate and zygomata rather narrow. Incisors slender, elongate, il with pos- Eptesicus nilssonii caucasicus (Satunin) terior accessory cusp; i2 slightly shorter Vesperugo (Vesperus) cauca.sicus SATUNIN, than accessory cusp of il; canine slender, 1901, Zool. Anz., XXIV, p. 649. with sharp posterior cutting edge; m3 TYPE LOCALITY.-Tiflis. complete, the two posterior cornmissures By Satunin in 1896 this bat was regarded relative to the anterior one, 0.7, 0.7 :1.0, as a "variety" of nilssonii (= borealis). and length of m3 (in toothrow) 1.0 mm. Forearm, 35 mm., according to Ognev, 1928 Lower incisors delicate, moderately im- (not given by Satunin). Color dark brown, bricated, incisor slightly thickened; P2 washed with yellowish, beneath gray brown, slender, compressed, its blade about one- with silvery tips. third of height of blade of P4. The numerous named forms referable to Eptesicus nilssonii pallescens Bobrinskoj this group are apparently confined to the Eptesicus caucasicus pallescens BOBRINSKOJ, Eurasian continental mass and to the 1926, Comptes Rendus Acad. Sci. U.R.S.S., ser. A, pp. 97-98. northern parts of Africa. TYPE LOCALITY.-River Moldja (Maldj), I have been able to see only a few rep- mountain ridge of Russki. resentatives of the group: nilssonii, walli The description indicates a pallid form of and "jobiensis." The last has forearm caucasicus Satunin. 38-42 mm. and c-m3, 5.6-5.9. Forearm, 35.0-35.5 mm. Of those forms described by various authors but not seen by me caucasicus Eptesicus nilssonii tamerlani Bobrinskoj from Tiflis, pallescens from the River Eptesicus tamerlani BOBRINSKOJ, 1918, Fauna Moldja, tamerlani from east Turkestan, and Flora of Russia, XV, pp. 13-16. and velox from Vladivostok, all with fore- TYPE LOCALITY.-Baisunski Bay, Bucharia, arm 35 mm., seem closest to nilssonii. east Turkestan. Larger bats (forearms near 40 mm.) in- Forearm, 33.5-35 mm. Ognev (1928) clude kashgaricus from near Kashgar, considered tamerlani a race of caucasicus centrasiaticus from Tibet, and gobiensis but somewhat paler in color. from Mongolia. The small tauricus may not belong with Eptesicus nilssonii velox Ognev nilssonii at all but to one of the African Eptesicus velox OGNEV, 1927, J. Mamm., VIII, groups of small-sized Eptesicus. p. 154. Walli, which I have described beyond TYPE LOCALITY.-Vladivostok. (from a supposedly identical specimen) in Forearm, 35 mm.; c-m3, 4.8. company with matschiei and pellucens, may represent a wholly distinct species. Eptesicus nilssonii kashgaricus Bobrin- E. alashanicus from Mongolia was dis- skoj tinguished by Bobrinskoj by special char- Eptesicus nilssonii kashjaricus BOBRINSKOJ, acters of the upper incisors. 1926, Comptes Rendus Acad. Sci. U.R.S.S., ser. A, p. 97. TYPE LOCALITY.-Khotan-Tagh, mountains Eptesicus nilssonii (Keyserling and of Russki, Asiatic Russia (near Kashgar). Blasius) Forearm, 40.5-41.7 mm.; c-m3, 6.0. Vespertilio nilssonii KEYSERLING AND BLAsius, Lighter in color than typical nilssonii- 1839, Archiv. f. Naturg., V, Part 1, p. 315. dorsally straw yellow, with bases of hairs TYPE REGION.-Sweden. dark brown, ventrally buffy white. This bat has been described by many authors. One of the best descriptions ap- 1 Cat. Mammals of Western Europe, p. 234. 274 Bulletin American Museum of Natural History [Vol. LXXX

Eptesicus nilssonii centrasiaticus Bobrin- Forearm, 35 mm.; ". . . uniform sandy skoj buffy above, rather paler below . . . skull Eptesicus nilssonii centrasiaticus BOBRINSKOJ, slender, with small braincase, widely ex- 1926, Comptes Rendus Acad. Sci. U.R.S.S., panded zygomata. Muzzle remarkably ser. A, p. 96. flattened . . . " TYPE REGION-.Tibet. No specimens of matschiei and pellucens Forearm, 40.5 mm.; c-m3, 5.9. are available. They seem from the descriptions most nearly referable to the nilssonii Eptesicus nilssonii gobiensis Bobrinskoj group, but like many other mammals from Eptesicus nilssonii qobiensis BOBRINSKOJ, 1926, that part of the world (the southeastern Comptes Rendus Acad. Sci. U.R.S.S., ser. A, p. 96. arid belt of Asia) they may have closer TYPE REGION.-Mongolia. relatives in Africa. Forearm, 38.5 mm.; c-m3, 5.5 Eptesicus matschiei pellucens (Thomas) Eptesicus tauricus (Ognev) Vespertilio matschiei pellucens THOMAS, 1905, Proc. Zool. Soc. London, p. 520. Amblyotus tauricus OGNEV, 1927, J. Mamm., TYPE LOCALITY.-Ahwaz, Karun River, south- VIII, p. 152. west Persia. TYPE REGION.-Crimea, Russia. This is one of the unusually small mem- This form was described as a race of bers of the nilssonii group, according to Og- matschiei but larger and paler. However, nev. Forearm, 30 mm.; c-im3, 4.5. Ognev its forearm measured only 35.7 mm. against contrasted it with tamerlani. 35.0 of matschiei. Eptesicus bobrinskoi Kusjakin Eptesicus waffi Thomas Eptesicus bobrinskoi KUSJAKIN, 1935, Bull. Eptesicus walli THOMAS, 1919, J. Bombay Soc. Nat. Moscou, XLIV, 7-8, pp. 435, 437. Nat. Hist. Soc., XXVI, p. 746. TYPE LOCALITY.-Brunnen Tjulek in the Aral TYPE LOCALITY. Basra, Mesopotamia. Sea, Kaza Kum desert (65 km. east of C;ty "...Allied to E. pellucens and matschiei. Aralskoje More), Central Asia. Size rather larger . . . [forearm, 40 mm., Forearm, 35; c-m3, 5.4 mm. Fur long, against 35.7 and 35.0] . .. c-im3, 5.3." dark brownish black, with tips olive yel- In a specimen from Nasiriyah, Iraq, low; "underparts whitish." Kusjakin F.M.N.H. 43767, the foregoing dimensions states that this species is nearest alashani- are exactly duplicated. This skull, com- cus, differing "by greater size and more pared with topotypical nilssonii, shows sev- flattened skull, by the smaller outer and eral outstanding differences which must upper incisor .... . be considered tt set walli off as a full species; the zygoma is weak and without any Eptesicus alashanicus Bobrinskoj sign of postorbital process, much as in Eptesicus alashanicus BOBRINSKOJ, 1926, Comptes Rendus Acad. Sci. U.R.S.S., ser. A, p. the Australian pumilus group. The bullae 98. are so large that they cover the cochleae; TYPE REGION.-Mongolia. no basial pits are developed (they are well 'Upper outer incisor said to equal upper developed in nilssonii); the internal cingu- inner incisor in extent. This indicates a lum of the canine and the regions of proto- peculiar form, if so, since in all groups of cone and hypocone are more heavily de- Eptesicus seen i2 is markedly shorter than veloped than in nilssonii. The length in the iP. Forearm, 37 mm. toothrow of m3 is considerably greater (0.95: 0.8). In the lower jaw P2 iS slightly Eptesicus nasutus Group (Subgenus smaller and slightly moved outward from Rhyneptesicus) the dental alignment. This group apparently contains only matschiei (Thomas) nasutus of the Himalayas. Bianchi (1916) Eptesicus with Vespertilio matschiei THOMAS, 1905, Ann. erected Rhyneptesicus type species Mag. Nat. Hist., (7) XVI, p. 573. nasutus Dobson. No subgeneric descrip- TYPE LOCALITY.-Aden, Arabia. tion was offered. 1942] Tate, Results of the Archbold Expeditions. 47 275

Eptesicus nasutus (Dobson) missure is much shorter than the middle Vesperugo (Vesperus) nasutus DOBSON, 1877, commissure, which in turn is very much J. Asiatic Soc. Bengal, XVI, p. 311. shorter than the anterior one; their propor- TYPE LOCALITY.-Sinde, western India. tions (in serotinus) are 0.4 : 0.7: 1.2 mm. "'Upper inner incisors moderately long The body of the tooth in the toothrow is and unicuspidate. Outer incisors very correspondingly shortened. In serotinus it short ... forearm, 1.45 inches (37-38 mm.) measures 0.7-0.8 mm. One primitive char- . . .no postcalcareal lobes; wings from acter retained by members of this group is base of toes ....." the well defined postorbital process of the The "unicuspidate" il, if substantiated, zygoma. would suggest relationship to Scoteinus The lower incisors are strongly imbri- and allies. cated, i2 and i3 each with a small low pos- A specimen, M.C.Z. 5147, from Amballa, terior cusp set directly behind the middle in the Siwalik Hills, is of moderate size cusp of the three which form the cutting (the forearm, 37 mm.), the skull without blade of the tooth. "helmet," rostrum broad, supraorbital A number of forms of the large, conspic- ridges prominent, and the large rostral de- uous bats of this group have received pressions anterior to the orbits and one names. Mainly temperate and subtropical either side of the midline of the rostrum in latitudinal range, the group is circum- emphasized. Zygoma with anterior por- polar in distribution. In America it has tion deep, no sign of postorbital process. also representatives in the tropics. It is Paroccipital processes proportionately short possible that the entire group represents (much shorter than in the fuscus group). only one species. The skulls of fuscus of Incisors and canines greatly worn; m3 com- the eastern UJnited States and serotinus of plete, the posterior, middle and anterior Europe, as stated earlier, greatly resemble commissures measuring 0.8, 0.7, 1.0 mm., each other. respectively; the length of m3 in the tooth- In the Old World members of the sero- row, 0.9-1.0 mm. tinus group (or fuscus group?) there ap- Lower incisors with i2 and i3 slightly and pear to be two lots of named forms distin- progressively thickened; cusp of P2 one- guishable by no morphological differences third of height of cusp of p4. other than size. The larger ones are primarily northern, reaching from Europe Eptesicus fuscus Group (Subgenus through to India, China and Japan. The Eptesicus) smaller ones are southern and extend from Although this group includes the type Roumania to Tripoli, Arabia, Persia and species of the genus, fuscus,l it is in reality southern Russia. rather specialized. Besides comprising species of unusually large size, it is char- "LARGE" SECTION Form Locality Forearm in mm., acterized by its broad, flat skull and strong serotinus Western Europe 49-53 development of an occipital "helmet," turcomanus Asia Minor 47-54 (Ognev) formed by confluence of lambdoid and mirza Persia sagittal crests, by rather pronounced de- shirazensis Shiraz, Persia 54 intermedius Russia velopment of the supraorbital ridges, and pachyomus India 54 by moderate depressions on the rostrum at andersoni Yunnan 53 either side of the midline anterior to the sinensis Peking 49 orbits. The teeth are large and strong; pallens Kansu 49 il has its posterior cusps almost as well de- brachydigitus Korea 47.5 veloped as the main cusps; the main cusp "SMALLER" SECTION of i2 is about one-half as deep as the poste- sodalis Roumania 45-48 rior cusp of il; in m3 the posterior com- ognevi Bucharia 45-46 (Ogne v) isabellinus Tripoli 42 1 The American members of the group treated by G. M. Allen, 1933, J. Mamm., XIV, p. 150; and by bottae Botta, Arabia 40 Engels, 1936, Amer. Midl. Nat., XVII, pp. 653-660. hingstoni Bagdad 45 276 Bulletin American Museum of Natural History [Vol. LXXX Thomas suggested that turcomanus and Eptesicus serotinus mirza (Filippi) mirza may be synonymous. I can offer Vesperus mirza FILIPPI, 1865, Note di un no evidence on the matter. Pachyomus of Viaggio in Persia nel 1862, p. 342. India and andersoni of Yunnan and Szech- I have not seen this work but Filippi wan have skulls which are indistinguish- is stated in the, Zoological Record (II1 able, though the tips of the hairs in the Mamm., p. 3) to have visited Tiflis and former (U.S.N.M. 37792 and 37306 from worked along the south shore of the Caspian Kashmir) are more strongly marked with Sea. Thomas (1919) has suggested that whitish than in our good series of andersoni. mirza is probably synonymous with turco- Pallens from Kansu appears to be equal or manus. very closely related to sinensis from Pe- king. Possibly brachydigitus of Korea be- Eptesicus serotinus shirazensis (Dobson) longs here too. Vesperugo shirazensis DOBSON, 1871, J. Asiatic Of the "smaller" bats isabellinus and bot- Soc. Bengal, XL, p. 459. tae should be compared for possible iden- TYPE LoCALITY.-Shiraz, east of Persian Gulf. tity. Hingstoni is the smallest of the spe- Another large form (forearm, 54 mm.), cies available to me for examination. perhaps also close to turcomanus and mirza The American species (or races) fuscus, but equally likely to belong with pachyo- bernardinus, miradorensis and wetmorei mus of northwest India. are members of the serotinus group, possibly only subspecies. None of the "small" Eptesicus serotinus pachyomus (Tomes) members is represented in America. Vespertilio pachyomus TOMES, 1857, Proc. Zool. Soc. London, p. 50. Eptesicus serotinus Section TYPE REGION.-India. Gray (1866) made of this bat the genus Eptesicus serotinus (Schreber) Pachyomus, recognized recently (1916) Vespertilio serotinus SCHREBER, 1774, Sauge- as a subgenus by Bianchi. Dobson (1878) thiere, I, P1. III; description, op. cit., I, p. 167, 1775. referred pachyomus to the synonymy of TYPE REGION.-France. serotinus. I am unable to distinguish Miller (1912) gives the forearm as 49- generic or even specific characters to sep- 53 mm., c-mi3, 7.6-8.2. The species has arate pachyomus from serotinus. The been described frequently. bat is larger and heavier than true serotinus of western Europe and should be considered Eptesicus serotinus intermedius Ognev a geographical race. It is closely allied Eptesicus serotinus intermedius OGNEV, 1927, to shirazensis and andersoni. J. Mamm., VIII, p. 152. Forearm, 55 mm. TYPE LOCALITY.-Terek region, near Vladi- kavkaz, Murtasovo Sta., Russia. Eptesicus serotinus andersoni (Dobson) Intermediate (in color) between serotinus Vesperugo andersoni DOBSON, 1871, Proc. and turcomanus. Asiatic Soc. Bengal, p. 211. TYPE REGION.-Yunnan. Eptesicus serotinus turcomanus (Evers- Forearm, 53 mm. A rather dark-colored mann) serotine with pale tips to hairs, related to Vespertilio turcomanus EVERSMANN, 1840, but darker than pachyomus. Bull. Soc. Imp. Nat. Moscou, I, p. 21. TYPE REGION. . Between Caspian and Eptesicus serotinus sinensis (Peters) Aral Seas." Vesperus sinensis PETERS, 1880, Monatsber. " ... Ashy ferruginous ... whitish be- Akad. Wiss. Berlin, p. 258. TYPE LOCALITY.-Peking, China. neath . . . front and sides of muzzle nearly bare ... teeth quite large ... ears exceeding Forearm, 49 mm. More nearly the size half length of head, nearly as broad as of true serotinus; smaller than the southern long .... " Length of forearm not given races shirazensis, pachyomus and ander- by Eversmann, but Ognev (1927) gives it soni. Peters gave an illustration of the as 47-54.2 mm. ear of this race. 19421 Tate, Results of the Archbold Expeditions. 47 277 Eptesicus serotinus pallens Miller Eptesicus demissus Group Eptesicus serotinus pallens MILLER, 191 1, Proc. Eptesicus demissus is distinguished by Biol. Soc. Washington, XXIV, p. 53. its moderate size and its "helmet" and TYPE REGIoN.-Kansu, China. broad zygomata like those of the fuscus Miller did not compare pallens with group. The zygomata are broken in the sinensis. The forearm lengths are identi- type. The basicranial region is conspicu- cal. But the type region is 500 miles west ous by having a pair of deep basial pits be- of Peking, so some racial differences may tween the bullae. (In fuscus and serotinus be discernible. the pits, though present, are very weak.) In addition, the photograph of the type Eptesicus serotinus brachydigitus Mori skull of demissus shows the roof of the Eptesicus trachydigitus MORI, 1928, Zool. Soc. posterior narial canal and anterior part of Japan, p. 291. the mesopterygoid fossa to be raised high TYPE REGION.-Korea. in the skull. M3 is also complete, its "Allied to serotinus but smaller . . . length (in the toothrow) being almost forearm, 47.5, c-m3, 7.3 mm." 1 mm. Eptesicus sodalis Section Eptesicus demissus Thomas Eptesicus demissus THOMAS, 1916, J. Fed. Eptesicus sodalis (Barrett-Hamilton) Malay States Mus., VII, p. 1. Vespertilio sodaiss BARRETT-HAMILTON, 1910, TYPE REGION.-Malaya. Ann. Mag. Nat. Hist., (8) V, p. 291. " . . . Related to E. pachyotis [see under TYPE REGION.-Roumania. incertae sedis] . . . color chestnut brown A small species very like serotinus; fore- above, lighter below . . . skull with well arm, 45-48 mm.; c-m3, 7.2 (Miller, 1912). marked occipital helmet . . . forearm, 42, c-mi3, 6.2." Eptesicus sodalis ognevi Bobrinskoj Zygomata very wide; basial pits very Eptesicus ognevi BOBRINSKOJ, 1918, Fauna and deep. Flora of Russia, XV, p. 12. I have not seen Bobrinskoj's paper. Eptesicus pachyotis Group (Subgenus Ognev (1927) made ognevi a race of sodalis, Paraeptesicus Bianchi) giving forearm length as 45-46.6 mm.; Seemingly only Vesperugo pachyotis c-im3, 6.4-6.7; " . . . color very pale, the Dobson is referable to this group. The only back sandy-gray, sometimes with a chest- descriptive matter related to the thickened nut tint; below white." basal part of the ear pinna posteriorly, a character of perhaps only specific value. Eptesicus bottae (Peters) However, Chasen compared verecundus Vesperus bottae PETERS, 1869, Monatsber. with "pachyotis" and "demisssus." Akad. Wiss. Berlin, p. 406. TYPE LOCALITY.-Botta, Arabia. Eptesicus pachyotis (Dobson) Forearm, 40 mm., too short to be related Vesperugo (Vesperus) pachyotis DOBSON, 1871, -to serotinus as indicated by Trouessart. Proc. Asiatic Soc. Bengal, p. 213. More probably either near sodalis and og- TYPE LoCALITY.-Khasia Hills, Assam. -nevi or else derived from an African group. Forearm, "1.6 inches" (38-39 mm.). Color dark brown, a lighter shade beneath. Eptesicus hingstoni Thomas Readily distinguished by the Eptesicus hingstoni THOMAS, 1919, J. Bombay peculiar thickness of the lower half of the Nat. Hist. Soc., XXVI, p. 745. outer side of the ear conch ... minute upper TYPE LoCALITY.-Bagdad. premolar ... absent.... " Forearm, 45 mm.; " . . . intermediate Most of the additional general char- -in size between the small E. innesi of Egypt acters given by Dobson apply to any spe- .and E. s. turcomanus of Persia ....." cies of Eptesicus. The type is supposedly 278 Bulletin American Museum of Natural History [Vol. LXXX in the Indian Museum, Calcutta, and A paratype, M.C.Z. 29113, is dark brown, should be examined carefully for further above and below, with blackish hair bases. characters. C-m3, 4.9 mm.; forearm, 34. This makes darlingtoni the largest of the weakly dif- Eptesicus pumilus Group' ferentiated races of pumilus. The Australian forms of Eptesicus are typified by E. pumilus. Compared with Eptesicus pumilus vulturnus Thomas E. Eptesicus pumilus vulturnus THOMAS, 1914, the conservatively built (Amblyotus) Ann. Mag. Nat. Hist., (8) XIII, p. 440. nilssonii, the braincase is unflattened, the TYPE REGION.-Tasmania. rostrum shorter and palate wider, but the The photograph of the type skull, B.M. zygomata have been reduced to slender 7.1.1.375, shows an animal decidedly larger shallow rods with scarcely a sign of post- than either pumilus or caurinus. Forearm, orbital enlargement. In il a well formed 33 mm.; c-m3, 4.6. posterior cusp can be observed, but i2 is much reduced, its depth less than half of Eptesicus pygmaeus (Becker) that of the accessory cusp of iP. Lower Vespertilio pygmaeus BECKER, 1858, Trans. teeth much as in E. (Amblyotus) but height Phil. Inst. Victoria, III, pp. 38-40, P1. of crown of P2 about one-half of height of TYPE LOCALITY.-Oakleight, Danegong crown of P4* ranges, 30 miles north of Melbourne, Victoria. These tiny bats include several races and Becker's plate shows unmistakably the are the only representatives of Eptesicus in skin, skull and dentition of an Eptesicus of Australia. the pumilus group. The form may be merely a synonym of pumilus from New Eptesicus pumilus (Gray) South Wales or it may equal vulturnus of Scotophilus pumilus GRAY, 1841, in Grey, Tasmania. Becker did not give the length Journals Two Exped. Australia, II, pp. 400, 406. of the forearm. TYPE LOCALITY.-Yarrundi, New South Wales. I have photographs and notes of the type Eptesicus, incertae sedis skull, B.M. 41.1523 (119c). Back of skull Besides the three species first treated, of type broken. Forearm, 31 mm.; c-m3, p7atyrhinus Dobson, atratus Blyth and 4.2. verecundus Chasen, there are a number of names which have been proposed by Eptesicus pumilus caurinus Thomas Japanese authors: aurijuncus Mori, hori- Eptesicus pumilus caurinus THOMAS, 1914, kawai Kishida, kobayashii Mori, parvus Ann. Mag. Nat. Hist., (8) XIII, p. 439. TYPE REGION.-Kimberley, Western Aus- Kishida, renanensis Kishida and Mori tralia. (nomen nudum) concerning which I have Here also we have a photograph of the little or no information valuable for the type skull, B.M. 14.3.9.1 (10473). purpose of referring the forms to species This western form is considerably smaller groups. than pumilus, with forearm, 30 mm.; Eptesicus platyrhinus (Dobson) c-m3, 3.9. Vesperugo platyrhinus DOBSON, 1875, Ann. Mag. Nat. Hist., (4) XVI, p. 262. Eptesicus pumilus darlingtoni G. M. TYPE LOCALITY.-Unknown. Allen Marked by "absence of the first minute Eptesicus darlingtoni G. M. ALLEN, 1933, J. upper premolar." Apparently an Eptesi- Mamm., XIV, p. 150. CUS. TYPE REGION.-Queensland. 1 Vespadellus Iredale and Troughton, 1934, Mem. [Eptesicus atratus (Blyth)] Australian Mus., VI, p. 95, was proposed without de- atratus Cat. Mamm. scription to include the Australian species of Eptesi- Nycticejus BLYTH, 1863, cus. According to the "International Congress, Mus. Asiatic Soc. Bengal, No. 96. Budapest," p. 1589, it is invalid. Troughton em- TYPE LOCALITY.-Darjeling, India. ployed the name again in his recent book "Furred Animals of Australia," still without generic descrip- Forearm, 1.7 inches (43-44 mm.). tion. I am not aware that a description has been published elsewhere. This name is a homonym of Amblyotus 19421 Tate, Results of the Archbold Expeditions. 47 279 atratus Kolenati, 1858, a synonym of E. are well imbricated, with their outer lobes nilssonii. Dobson placed atratus Blyth reduced as in Rhogulssa; P2 iS in the tooth- between discolor Kuhl (true Vespertilio, be- row but much compressed and reduced. cause a synonym of V. murinus) and pachy- The ears appear much as Eptesicus, and otis. the pelage is a pale straw color with dark bases. Eptesicus verecundus Chasen Allen lists three species, two from Eptesicus verecundus CHASEN, 1940, Bull. Anglo-Egyptian Sudan, one from South Raffles Mus., XV, p. 53. Africa. TYPE LOCALITY.-Perak, Malaya. " . . . General characters of E. pachyotis LAEPHOTIS THOMAS Dobson (not studied) and E. demissus Laephotis THOM.AS, 1901, Ann. Mag. Nat. Thomas (photograph of skull of type Hist., (7) VII, p. 460. studied) but smaller than both . . . inner GENOTYPE.-Laephotis wintoni Thomas. incisor much larger [than in pachyotis] This genus is in many respects a smaller and tricuspid. TUpper canine with small copy of Histiotus. It is less specialized. secondary cusp ... forearm, 34.5 ... c-m3, The ears and bullae, although larger than 4.8mm." those of Eptesicus, do not attain the ex- Verecundus obviously has peculiarities treme size seen in Histiotus. In the teeth not seen elsewhere in Eptesicus. The type il is bifid as in both Eptesicus and Histio- was kept at Singapore. tus; m3 is unreduced. The dorsal profile The following references apply to Japa- of the skull is high as in Histiotus. The nese names concerning which sufficient data palate is very strongly domed. are not available: Laephotis is known only from Africa. E. horikawai Kishida, 1924, Zool. Mag., Tokyo, XXXVI, p. 127; Dobuts. Zasshi, XXXVI, HISTIOTus GERVAIS No. 425. Formosa. E. kobayashii Mori, 1928, Zool. Soc. Japan, p. Histiotus GERVAIS, 1855, Exped. Comte de 292; Annot. Zool. Japon., XI, 4, p. 392 (Eng- Castelnau, Zool. Mamm., p. 77. lish). Korea. GENOTYPE.-Plecotus velatus I. Geoffroy. E. parvus Kishida, 1932, Lansania, IV, No. 31, p. 2. North Korea. Derived from the line leading to Eptesicus E. aurijuncus Mori, 1928, Annot. Zool. Japon., and showing the same dental formula, but XI, 4, p. 393. Korea. with i2 even more reduced. The last molar " . . . Ears firmly joined by dermal ridge is unreduced. between their inner margins on forehead ... In the skull the rostrum and temporal forearm, 49; c-m3, 6.8." This description regions are rather high, combining to form seems inapplicable to Eptesicus. a gently arching profile leading to the braincase. RHINOPTERUS MILLER The outstanding specialization of Histio- Rhinopterus MILLER, 1906, Proc. Biol. Soc. tus occurs in the ears. Both external and Washington, XIX, p. 85. skeletal parts are greatly enlarged. The Scabrifer G. M. ALLEN, 1908, Bull. Mus. very large, forward projecting pinnae are Comp. Zo6l., LII, p. 46. united by a lower frontal band. The GENoTruPsl-G1auconycteris floweri DeWinton. width of each bulla is more than twice The rostrum is rather flat and broad; the their distance apart. braincase shows no increase in depth such Further distinctive characters of Histio- as appears in Glauconycteris and Chalinolo- tus are the large size of the anteorbital fo- bus; the zygoma is moderately deep but ap- ramina and of the eyes. parently without postorbital processes. Histiotus is a purely American offshoot In the dentition, il is bifid, the secondary of the Eptesicus stem and is found chiefly cusp lateral from the primary one; i2 is in the warmer latitudes. I have taken it in quite small; p2 is absent as in Eptesicus Ecuador at 8000 feet. Several species have (and Glauconycteris). The lower incisors been named. 280 Bulletin American Museum of Natural History [Vol. LXXX Nycticeini All general of this division, named are Nycticeius in America and Scoteinus after Nycticeius, possess the common char- in Asia. A small American offshoot cul- acter that their upper incisor toothrows minates in Rhoge6ssa and Baeodon. In are strongly modified from the bicuspid Asia and Africa considerable specialization condition of Myotis, Pipistrellus and has resulted in the production of Scotophilus Eptesicus. The outer incisor is obsolete; and allies, and Otonycteris. The nycticeine the inner one usually lacks the supple- genera are shown in relation to each other mentary cusp and is simply conical. There diagrammatically (Fig. 5).

Scotophilus Scotoecus Scoteinus Nycticeius ScotomanesIl Skull relat:ively Skull broadened; unbroadened, teeth massive I .A-1 4 -111., \\/

Rhogelssa Baeodon

i3 reduced ij3 unreduced

Skull of Scotophilus Ears not characters; ears enlarged | greatly enlarged

Nycticeini Incisors and premolars reduced in number

Fig. 5. Suggested phylogeny of the nyeticeine genera. See also Fig. 1. is a strong resemblance between the incisor SCOTEINUS DOBSON teeth of Australian Scoteinus and those Scoteinus DoBsoN, 1875, Proc. Zool. Soc. of the nyctophiline bats of that country. London, p. 371; 1876, Monogr. Asiatic Chirop- P- are invariably absent; m3 often much tera, pp. 120, 123-124; 1878, Cat. Chiropt. Brit. Mus., pp. 257, 262-264. reduced. GENOTYPE.-Nycticejus emarginatus Dobson Less specialized members of this division (designated by Miller, 1907; Palmer, 1904, 1 Except Scoteinus pallidus. The status of Sco- indicated no previous selection of the type teinus, which may be diphyletic, needs clarifying. species). 1942] Tate, Results of the Archbold Expeditions. 47 281

In this genus, as in Hesperoptenus, we lian species are more specialized than is the may be dealing with a polyphyletic group. Indian pallidus. Whether emarginatus, the Unfortunately there appears to be no genotype, conforms to this situation may Indian material in our collections which is be shown eventually, when the skull of the undisputably referable to S. emarginatus, type specimen has been studied. the type. One skull from India, A.M.N.H. 54420, labeled S. pallidus, has m3 unreduced Scoteinus emarginatus (Dobson) and so does not come within Miller's Nycticejus emarginatus DOBSON, 1871, Proc. (1907) definition. But that definition, it Asiatic Soc. Bengal, p. 211. was Scotophilus emarginatus DOBsON, 1875, Proc. will be noted, based upon the Austra- Zool. Soc. London, p. 371; 1876, Monogr. lian balstoni and greyii and the African Asiatic Chiroptera, pp. 123-124; 1878, Cat. schlieffenii. Actually, until we have a good Chiropt. Brit. Mus., p. 262. description of the skull of the genotype TYPE LOCALITY.-"India, precise locality emarginatus we cannot correctly under- unknown. Type in the collection of the Indian stand Scoteinus. Museum, Calcutta." Provisionally Scoteinus may be con- Dobson accompanied his first descrip- sidered a less specialized genus coming from tion of emarginatus with the words "sub- the stem which also produced the more genus Nycticejus: premolars, 1_2 wing specialized Scotophilus. Indication of af- membranes attached to base of toes." His finities between the two genera is shown description of N. emarginatus was based by the character of the upper incisors. otherwise upon external characters. The Of the Australian species one, rdppellii, is hair "above, tricolored, at the base dark strongly divergent. It is a large species (as ferruginous brown, then buff, the tips large as the Indian emarginatus) with light yellowish brown; beneath, dark fer- forearm, 52-53 mm. It offers only one ruginous brown at the base, the remaining special character, the unusual length of the portion buff . . . forearm, 2.2 inches (54 postpalatal spine. The remaining Austra- mm.)." lian species fall into three groups: larger In his monograph (1876) he added a line species with broadened mastoid processes drawing of the head of emarginatus. His and prominent supraorbital ridges, in- remarks in the "Catalogue" (1878) added fluatus and orion; smaller forms with mas- nothing new. toids less prominent and the supraorbital Actually the skull of emarginatus remains eminences virtually obsolete, so that the undescribed. We must accept Dobson's margin of the orbital fossa passes back indirect statement that p2 is absent. smoothly from the lacrimal region to the Nevertheless it is desirable that the type intertemporal constriction, balstoni, aquilo, be restudied so that the characters of the caprenus; and a considerably smaller series type skull may be clearly defined. from Papua and north Queensland, sanborni and greyii. Scoteinus riuppellii (Peters) All of these Scoteinws, including rfppeliii, Nycticejus riuppellii PETERS, 1866, Monatsber. are distinguished from the Indian pallidus Akad. Wiss. Berlin, p. 21. TYPE LOCALITY.-Sydney, New South Wales by the fact that in them m3 is more re- (by Peters stated "Sydney, West Australia"). duced, though far from reaching the extreme condition seen in Scotophilus. The Color dark rust red, beneath lengths of the two posterior commissures of paler .... Forearm, 52 mm....." This the Z-pattern (in orion) compared to that large Australian species is wholly unlike of the anterior one is approximately 0.6, any other species of that continent. Fore- 0.7: 1.1; whereas in paUlidus m3 is scarcely arm, 52 mm. reduced at all, and the commissures measure 0.8, 0.8: 1.0. The length of the body Scoteinus pallidus (Dobson) Scotophilus pallidus DOBSON, 1876, Monogr. of the tooth (m3) in the toothrow is 0.7 Asiatic Chiroptera, Appendix D, p. 186. mm. in orion and in pallidus 1.0. It be- TYPE LOCALITY.-Mian Mir, near Lahore, comes clear that, in this respect, all Austra- Punjab, northwest India. 282 Bulletin American Museum of Natural History [Vol. LXXX

" .. . Teeth as in S(cotophilus) temminckii perhaps the eastern representative of S. [i.e., with i2 and p2 absent] . . . . Fur, balstoni.... . integuments of the body, wings and interfemoral membrane pale buff through- Scoteinus orion Troughton out. . . Forearm, 1.4 inches (35-36 Scoteinus orion TROUGHTON, 1937, Austral. mm.)." Zool., VIII, 4, p. 277. Specimens from Kooloo Valley, India, TYPE LoCALITY.-Sydney, New South Wales. exist in American museums (M.C.Z. "Allied to balstoni . . . forearm, 34.5- 5337, 5341; A.M.N.H. 54419-20) which 36.3 . . . ear definitely smaller than in agree well with the description of pallidus. balstoni." Forearm, 34-36 mm.; rostrum broad, supraorbital tubercles small; zygomata weak; anteorbital foramina not enlarged; Scoteinus balstoni Thomas sagittal crest undeveloped (back of brain- Scoteinus balstoni THOMAS, 1906, Proc. Zool. Soc. London, p. 472. case broken away); dentition of type TYPF LOCALITY.-Hawknest, Laverton, south- of Scotophilus, i2 and p2 obsolete, but differ- west Australia. ing in (1) unreduced pattern of m2-, (2) lower incisors unimbricated and almost "Allied to greyii, hair bicolor instead of equally thin, and (3) P2, though the height unicolor ... pale brown ... Below, hairs of its crown is only one-third of that of dark at base . .. tips pale pinkish buff ... p4, is completely in the toothrow, un- upper incisors standing rather farther from crowded, and shows the crown outline canines (than in greyii) . . . forearm, 36 very slightly compressed laterally. mm. (4 others 34-35) ... zyg. br., 10.1; c- Before we can feel sure of the identifica- m3 5.2.. tion the type skull (in Indian Museum) must be examined. Scoteinus orion aquilo Troughton Obviously, from its much smaller size, Scoteinus orion aquilo TROUGHTON, 1937, pallidus must be entirely different from Austral. Zool., VIII, 4, p. 278. TYPE REGIoN.-East coast of Queensland, emarginatus. Bowen to Rockhampton. Scoteinus noctulinus (I. Geoffroy) Compared with orion, "size generally smaller . . . skull of similar outline but Vespertilio noctulinus I. GEOFFROY, in Bel- smaller proportions . . . cheek-teeth rows anger, 1834, Voyage aux Indes-Orientales . . . Zoologie, p. 92, P1. III. decidedly shorter . . . forearm (holotype), TYPE REGION.-Bengal. 34.5; zygomata breadth, 10.4 . . . c-m3, " . . . Head and body above reddish fawn, 5.1 . . . the longer ear, larger dimensions beneath clear fawn . .. interfemoral mem- and more robust skull leave no doubt of its brane partly hairy . . . forearm, 1 inch, distinction from greyii. . . . 4 lines (34 mm.) ... muzzle naked. ..." It appears probable that noctulinus is Scoteinus balstoni caprenus Troughton related to pallidus. It is far too small to Scoteinus balstoni caprenus TROUGHTON, 1937, equal Scotophilus temminckii, in whose Austral. Zool., VIII, 4, p. 279. synonymy it has usually been placed. TYPE LoCALITY.-Roebuck Bay, Kimberley region, northwest Australia. Scoteinus influatus Thomas "Size smaller generally than southern ... Scoteinus influatus THOMAS, 1924, Ann. Mag. balstoni ... fur not strongly bicolored Nat. Hist., (9) XIII, p. 540. forearm, 32.5-34 mm... color ... appar- TYPE LOCALITY.-Prairie, central south ently a dull drabby or more olivaceous Queensland. brown (than balstoni) . . . lower incisors " . . . More nearly approaching ... bal- more crowded, distorted, and less distinctly stoni ... forearm, 39 mm.; condylo-canine trilobate than in true balstoni or greyii; length, 15; zygomatic breadth, 11.5; mas- first lower premolar smaller and more toid breadth, 9.4; maxillary toothrow, 6 ... rounded. ... 1942] Tate, Results of the Archbold Expeditions. 47 283

Scoteinus greyii (Gray) all trace of the posterior sagittal crest of Scotophilus greyii GRAY, 1843, List Mamm. Australian Scoteinus. Brit. Mus., p. 30; 1844, Voyage Erebus and Nycticeius is represented by the single Terror, P1. xx. TYPE LoCALITY.-Port Essington, Northern American species humeralis. Territory, Au sralia. RHOGEESSA H. ALLEN "... . Back and beneath brighter chest- Rhogeessa H. ALLEN, 1866, Proc. Acad. Nat. nut..."(1843). Sci. Philadelphia, p. 285. The forearm length of the female de- GENOTPYPE.-RhoJeS&sa tumida H. Allen. picted on P1. xx (1844) is 34 mm. How- "Like Nycticeius, but outer lobe of all ever, no statement is made that the picture three lower incisors obsolete and inner lobe is of natural size. I have not seen the text practically absent also in outer tooth.... " which this plate accompanies. Known only from tropical America. Thomas (1906) designated specimen "b" Rhogeessa and its derivative Baeodon of P1. xx type of S. greyii. constitute a primitive, isolated American The skull of the type, B.M. 42.8.17.12, offshoot of the nycticeine bats of Asia. is badly smashed, but its toothrows are Contrary to the usual trend in the Ves- intact and measurable (c-m3, 5.2 mm.). pertilioninae, i3 in this genus and the next becomes progressively smaller. Scoteinus sanborni Troughton Scoteinus sanborni TROUGHTON, 1937, Austral. BAEODON MILLER Zool., VIII, p. 280. Baeodon MILLER, 1906, Proc. Biol. Soc. TYPE LOCALITY.-East Cape, Papua. Washington, XIX, p. 85. Compared chiefly with greyii, "ear de- GENOTYPE.-Rhoge8ssa alleni Thomas. cidedly smaller and forearm and pes Specialized offshoot of the line leading longer ... forearm, 32-33.3 mm....." to Rhogeessa; " . . . reduction of the outer lower incisor carried so far that the tooth NYCTICEIUS RAFINESQUE has become a mere functionless spicule Nycticeius RAFINESQUE, 1819, Journ. de Physique, LXXXVIII, p. 417. less than one-twentieth as large as the first GENoTYPE.-Vespertilio humeralis Rafinesque. or second incisor, nearly concealed beneath The American Nycticeius was the first cingulum of canine...." of the vespertilionid group of genera char- R. alleni was taken first in Mexico. acterized by simplified first upper incisors to be recognized. Immediately afterward SCOTOECuS THOMAS came Scotophilus of the oriental tropics. Scotoecus THOMAS, 1901, Ann. Mag. Nat. It was long usual for writers describing Hist., (7) VII, p. 263. bats of this group in India and adjoining GENOTYPE.-Scotophilus albofuscus Thomas. areas to refer them to Rafinesque's genus. Exclusively African in distribution. In- Nycticeius, though its relationship to termediate between Nycticeius and Sco- Scotophilus and allies is demonstrated by teinus on the one hand and Scotophilus and the loss of i2 and by reduction of il by loss Scotomanes on the other. Zygomatic and of the posterior cusp to a simple, peg-like rostral widths exaggerated. Anteorbital tooth, is far less specialized. The skull foramen enlarged as in Scotophilus. Teeth shows none of the extreme broadening of much as in Nyctophilus. the rostrum and lacrimal region or the en- G. M. Allen listed eight species. largement of the supraorbital tubercles usually found in the oriental representatives SCOTOPHILUS LEACH of the group of species. The zygoma is Scotophilus LEACH, 1821, Trans. Linn. Soc. London, XIII, pp. 69, 71-72. weak. Pachyotus GRAY, 1831, Zool. Miscellany, I, Compared with Australian Scoteinus less p. 38 (part); 1838, Mag. Zool. Bot., II, p. 498. dissimilarity may be noted than in the case GENOTYPE.-S. kuhlii Leach (for both generic of Scotophilus. The skull of Nycticeius is names). See remarks on status of kuhlii. unbroadened across the rostrum and lacks Name Scotophilus not preoccupied by 284 Bulletin American Museum of Natural History [Vol. LXXX

Scotophila Hftbner.I Nycticeius, now re- tion to the shortened face, present in stricted to American bats, was used com- Pipistrellus-like bats, it has developed the monly for Asiatic species by authors during posterior part of the sagittal crest, together the 19th century. Scotophilus has been with the lambdoid crests, to form a distinct restricted since Dobson wrote in 1878. At helmet such as one sees in Taphozous; the that time it included as subgenera, Scoto- zygomata are weak; the anteorbital philus, Scoteinus and Scotomanes. foramen on the face is unusually large- Pachyotus Gray included Nycticeius larger than any other Vespertilionid known Rafinesque, 1819, with type N. humeralis to me except Scotoecus. from the United States, and Scotophilus The dentition is massive and crowded: Leach, 1821, with type S. kuhlii. Gray il unicuspid; i2 and p2 obsolete; m3 greatly (1831) wrote, "The bats, the Vespertiliones modified by shortening in the toothrow of Geoffroy, might for convenience be and comprise only protocone, paracone and divided into three genera, the true bats, parastyle. Vespertilio, with thin ears and membranes Lower incisors thickened, much imbri- and a hairy face, the Pachyotus with thick cated, i1-2 trilobate, the inner and middle ears and membranes and bald swollen lobes higher than the outer, i3 as i2 but with cheeks, including the genera Nycticejus small anterior and posterior cuspules added and Scotophilus, and the hairy-tailed species which rise only. slightly above the cingu- of America (Lasiurus)." And again (1838) lum; lower canine with large cingulum heel under Scotophilus, "upper surface of inter- extending backward, tending to embrace femoral membrane slightly covered with the inner wall of P2 and touch p4; P2 com- hair (Pachyotus)," followed by two species pressed between c and p4; talonid of m3 Vespertilio polythrix I. Geoffroy and Scoto- much reduced in conformity with condi- philus levis I. Geoffroy from Brazil.2 The tion of m3. descriptions of these bats form part of a paper on the Vespertilionidae of Brazil. Status of Scotophilus kuhlii Leach Polythrix was apparently a species of Scotophilus kuhlii LEACH, 1822, Trans. Linn. Dasypterus; levis perhaps a Myotis. Soc. London, XIII, p. 72. Dobson, when writing his monograph,3 TYPE LOCALITY.-Unknown. divided Scotophilus into three subgenera: Peters and Dobson have explained that Scotophilus, Scoteinus and Scotomanes. All the incisive formula - given by Leach have since been given generic standing. In related to milk dentition in the extremely the restricted Scotophilus Dobson included young type specimen. Presumably the only temminckii (Java) and borbonicus molar (plus premolar) formula , instead (Africa). of k and the canine peculiarities are ex- The next year4 he dealt with temminckii plainable in the same way. and "subspecies heathii," showing wood- The only specific character (and one cuts of the heads of both. Two years without value in systematic work on this later5 he treated temmiinckii, with variety species) given by Leach was the color, heathii, borbonicus (Africa) and gigas "ferruginous, ears, nose and wings fuscous." (Africa). Under temminckii he grouped a Dobson, in the monograph cited, claimed number of alleged synonyms: kuhlii, that the specific identity of the animal belangeri, castaneus, luteus, flaveolus and which Leach named Scotophilus kuhlii could fulvus. not be determined. Peters had written Scotophilus is specialized in regard to earlier6 that he regarded the t-ype as equal certain characters of the skull. In addi- to temminckii Horsfield, that it retained the milk incisives, and its forearm meas- 1 G. M. Allen, 1939, Bull. Mus. Comp. ured only 41 mm. I have seen the types LXXXIII, p. 99. Zool., 2 1824, Annales des Sciences Naturelles, (1) III, of neither. pp. 440-447. In more recent times, and in spite of 3 1875, "On the Genus Scotophilus ...," Proc. Zool. Soc. London, pp. 368-373. Peters' expressed opinion, authors have 41876, Monogr. Asiatic Chiroptera, pp. 119-125. 5 1878, Cat. Chiropt. Brit. Mus., pp. 256-266. 6 1866, Monatsber. Akad. Wiss. Berlin, p. 679. 1942] Tate, Results of the Archbold Expeditions. 47 285 come to apply kuldii to the larger forms The races and synonyms are presented in which the forearin exceeds 54 mm. I can from west to east and north to south, be- find no more justification for this than I ginning at India and ending at Philippines. can for Peters' conclusion, and until some- one has measured the size ot the teeth in Scotophilus temminckii wroughtoni the juvenal type amd checked the results Thomas against teeth of known "large" or "small" Scotophilus wroughtoni THOMAS, 1897, J. Scotophilus, it will be best to pass by kuhlii Bombay Nat. Hist. Soc., XI, p. 275. and use the names heathii and temminckii. TYPB LOCALITY.-Kim, Surat District, north In the list following the heathii and of Bombay, western India. temminckii groups are separated. It will "Fur . . . uniform brownish above, be seen that the smaller forms appear to without any tinge of yellowish, the hairs have a slightly difmerent total range from white at their bases and gradually dark- the larger. A bat of the temminckii group ening to brownish tips . . under surface occurs on Formosa. very pale fawn, almost white . . . forearm, 50mm....." LIST OF NAMED FoRMS OF ASIATIC Scotophilus This bat was compared with "kuhlii" and Form Locality Forearm emarginatus (a Scoteinus). temminckii The photograph of the type skull shows a wroughtoni Peninsular India 50 slightly narrower zygomatic expanse than gairdneri Central Siam 48 that of true temminckii. Pelage paler consobrinus Hai:nan 52 brown dorsally, and ventrally buffy white. castaneus Mal 50 temminckii Wee;tJavaJacca 51 All over peninsular India and common on = fulVus Ceylon where Phillips2 records it from the solutatus East Java 54-55 drier parts of the island. Its pallid tones collinus Bali 48-54 give it the appearance of an arid country panayensms Phil 48 bat. heathii ~ippines heathii South peninsular 64-69 India Scotophilus temminckii gairdneri Kloss belangeri South peninsular 54 India Scotophilus gairdneri KLOSs, 1917, J. Nat. luteus Coromandel- 59 Hist. Soc., Siam, II, p. 284. Bengal TYPE LOCALITY.-Paknampo, central Siam. flaveolus Continental India 54-56.5 "About size of S. castaneus and wrought- insularis Hainan, South 64-67 oni ... rostrum narrower than in S. casta- China celebensis Celebes neus ... above bister, the head darker.... 621/2-64 Below pale drab, the base of the fur whitish. . . . In its pale underparts this bat Scotophilus temminckii Horsfieldi bears some resemblance to S. wroughtoni ... (References under subspecies.) but it has the upper pelage much A term used here to include a number darker...... of named races and color forms of "small" This a small species with forearm only Scotophilus with forearm 54 mm. or less. 48 mm. I have seen no undoubted speci- Anatomically almost identical to heathii mens. (which by Dobson was considered a subspecies of temminckii) but with the occipital "helmet" less developed. Anterior lower Scotophilus temminckii consobrinus premolar invariably compressed longitu- (Allen) Scotophilus castaneus consobrinus J. A. ALLEN, dinally. 1906, Bull. Amer. Mus. Nat. Hist., XXII, p. 485. Distribution from Bombay to Formosa, TYPE LOCALITY.-Hainan Island, South China. Philippine Islands, Java and Bali. Seem- Of this form we have the type and numer- ingly absent from Celebes. ous paratypes. I find myself unable to 1 See Sody, 1928, Natuurk. Tijdschr. Ned. Ind., distinguish between these bats and the XC, pp. 271-272, in which he reviewed Pachyotus temminckii. 2 1935, Manual of Mammals of Ceylon, p. 124. 286 Bulletin American MlIsecuml of Naturol Jhistory [VLX,1. I ,XXX large series we now have of true temnminckii [Scotophilus fulvus Gray I from Java. This similarity supports to Scotophilus foilvcos GRAY, 1843, List Mainiiii. some extent my idea that intervening Col. Brit. Mus., p. 31. forms, gairdneri, castaneus, panayensis, are TYPE LOCALITY. Java. merely weak variant races or even indi- Gray's terse description serves little ex- vidual variants from a single widely dis- cept to validate the name fulvus: persed species. "The Foxy Noctule. Scotophilus fulvus: a. Yellowish brown beneath. Java. Scotophilus temminckii castaneus b. Yellowish beneath. Java.... (Horsfield) c. India, Madras." Nycticejris castaneus HORSFIELD, 1851, Cat. Since descriptive matter is provided only Mamm. Mus. East Iindia Cornp., p. 38. for Javanese specimens, the type locality TYPE REGION. "Singapore, Penianig, Malay may be restricted to Java. Fulvus then Penini,tula and Islands." becomes a probable synonym of temminckii. Characteristic feature . . . a uniform deep chestnut color of the body, above and beneath . . . head blackish. Scotophilus temminckii collinus Sody Size of temminckii." Scotophilus temminckii collinus SODY, 1936, A cotype of castaneus, a female, labeled Natuurk. Tijdschr. Ned. Ind., XCVI, p. 48. "Malacca," B.M. 79.11.21.11b, has the TYPE LOCALITY. Bali. forearm 50.5 mm. in length. "Balinese (and East Javanese) ... aver- In this case too I am constrained to the age slightly smaller than the typical race belief that castaneus of the Malacca region [temminckii]...... is merely a rufescent phase of temminckii. Sody gave: forearm, 48-54 mm.; greatest Bonhote, 1 comparing castaneus of Malaya length of skull, 18.4-19.3; upper toothrow with "kuhlii," gave the "average length of (+ c), 6.5-6.9. He compared collinus from forearm . . . 50.7 (49-52)." These meas- east Java and Bali with temminckii from urements correspond exactly with those of west Java, giving for the latter the meas- temminckii given by Sody in 1936 and measurements: forearm, 50-56 mm.; greatest ured on our series of specimens from Cheri- length of skull, 18.0-19.7; upper toothrow, bon, Java. 6.6-7.2. In the Archbold Collections is a series of Scotophilus temminckii temminckii fourteen specimens from Bali. I cannot (Horsfield) agree with Sody that they average smaller Vespertilio temminckii HORSFIELD, 1824, Zool. than our material from Cheribon, Java. Researlches in Java. TYPE LocALITY.-Java. Color "pure dark brown above, grayish Scotophilus temminckii solutatus Sody brown, somewhat dusky beneath, with a Scotophil Os castaneos solotat os SODY, 1 936, rufous tint exten(ling laterally from the Natuurk. Tijdschi. Nod. Iimd., XCVI, p. 49. nose to the tail." TYPE LOC.\T,ITY.-Tjalldiroto, cast Java. No significant measurements were of- In this race Sody claimed superior size fered by Horsfield. over castaneus of Malacca and true tern- Temminck2 recognized under "Nyctice- mninckii. He slhowed (with rather few speci- jus temrinckii" five varieties in more than mens) that whereas size gradient ran up- 100 specimens taken from "Java, Sumatra, ward in temminckii from west to east, it Borneo, Banda and Timor." ran downward from west to east in solu- In the Archbold Collections is a series of tatus. On this somewhat tenuous basis he twenty-three specimens of temminckii, all claimed specific rank for castaneuts. from Cheribon, Java. Our forearm meas- In two strongly rufescent specilnens, urements extend from 47-51 mm. M.C.Z. 12909 and 12919, fromn Pelahoean Ratoe, Java, the forearm lengths are 55 1 1900, PIroc. Zool. Soc. London, pp. 191-192. 2 1835. M\onogr. Marnnm., II, p. 149). mm. 1942 ] Tate, Results of the Archbold Expeditions. 47 287

Scotophilus temminckii panayensis Sody sented in Sumatra, Java, Borneo and Scotophilus panayensis SODY, 1928, Natuurk. Philippines. 'Iijdschr. Ned. Ind., LXXXVIII, p. 90. TYPE LOCALITY. Panay Island, Philippines. Scotophilus heathii belangeri Forearm only 48 mm. Virtually no de- (I. Geoffroy) scription offered. I have no Philippine Ve3pertilio belangeri I. GEOFFROY, in Belaniger, specimens. Taylor referred such material 1834, Voyage aux Inde--Orienitales ... Zoologie, pp. 87-92. to temminckii, but he seems not to have TYPE LOCALITY. Found commonly ill the come across Sody's name panayensis, which towins near Pondichslrry, Coromandel Coast, was applied to four animals which the latter India. had never seen, said by Elliot to have "Body fawn or white, yellowish beneath, forearms only 48 mm. in length. Those maroon olive or brown above.... Length animals were females. The low limit for of forearm, 2 inches (54-55 mm.)." This length of forearm in Javanese temrninckii, form appears to be annectant in size be- according to Sody,1 is 50 mm. tween heathii and temminckii. The de- Miss Lawrence2 mentioned specimens scription almost fits that of wroughtoni. of larger size than the type of panayensis. The type skull is probably lost. She gave the forearm length as 48 to 50 mm. The unusual shortness of the caudal [Scotophilus luteus (Blyth)] vertebrae, mentioned by her, may prove Nycticejus luteus BLYTH, 1851, J. Asiatic Soc. to be a useful eharacter. Bengal, XX, p. 157; 1852, J. Asiatic Soc. Bengal, XXI, p. 345. TYPE REGION. "Beigal: Coromandel" (eaAt- Scotophilus heathii (Horsfield) em-n peniinsular India). Nycticejuts heathii HORSFIELD, 1831, Proc. "Length (of a large male) 55/8 incl1es... Zool. Soc. London, p. 113. tail . . . 2'/ inches . . . forearm, 2'/4 in. TYPE LOC.ALITY. Madras, Inidia. (57 mm.) ... entire length of skull is barely Considerably larger than the 1 in., inclusive of the greatly developed Javanese species [temminckii], from which sagittal crest" (1851, p. 157). it differs also remarkably in its coloring ... Underparts of luteus being of a covered with short . . . silky hair . . . much more rufescent hue than those of . .. color . . above is brown with a tawny hue; ternminckii. The length of forearm in underneath fulvous with a slight tendency temminckii is very regularly 2 inches (51 to gray ...... mm.), in luteus 23/8 inches (58 mm.), in Larger bats (forearm, 64-69 mm.) of the heathii 23/4 inches (69 mm.)" (1852, p. Indian peninsula, generally colored yellow- 346). ish brown, slightly paler beneath. In the Luteus appears to be a synonym of skulls I can find no difference from the belangeri I. Geoffroy. skulls of temminckii, other than those caused by their greater size. In both [Scotophilus flaveolus (Horsfield)] species P2 is compressed longitudinally in Nycticejus flaveolhts HORSFIELD, 1851, Cat. Mamm. Mus. East Inidia Coirnp., p. 37. the lower tioothrow. Specimens from Cey- TYPE REGION. "Many parts of Continental lon appear to be consistently darker colored India." than those from the mainland. Phillips Horsfield listed five specimens: A, (1935) states of the Ceylonese race "color "bright rufous beneath"; B, yellowish exceedingly variable, commonly dark yel- gray beneath; C, pale. D aind E were not lowish bronze-brown above . . . another described. "This species represents the variety ... deep chestnut .... Between former [temminckii of Java] on the con- these extremes all variations of color are tinent of India, being fully one-third to be found." larger. The color varies considerably in This large species is apparently unrepre- different individuals, being dark brown 1936, Natuurk. Tijdschr. Ned. Ind., XCVI, p. 48. above, in different shades, and rufous or 2 1939, Bull. Mlus. Cornp. Zool., T,XXXVi, pp. 18-59. yellowish underneath." 288 Bulletin American Museum of Natural History [Vol. LXXX

We have photographs of the skulls of two than those from South India." The rostral of Horsfield's cotypes, B.M. 60.5.4.31-32. sinus in insularis is deeper than the sinus Both are marked "India." In both, P2 iS in Ceylonese specimens of heathii. strongly compressed longitudinally in the Forearm, 60-62 mm. toothrow. The forearm length of two cotypes, B.M. 60.5.4.30 and 32, is 56.5 and Scotophilus heathii celebensis Sody 54 mm., respectively. Scotophilus celebensis SODY, 1928, Natuurk. The type locality of flaveolus should be Tijdschr. Ned. Ind., LXXXVIII, p. 90. restricted. Provisionally I treat it as syn- TYPF LoCALITY.-Toli-toli, Celebe3. onymous with belangeri and luteus. As in Forearms of three specimens, 621/2-64 belangeri, the relative shortness of the fore- mm. Very weak description. arm suggests the temminckii group, but the I have included celebensis with the heathii large size of the teeth (see table) indicates division chiefly because of its great size heathii. compared with temminckii of nearby

DETAILED MEASUREMENTS OF DENTITION IN Scotophilus, TENDING TO SHOW DISTINCTION BETWEEN heathii (= kuhlii?) GROUP AND temminckii GROUP Cingulum Distance length apart of Crown di- lower lower Width, Width, mensions, Forearm C-m3 canine canines m4 m3 p2 A.M.N.H. 83449 Kashmir 58 8.0 1.0 1.5 2.5 2.5 0.9 X 1.0 A.M.N.H. 44544 Salween River 61 7.8 1.2 2.3 2.6 0.6 X 1.2 A.M.N.H. 54796 Burma (red) 7.6 1.3 1.7 2.3 2.5 0.7 X 1.0 26787 Hainan (insularis) 61 8.4 1.5 1.5 2.7 2.8 0.7 X 1.4 26784 Hainan (insularis) 66 8.7 1.6 1.9 2.9 2.9 0.8 X 1.3 M.C.Z. 27519 Ceylon (heathii) 63 8.0 1.4 2.4 2.6 0.7 X 1.2 M.C.Z. 27518 Ceylon (heathii) 63 8.0 1.4 1.9 2.5 2.7 0.6 X 1.2 M.C.Z. 12909 Java (solutatus) (red) 55 7.1 1.1 1.4 2.2 2.2 0.5 X 1.0 A.M.N.H. 101869 Java (temminckii) (brown) 48 6.5 1.2 1.3 2.1 2.1 0.6 X 0.9 A.M.N.H. 107452 Bali (collinus) (brown) 49 6.9 1.3 1.5 2.2 2.2 0.7 X 1.0 IJ.S.N.M. 144817 Luzon (panayensis) (brown) 50 6.5 1.2 1.6 2.3 2.1 0.5 X 0.9 U.S.N.M. 144819 Luzon (panayensis) (brown) 50 6.6 1.2 1.7 2.1 2.1 0.5 X 0.8 U.S.N.M. 199596 Hainan (consobrinus) (brown) 53 6.7 1.3 1.7 2.2 2.1 0.5 X 0.9 F.M.N.H. 45998 Formosa (brown) 6.5 1.2 1.8 2.1 2.1 0.6 X 1.0 Scotophilus heathii insularis Allen islands. There exists, however, a geo- Scotophilus kuhlii insularis J. A. ALLEN, 1906, graphical hiatus since, according to Bon- Bull. Amer. Mus. Nat. Hist., XXII, p. 485. hote and Chasen,1 "kuhlii" (meaning TYPE LOCALITY.-Hainan Island, South China. heathii) has not been recorded south of "Color above uniform olive brown; below Patani, Malay Peninsula. pale brownish buff....." These large bats are apparently the SCOTOMANES DOBSON northeastern representatives of the Indian Scotomanes (subgenus of Scotophilus) DoBsoN, heathii. They are essentially similar to 1875, Proc. Zool. Soc. London, p. 37. the continental form rather than to the GENOTYPE.-Nycticejus ornatu, Blyth. Ceylonese race which latter, as Blyth re- This is a monotypic genus with one marked (1852), "are a good deal darker 1 1940, Bull. Raffles Mus., XV, p. 54. 19421 Tate, Results of the Archbold Expeditions. 47 289 species, ornatus, and two named forms, contrasted with rest of under-surface .... imbrensis and sinensis, both weak races Forearm of type 56 mm....." which perhaps cannot be sustained. - The type locality of this "race" is only a Its relationships seem to be with the same few miles from Cherrapunji, type locality line which gave rise to Scotophilus. The of true ornatus. skin is less specialized as regards the ear, in which the tragus is of simple Pipistrellus Scotomanes ornatus sinensis Thomas form instead of being elongate pointed and Scotomanes ornatus sinensis THOMAS, 1921, J. broadened in the middle as in Scotophilus. Bombay Nat. Hist. Soc., XXVII, p. 772. The ornate white pattern may be con- TYPE LOCALITY.-Kuatun, northwest Fukien, sidered as specialization. China. In the skull, shortening of the rostrum "Size smaller, forearm 50-55, rarely has gone farther than in Scotophilus, similar reaching 60. Color deeper . . . forearm of broadening of the supraorbital area has type 55 mm....." occurred, but widening across the anterior It is very doubtful whether the "smaller" margins of the orbits in the region of the size of this Chinese race can be sustained. anteorbital foramen (circular instead of A.M.N.H. 84847 from Fukien has the oval) has taken place. The rostral sinus forearm 59 mm. A second specimen meas- is relatively small. The palatal sinus, ures 54 and a third 56 mm. too, is little developed. The posterior part of the skull remains low, and the sagittal and lambdoid crests weak. Mastoid OTONYCTERIS PETERS processes comparatively unexpanded. Otonycteris PETERS, 1859, Monatsber. Akad. 223. Dentition much as in Scotophilus but m3 Wiss. Berlin, p. slightly less reduced, lower incisors simply GENOTYPE.-Otonycteris hemprichii Peters. trilobate as in Pipistrellus. P2 proportion- In this genus we see the third independ- ately as small, but less compressed between ent evolution of "big-eared-ness" in the c and P4* Vespertilioninae. The other cases were Plecotus and allies, derived from the Myotis stem, in which little reduction of the Scotomanes ornatus (Blyth) dentition is evident, and Histiotus (with Nycticejus ornatus BLYTH, 1851, J. Asiatic Laephotis), an offshoot of the Eptesicus Soc. Bengal, XX, p. 517. line, showing sp3cialization of the premolar TYPE LOCALITY.-Cherrapunji, Burmese India. area but none of the incisors. Otonycteris " demonstrates enlargement of the auditory . . . Bright pale rusty isabelline-brown above (the piles black for the basal fourth, apparatus in the nycticeine bats, where then whitish, with rusty extremities) less already marked modification of the upper vivid on the lower half of back, and incisors is an established fact. somewhat paler below; a pure silky white Divergence probably took place at or spot on the center of forehead, others on before the Nycticeius-Scoteinus stage, be- each shoulder and axilla above, and a nar- cause marked broadening of the rostrum is row stripe of the same along the middle of not evident, and the, occipital helmet is the back ... forearm 21/2 inches [57 mm.]." merely incipient. Moreover, the zygomata are still fairly strongly developed. Otonycteris inhabits the arid regions of Scotomanes ornatus imbrensis Thomas north and east Africa, Arabia and adjoin- Scotomanes ornatus imbrensis THOMAS, 1921, ing territories. J. Bombay Nat. Hist. Soc., XXVII, p. 772. In Asiatic regions the names petersi TYPE LOCALITY.-Konshnong, Jaintia Hills, (Persian Gulf), jin (Hasa, Arabia), cinereus Assam. (Baluchistan) and cheesemani (Arabia) "Size ... as in ornatus or rather smaller. have been applied. I am not in a position Color distinctly darker and browner ... to attempt to evaluate these names. Prob- dark collar on throat blackish, strongly ably some, at least, are synonyms or races, 290 Bulletin American Museum of Natural History [Vol. LXXX Lasiurini American vespertilionid bats, specialized il, obsolete or vestigial state of p2, great to an extraordinary degree. The elevated reduction of m3 (longitudinally). In the braincase provides a superficial likeness lower incisors iP is the major functioning to Chalinolobus of Australia and Africa. tooth, j2 and il being smaller, a condition The densely pilose uropatagia resemble wholly different from Myotis. Imbrication those of the Murininae of southeastern occurs. Asia. Two genera (which perhaps should be subgenera) only are distinguished: LASIURUS GRAY Lasiurus and Dasypterus. Within Lasiurus Lasiurus GRAY, 1831, Zool. Miscellany, I, two chief divisions appear, the cinereus p. 38. group, large bats with greatly elongated GENOTYPE.-Vespertilio borealis Mutller (Pal- basal pollical phalanx, under surface of wing mer, 1904). adjoining forearm densely pilose, etc., and "The hairy-tailed species of America"2 the borealis group, smaller bats with basal' (Gray). Only two species, cinereus and phalanx of thumb less elongated and wing borealis, are listed by Miller for North and near forearm thinly haired beneath. This ,Central America. Several others related latter group extends to South America as to borealis occur in South America. far south as Chile, where it is represented by a somewhat larger form (varius). DASYPTERUS PETERS Dasypterus, although it is distinguished Dasypterus PETERS, 1871, Monatsber. Akad. from Lasiurus by the absence of p2, re- Wiss. Berlin, p. 912. sembles L. cinerea otherwise in many GENOTYPE.-Lasiurus intermedius H. Allen. respects. Miller lists three species of Dasypterus The bats of this supergeneric group to- north of the Isthmus of Panama. The gether form a very distinct offshoot of the genus is represented in tropical Soutb Vespertilionidae. They are set apart by America. Dasypterus differs from Lasiurus the broad, very short, elevated rostrum obviously by the loss of p2. This distinc- and braincase, which is illustrated by the tion, although perfectly definite, is less proximity of the orbital fossa to the canine, important than may be suspected, because by the great breadth of the anterior rostral p2 in Lasiurus is a disappearing structure and palatal sinuses, reduction of incisors to already greatly reduced in size.

TABLES OF MEASUREMENTS OF VESPERTILIONINE BATS The tables which follow are made up for cus pumilus group, both measurements are the most part of the measurements of type shown. In each case an apparent increase specimens and cotypes. A great many of in the size of skull varying from 2 per cent the sets of measurements were made on to 10 per cent results from the use of photog- actual specimens in European museums raphy. This means that in photograpbs during the summer of 1937, but a large made with my Contax camera and en- number also have been taken from photo- larged with a commercial enlarger, tooth- graphs of the types with which a millimeter row lengths of 5 mm. usually require a cor- scale had been included. Slight lenticular rective of -0.4 mm. In a few instances aberration has introduced errors in these measurements are quoted from publica- latter groups of measurements. In certain tions. Forearm lengths are generally taken instances, Pipistrellus abramus, P. banca- from actual type specimens. A few are nus, P. regulus, P. tasmaniensis, Nyctalus copied from text or transposed into milli- molossus and some members of the Eptesi- meters from inches. sluatuxais'equa SoSSooo0 0 0 0 0 0 0 0 0 0 0 0 0 jo aoinoS - 0) 0) 0)

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SYNOPTIC LIST OF VESPERTILIONINAE (EXCEPT MYOTIS) IN THE ARCHBOLD COLLECTIONS SKIN AND ALCO- SPECIES OR SUBSPECIES LOCALITY SKULL HOLIC Pipistrellus imbricatus South Celebes 15 " I Bali 6 imbricatus collinus Balim River, Netherlands New Guinea 2 Idenburg River, Netherlands New Guinea 1 Mt. Tafa, Papua 1 Lake Habbema, Netherlands New Guinea 1 abramus bancanus Cheribon, Java 12 papuanus Sogeri, Papua 13 Kemp Welch River, Papua 1 Hollandia, Netherlands New Guinea 1 Idenburg River, Netherlands New Guinea 10 5 Wassi Kussa, western Papua 7 Daru, western Papua 2 4 Dogwa, western Papua 25 Mabaduane, western Papua 1 Madiri, Fly River, western Papua 2 tenuis Bratan, Bali 1 macrotis Koeta, Bali 3 minahassae Roeroeken, north Celebes 1 Glischropus tylopus Southwest Borneo 19 I "4 I Northwest Borneo 18 "4 di South Borneo 14 Philetor rohui Idenburg River, Netherlands New Guinea 1 66 46 Papua 4 Tylonycteris pachypus Palembang, Sumatra 16 $I 64 Bali 4 robustula Peleng, east Celebes 15 "" Palembang, Sumatra 6 Eptesicus pumilus Quamby, Queensland 23 "4 di Pentland, Queensland 7 Scoteinus influatus Wassi Kussa, Papua 6 it "4 Mabaduane, Papua 1 is di Madiri, Papua 1 di "6 Dogwa, Papua 12 Daru, Papua 1 greyji Pentland, Queensland 4 It "Malbon, Queensland 1 Scotophilus temminckii temminckii Cheribon, Java 22 619 id collinus Bali 15

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