Catalogue of the Chiroptera in the Collection of the British Museum

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Catalogue of the Chiroptera in the Collection of the British Museum

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Catalogue of the Chiroptera in the Collection of the British Museum
Andersen, Knud
Dobson, George Edward
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Britisb museum (Ratiiral l)lstorp). This IS No. 25. of 25 copies of the " Catalogue of Chiroptera " {Second Edition), Vol. 1, printed on special paper.




'A 1 1 y.i'xy^s'Ms: M, pr. iK;-i-n r, y tayloe .and francis, ntD LION COur.T, FLEET SI BELT.


PREFACE. The Cntalogjiie oi Chiroptem in the British Museum by Dr. G. E. Dobson, published b}the Trustees in 1878, proved to be, as was expected, a work of great utility to zoologists, the systematic arrangemeut of these animals having bceu previously in a state of the utmost confusion. The Catalogue was therefore generally welcomed, and as a consequence soon ran out of print and became almost unobtainaMe. Iforeover, the stimulus it gave to the study of Bats rapidly tended to render it obsolete, and a new edition has therefore long been a desideratum. But owing to the difficulties of the subject and the necessity that the writer should be able to give his whole time to the work, undistracted by official duties, it has not previously been possible to arrange for tlie publication of a second edition. Two or three years ago, however, the Trustees were fortunately able to secure the services of Dr. Ivnud Andersen, who had already made a special study of Bats, and who has been able to devote himself uninterruptedly to the preparation of the present woik. This Kdition is in reality a completely new and (irigiiial Monograjdi of the Order Cbirnptera, ior materiids. metliods of work, and ideas on si>ecies have all so radically ch:ingcd since 1878, that notliing remains of the first edition — good as that was for its date— but the title. So great is the increase in the general knowledge of the subject and also in the material examined and described, that whereas in a2


1878 one volume was fouud sufficient for the whole of the Order, rather less than one fifth of it being devoted to the Megachiroptera, a volume of somewhat greater size is now needed for the Megachiroptera only, while a corresponding enlargement will probably take place in the part dealing with the other groups of the Order. The Author indicates in his Preface that the number of specimens belonging to the Museum has greatly increased since 1878, while in addition he has been able to examine all the chief collections of Chiroptera in Europe, notably those of Paris, Berlin, and Leyden, and to borrow a large part of that of the U.S. National Museum at Washington. The thanks of the Trustees are due to the Authorities of the various Museums who have helped him in this respect. The volume also' owes much to Mr. Oldfield Thomas, F.R.S., who has interested himself in its progress since its commencement, and has assisted the Author by discussing difficulties from time to time as they arose, and in other wars. British Museum (Natural History). February 22nd, 191--'. SIDNEY F. HAIIMER, Keeper of Zoology. J.?.'ii,"ilii[. '..L» rr! -,ri;,7'> ;!: ii(>iii!;»ii^\ rt'ii,niv\ J, ii! > :•! ., ly.f.l,,, u\\ (,, .„:}. .f/-,,/f,' \p' T>in() 'ill J Vi litfMTjU •ii.t (*iit luii -jii-h


1»fO If ft-.. AUTHOR'S PEEFACE. SpECiMhw.s EXAMiXED. — III 1843, wheii J. E. Gray published his ' List of the Specimens of Mammalia in the British Museum,' the Megachiroptera were represented in the National Collection by 69 specimens. Thirty-five years later (Dobson's ' Catalogue of Chiroptera,' 1878) the number had increased to 425. After another period of thirty-four years (this Catalogue) the total reaches 1470. The skins enumerated in this volume amount to 956, the specimens preserved in alcohol to 444, and the skulls to 1228. Exactly half a century ago (Gerrard's 'Catalogue of the Bones of Mammalia in the British Museum,' 1862) the number of skulls was only 39 ; in 1878 it was 82. That the number of specimens is now three and a half times, but that of skulls fifteen times, greater than in 1878 is due to the fact that during the preparation of this Catalogue the skulls have been extracted from nearly all the old skins in which they had hitherto been left as well as from about 60 per cent, of the alcoholic specimens. In Dobson's time there were separate skulls of only half the number of species of Megachiroptera then in the Collection, whereas now every species and subspecies in the Museum, with one exception, {Pteropus arncnsis), is represented by at least one and often by a series of skulls. Besides the specimens preserved in the British Museum I have had for inspection a large number from other Collections, and during two visits (in 1907 and 1909) to the Museums of Leyden, Berlin, and Paris I had the privilege of going through nearly the whole of the series of Megachirojjtera in those Collections, so that the total number of specimens examined for the purpose of this Catalogue amounts to about 2400.


AUTUOE S PREFACE. Number of genera, species, ami subspecies. — The growth of our knowledge of the existing forms of Megachiroptera, since Linnean times, is shown in the subjoined table : — Genera. 1758. Linno t . 1810. E.Geoffroyt 2 . 1825. Temiuinck{ 2 . 1837. TemminckJ 5 . 1867. Peters 1 9 . 1870. Gray* 14 . 1878, Dobson 10 . 1899. Matsohie + 20 1907. Millerl 30 1912. This Catalogue 35 . Genera and Subgenera. 11 15 13 35 38 Species. Forms. 1 12 , 20 37 49 . 57 . . 70 . 120 55 71 180 142 228 Of the 228 forms described in this Catalogue 207 (91 per cent.) tire represented in the Collection, of the 35 genera 32 (91 per cent.). The following twenty-one forms are at this moment desiderata (the throe missing genera, all mouotypic, are marked with an asterisk) : — rteropus /ii/j)omelanus geminontm, p. 1C6. „ pallidus, p. 136. „ pwmilus, p. 757. ,, ualanus, p. 177. „ mariantius, p. 178. „ va/iil-ore/isi^, p. 184. "" „ aldahmisis, p. 213. i; '„ tubcrculatus, p. 309. _ J ,, vampj/rua plidon, p. 353. ,, macrotU, p. 396. Aexrodon lucifer, p. 432. * Plcroies andiiota, p. 486. Epomoplorus pousarguesi, p. 543. Myoiigoteris {I'hygetis) brachyceph(da, p. 582. Cynoplerus bracliyotis miuutus, p. 625. ,, vuij&r, p. 029. ,, lyrinceps, p. (i33. * Pieuochincsjagori, p. 645. * Chironax melanocephalus, p. 660. Syconycteris anstralis, p. 781. Noiopieris neocaledonioa, p. 799. All forms, including those not represented iu the Collection, have been examined by me. t All Chiroptera were iu the tenth edition of the ' Systeiua Natura; ' referred to one genus, Vefperfilw. \ E. Geofiroy, ' Description des Roussettes et des Cephalotes, deux nouveaux genres de la faiuille des Chauve-souris ' ; Ann. Mus. d'Hist. Nat. Paris, xv. pp. 86-108 (1810). Teraniinck, ' Monographies de Mainmalogie,' i. pp. 166-204 (1825). Temminck, vp. cit. ii. pp. 49-112 (1837). Peters, " Ueber die Flederlmnde, Ptcrop't, und insbesondere iiber die Arten der Gattung Pterrpus s. s." ; MB. Ak. Bedin, 18G7, pp. 319-333, 86.>-872. Gray, 'Catalogue of the Monkeys, Lemurs, and Fruit-eating Bats in the Collection of the British Museum' (1870). Matschie, ' Die Megachiroptera des Berliner Museums fiir Naturkunde ' (1899). Miller, " The Families and Genera of Bats " ; U.S. National Museum, Bulletin 57 (1907).


authdks teeface. Types of valid names (liuti/pes).— The types of the 228 species and subspecies of Megachiroptera are distributed as follows: — Kidolon (3 forms) 1 H(),I^<;ttus (U) I 9 lioiK'ia (\) ' Pteiopus (103) I 49 Acerodoii (9) I 5 Pteralopex (2) j 2 Styloctcnium (1) 1 Dobsonia (18) j 8 Harpi-ionycteris (1) ... 1 Plerotes (1) I Epoiuops (4) 1 Hvpsigiiatlms (1) I Epoinopliorus (9) | 3 Mici'opteropus (1) Nanonyctoris (1) Scotouycteris (1) C'jwiuycteris (1) .. Myouycteris (4) ... Cyhopterus (16) ... Ptciiochinis (1) ... Me;ja;rops (1) I Dyacopterus (!) ... I Halionycteris (1) ... ! Chironax (1) j Tliooptevus (1) I I'oiithetor (1) j Splia;rias (1) I Xyctimeiic (13) ... j Koiiycteris (3) } Mcgaloplossus (1) Macioglossus (6)... ' Syconycteris (7) ... Mi'loiiycteiis (1) ... ] Nesonycteris (1) ... Notopteris (2) := t3 ^ 13 15 10 Not traced, 1. Calcutta, 2 ; Li,-,boii, 1. Caloitta, 1; Cambridge (Mass.), 1; Copenhagen, 1 ; Genoa, 3 ; Sydney, 1 ; ? Zi-ka-wei, 1 ; private possession, 1 ; not traced, 5. Chicago, 1 ; Dorpat, 1. Brisbane, 1 ; ? Zi-ka-wei, 1. Lisbon, liisbon, 1. Philadelphia. Philadelphia, Stuttgart, 1. Not traced. 1 ; Stockhohti, 1 Lisbon, 1. Calcutta, 1 ; Vienna, 1; not traced, 1. Genoa. Not traced, 1. Calcutta, 1. Hamburg. Not traced, 1. yummed up, this gives the following totals: — British MuseuiQt 118, or more than half the number of all cutypcs; Berlin Museum, 25 ; Leyden, 22; Paris, 17; U.S. Natioual, 15 ; Calcutta, 5; Lisbou, 4; Genoa, 3; two each in the Philadelphia and Zi-ka-wei (?) Museums; one each in the Brisbane, Cambridge (^Massachusetts), Chicago, Copenhagen, Dorpat, Hamburg, Stockholm, Stuttgart, Sydney, and Vienna Museums ; one {Ptcropas lijtleri) is said to be. in private possession ; while ten have not been traced (moat of


AUTHOR S I'KEFACE. these, if not all, are almost certainly not in existence), viz. Eiclohn heJvxim (once in the Mnseum Leverianum), Pteropus hypomelanus tomesi (once in the possession of Mr. L. L. Dillwyn), Pteroims suhniger (Cabinet lleaumur), Pteropvs rufus (Paris Museum), Pteropus nir/er (ancient Royal Cabinet, Paris), Pteropus polioceplialus (based on specimens in Leyden, Paris, and London, none of which can now be traced with certainty), Micropiteropus p>usillus (cotypes once in the Tomes Collection and Paris Museum), Cijnopterus sphinx (ancient " Naturhistorie-Selskab," Copenhagen), Nyciimene cephalot€s (described by Pallas), and Macror/lossus minimus (Paris Museum). — That the sum total (234) is a little in excess of the number of recognized forms (228) is due to the fact that in a few cases the cotypes of one form are preserved in different collections. The 228 forms have been described by the following fifty-seven writers (those, seventeen in number, who have described eutypes preserved in the I3ritish Museum are marked with an asterisk) : — Allen (G. M.), 1. Allen (H.), 1. * Andersen, 57. * Anderson & de Winton, 1. ' Blyth, 1. * Bocage, 'i. Briinnich, 1. Desniarest, 2. De Vis, 1. * Dobson, 13. Elliot, 1. Eschscboltz, 1. Geoffroy (E.), 7. * Gould, 1. * Gray, 16. Hallowell, 1. Heude, 2. Heuglin, 1. * Hodgson, 1. Hombron & Jacquijiot, 1. Jentink, 3. Kelaart, 1. Kerr, 3. * Lay, 1. * Linne, 1. * MacGillivray, 1. Mason, 1. Matscbie, 8. Mearns, 2. Miller, 10. Milne-Edwards, 1. Miiller (S.), 1. * Nicoll, 1. * Ogilby, 1. Pageustecher, 1. Pallas, 1. Peale, 1. Pelers, 20. Peters & Doria, 1. Quoy & Gaimard, 3. Ramsay, 1. Scblegel, I. * Sclater (P. L.), 1. Seabra, 1. * Smith (A.), 1. Sundevall, 1. Teinminck, 15. * Thomas, 24. * Thomas & Wroughton, 2 Tomes, 1. Tronessart, 2. True, 1. Vabl, 1. Zelebor, 1. The types of 185 forms have been examined by me, and of ten other forms I have had paratypes for inspection. With regard to the remaining thirty-three forms, I have seen topotypes of nineteen, of two types I have had photographs of the skull and dentition, and two have been examined for me by others. This leaves only ten forms, but of six of these the types are, either certainly or probably, not in existence, leaving finally the following four, viz. Pteropus papuanus, Dobsonia pannietensis, Epomops dobsoni, and Megalo(jlossus xvoermanni, but in none of these four cases is there any doubt as to the identification of the species.


AUTHOR S PREFACE. IX Types of synonyms {Hypotypes). — 127 names (variants not counted) occur iu the synonymy of the recognized forms of Megachiroptera. Eight are noraina nuda, and the types of thirty-nine names have not been traced. The remaining eighty hypotypes are distributed as follows : — British Museum, 38; Berlin, 10; Leyden, 7; U.S. National Museum, 6 ; Paris, 5 ; Zi-ka-wei (?), 3 ; two each in the Museums of Frankfort-on-the-Main, Lisbon, Philadelphia, and Sydney ; and one each in Calcutta, Dresden, Stockholm, and Vienna ; while one is probablj* in private possession. 63 of these SO types have been examined by me, while of two other forms I have seen paratypes. In thirteen other cases I have either had photographs of the skull and dentition of the types, or the t3-pe8 have been examined for me by local zoologists, or I have seen topotypes. The remaining two names are Pteropus masMrinus (= Ft. rodricensis) and Odontonycteris meyeri (= Macroglossus laffocliilus). Tlie Catalogue. — Nominally this is a second edition of Dobson's ' Catalogue of the Chiroptera in the British Museum ' (1878^ ; in reality it is an independent work, except of course in so far as every work of this kind must be based on its predecessors. The descriptions are new, the technical names are fixed in strict accordance with the principle of priority, the synonymy is in most cases worked out on the basis of an examination of the types and paratypes, and all references to literature have been compiled by myself. It has been my object on the one hand to make the descriptions of the genera, species, and subspecies reasonably complete, on the other hand to avoid repetitions as far as possible. For the latter reason I have often preferred to give a differential rather than a full description, that is, I have confined the description of a given form chiefly to those characters by which it differs from its nearest fully described relatives. To render the Catalogue easier for reference the descriptions are, whenever required, subdivided into specially headed paragraphs (Diagnosis ; Skull ; Dentition • Palate-ridges ; External characters ; Sexual differentiation ; Speci-' mens examined ; liange ; Habits ; Affinities ; History in literature ; Type, Nomenclature, and Synonymy, »tc.). Detailed measurements are given of the skull aod external dimensions, and as a rule also of the premolars and molajs, of every form described.


X AUTHORS PREFACE. Ill the Introduction to this volume (pp. xvii-ci) I have given a summary of the general characters of the Megachiroptera, of the jirincipal variations within the Suborder of the cranial, dental, and external characters, of the mutual affinities of the genera, of the geographical distribution of the genera, species, and subspecies, and finally, a sj-nopsis of the more important differential characters of the subfamilies and geuera, and an artificial " key " to the genera. The illustrations (80 in number ; see list p. 835) are original, with two exceptions (fig. 32, p. 4!)0, and fig. 51, p. 643), and figures are given of the skull and dentition of all genera and subgenera, except one (Chironax, p. 658). As this is the first Vertebrate Catalogue in which the register numbers of the specimens have been published, it may he explained that of the four successive items composing these numbers, the first represents the year, the second the month, and the third the day when registration took place, the fourth being the individual number of the specimen ; e. (/., means 189S, July, the sixth, number five. The same symbols are employed as have been customary in tliis series of Eritish Museum Catalogues. "[P.] "reads "Presented by," " [C.]" equals " Collected by," and "[E.]" signifies "Iteceived in Exchange." The printing of the Catalogue has taken place, at intervals, between March, 19U8, and March, 1912, but the 'Addenda' at the end of the volume carry the previously printed sheets up to date. The actual date of publication of the volume is March 23rd, 1912. • Aclcnoiuledijments. — My thanks are due, above all, to Mr. Oldtield Thomas, •without whose generous support I should not have been able to devote practically the whole of my time, for several years, to this work, I am also under special obligation to the Authorities of the United States National Museum, through Mr. Gerrit S. Miller, Jr., for the loan without limit of time of a considerable number of specimens (including many jtaratypes), chiefly of the genera


AUTHOK S PREFACE. SI Fleropiis aud Cynopterus, which have been of great help to me during ray work. Further, to Dr. P. A. Jeutink, Professor 1*. Matschie, and Professor Dr. E. Trouessart for much kind assistance during ray visits to the Museums of Leyden, Berlin, and Paris. For the loan of type specimens, for information on types or other specimens in foreign Museums, or for assistance in any other way, I am indebted to Dr. N. Aunandale, Calcutta, Dr. It. Anthony, Paris, the Authorities of the Pombay Natural History Society, Professor R. Collctt, Christiania, Dr. A. Fritze, Hanover, Professor Dr. J. von Kennel, Dorpat, Oberstudienrat Professor Dr. Kurt Lampert, Stuttgart, Professor Dr. Einar Lonnberg, Stockholm, Dr. Ludwig Ritter Lorenz von Liburnau, Vienna, Dr. Marcus W. Lyon, Jr., Washington, Dr. P. Chalmers Mitchell, London, Mr. James A. G. Eehn, Philadelphia, Dr. E. lioediger, Frankfort-on-the-Main, Senhor A. F. de Seabra, Lisbon, and Yiceinspector Mag. sc. Herluf Winge, Copenhagen. The only two illustrations that are not original (see above) have been copied by kind permission of, respectively, the Council of the Zoological Society, London, and Herr Georg lleimer, publisher, Berlin. In conclusion I desire to express my thanks to my artist, Mr. A. J. Engel Terzi, for the unremitting care and artistic skill bestowed by him upon the illustrations. K. A. British Museum (N.H.). January 16tb, 1912.


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CONTENTS. INTRODUCTION. I. General cliaracters of Megachiroptera ... II. Plastic characters of Megachiroptera . 1. Rostrum, p. xix. 11. 2. rremaxilhie, p. xx. 3. Tiifraorbital canal, p. xxi. 12. 4. Postorhital processes and poslorbitnl foramina, 1.3. p. xxi. 14. .5. Palate, p. xxii. IT). G. Tympanic bones, p. xxiii. l(i. 7. Facial axis, p. xxiii. 17. 8. Mandible, p. xxiii. 18. 9. Incisors, p. xxiv. 19. 10. Canines, p. xxvi. 20. Page xvii Premolars and molars, p. xxvii. Anomalies in dental formula, p. xxxi, Palate-ridfres, p. xxxiii. Tongue, p. xxxv. Wing-structure, p. xxxv. Tail, p. xli. Calcar, p. xlii. Colour of fur, p. xlii. Size, p. xliv. Secondary sexual characters, p. xl V. III. Interrelations of the genera of Mcgacliiroptera IV. Geographical distribution of Megachiroptera 1. Distribution of genera, p. Ixv. 2. Distribution of species and subspecies, p. Isvii. 3. Remarks on the geographical distribution, p. Ixxv. V. Synopsis of the subfamilies and genera of Megachiroptera (with a '^Key'' to the genera based on their dental and cranial cliaracters only) l.w


CONTEXTS. CATALOGUE OF CHIROITEEA. Order CHIROPTEKA. Suboi-derl. MEGACHIltOPTERA. Family PTEROFODIIVE. Page Subfamily I. Pteropodinae. 1 [RorsKTTUS section.] 1. Eidolon, linjiii 2 1. diipreanum, Schl. ^Poll. 7 2. helvimi, Kn-r 6, 809 3. siibjeiim, K. And 15 2. Koliseltus, Gray 1

^ Page 43. rayneri, Grnj/ S-'iS 44. rubianus, K. Awl :?oo 45. IftVfllaiius, K. And 258 4(). graiidis, T/io.t 259 47. chrysopvootu.s, Temm. . . 2(J0 H. I't. loinbocen«is group . 2(i5 48. lombocensi.i, Dob/i 2Gt> 49. solitarius, K. And., 269 50. rodricensis, Dohs 273 51. molossinu.", Temvi. .... 275 I. Pt. samoensis group . . 280 52. nawaien.sis, frrai/ 280 5-3. samoensis, Peale 284 54. anetianus, G?yn; 288 J. Pt. pselaphon group . . 293 •55. insularis, Homhr. 4" J«cq. 295 56. phreoceplialus, Thos 298 57. pselaphon, Lay 301 58. pilosus, K. And 306 59. tuberculatum, Pet 309 60. leucopteru.s, Temm 311 K. Pt. temmincki group . . 315 61. temmincki, Pet 316, 822 62. capistratus. Pet 319 63. personatus, Temm 321 L. I't. vampyrus group . . 324 64. piganteus, Briinn 326 a. gigauteus, Bribm 329 b. leucocephalus, Hodys. . 333 6.5, ariel, G. M. Allen 335 66. lylei, K. And 339 67. interniedius, K. And. . . 340 68. vampynis, L 343 fl. maiaccensis, A'. And. . 346 b. vampyrus, L 349 c. pluton, Temm 353 d. edulis, E. Geoff. 356 e. nataiife, K. And 358 f. ItinensLs, Mearns .... 359 M. I't. alecto group 3G3 69. aterrimus, Matxchie .... 363, 822 70. alecto, Temm 365 71. morio, K. And. 370 72. gouldi. Pet 370 N. Pt.couspicillatus group. 375 73. chrysauchen, Pet 375 74. conspiciilatu8, Gould. . . . 378, 823 75. ocularis. Pet 381, 823 O. Pt. neoliiberiiicus group 384 76. papuanus, Pet. ^Jioiia . 38-j, 823 77. neohibernicus. Pet 387 P. Pt. macrotis group .... 392 78. epulnriiis, Panisai/ 392 79. m.acroris, Pet. . . . " 396 80. poliocephalus, Temm. . . 397 Q. Pt. scapulatus group . . 402 81. scapulatus, Pet 403, 824 82. woodfoiHli, Thos 407 rase 5. Acerodon, Jour dan 412 1. celebensis, Pet 417 2. mackloti, T«>!?)i 418 n. mackloti, Temm 419 h. Horesii, Grnii 420 c. alorensis, K. And 423 3. gilvus. A'. And. 423 4. humilis, K. And 424 5. jiibatus, BseJiRch 426 a. jubatus, Eschsch 427, 824 h. miudanensi.s, K. And. . 429 6. lucifer, Elliot 432 6. Pteralopex, Thos 432 1. ancepf, K. And 437 2. atrata, Thos 439 7. Styloctenium, Mntschie . . 442 1. wallacei, Grai/ 44-5 8. Dobsonia, Palmer 448 1. minor, Dobs 460, 824 2. exoleta, K. And 461 3. pannietensis, T)e Vis .... 463 4. moluccensis, Q. S,G 464 5. magna, Thos 466, 825 6. pevoni, A^. Geoff. 467 7. surabana, K. And 471 8. vii-idis, Heude 471 , 825 n. umbrosa, Titos 825 h. viridis, Heude 826 9. crenulata, K. And 47-3 10. praedatri.-?, A'. And 474 11. inermis, K. And. 475 12. nesea, K. And. 476 [Epomophokus section.] 9. Plerotes, K. And 483 1. ancbietSR, Senbra 486, 837 Epomop.s, Gray 487 franqueti, Tomes 494 a. strepitans, A'. And. . . 496 h. franqueti, TodiC^ .... 497 bupttikoferi, Matschie . . 499 dobsoni, Bocage 500 IIyp*ignatluis, H. Alien . . 501 , monstrosus, H. Allen . . 506, 827 Epomophoru.", />^'7f?!(-/< .. 514 wahlbergi, Sund. ...... 521 a. haldomani, Hidl.owell . 522, 827 b. wahlbergi, Snnd 526 labiatus, Temm 529 minor, Dohs. .......... 531 anuru.a, HeugHn. ....... 532 crypturus. Pet 535, 827 g.ambianua, 0//('Wy ^^^'^ ' 7. angolensis, Gray 542 8. pou.'>arguesi, rz-ot/Ms. .. 543 13. Micropteropus, Matschie . 654 1. pusillus, Pet. 557 10. 11. 1 12. 1


Z;V1 CONTENTS. 14. Nanonycteris, Matschie . . 1. veldkanipi, jenttnic .... 15. Scotonycteris, Matschie . . 1. zenkeri, Mol>s. . . h. scherzeri, Zelebor .... 3. major. Miller 4. borsfieldi, Grai/ a. liorsfieldi, Gray .... h. lyoni, nom. n. 5. harpax, Thos 0. princeps. Miller 19. Ptenochirus, Pet.. ..... 1. jagori, Pet 20. Megferops, Pet 1. ecaudata, Temm 21. Dyacopterus, yew. M. .. 1. spadiceus, Thos., , , , . . 22. Balionycteris, Matschie 1. maculata, Thos 2;3. Chironax, gen. n 1. melanocephalus, 2'emm. 24. Thoopteriis, Matschie . . 1. nigrescens. Gray .... 25. Penthetor, gen. n 1. lucasi, Dobs. 20. Sphserias, Miller 1. blanfordi, Thos 27. Nyctimene, Borkhaiisen 1. papuanua, K. And. . . 2. albiveutei', Gray .... Paj;e 559 562 66.3 567 668 572 570 578 580 681 579 582 586 598 598 604 605 609 611 614 622 623 624 625 626 627 629 630 631 632, 827 633 633 643 645 646 650 651 654 654 657 058 660 662 665 665 669 671 674 681 698 700 Patre 3, minutus, A'. And 701 4. variiis, K. And 702 6. cyclotis, K. And 703 5 bis. certans. A', And 828 6. cephalotes. Pall 703 7. geiuinus. A'. And 709 8. major. Dobs 710 9. scitiilus, K. And 711 10. lullula;, Thos 713 ] 1. robiusoni, Thos 714 12. aello, Thos 715 Subfamily II. Macroglossinae. 723 [EoNYCTERis section.] 28. Eonycteris, Dols 728 1. spelaea, Dobs 734 2. major, K. A7id. , 730 3. rosenbergi, Je?itink .... 737 29. Megaloglossus, Payenstecher 738 1. woevrnMim, Payenstecher. 742 30. Macroglossus, F. Cuv. . . 746 1. minimus, E. Geoff. .... 755 a. minimus, E. Geoff. . . 757 b. sobrinus, A'. And 760 2 Iagochilu8, Matschie .... 7(52 a. lagochilus, Matschie . . 703 b. nanus, Matschie 765 c. pygmaius. A'. And. . . 707 d. microtus, K. And 767 31. Sjconycteris, Matschie .. 771 1. crassa, Thos 775 a. papuana, Matschie .... 777 b. keyensis, A'. And. .... 779 c. linschi, Matschie .... 779 d. crassa, Thos. 780 e. major, K. And 780 2. australis, Pet 781 3. naias, A'. And 785 [NoTOPTERis section.] 32. Melonycteris, Dobs 785 1. melauops, Dobs 789 33. Nesonycteris, Thos 790 1. woodfordi, Thos 792 34. Notopteris, Gray 793 1. macdonaldi. Gray 797 2. neocaledonica, Trouess.. . 709 Subfamily III. Harpyionycterinae 799 35. Ilarpyionycteris, Thos. . , 799 1. whiteheadi, Thos 805 List of Illustrations 835 Alphabetical Index = 839


INTRODUCTIOX. I. General ckakacters of Mkgacjiiroptera. The prototype of tlio Chiroptera must have possessed all tlio mo.^t primitive features preserved by any living or extinct form of !Me.gachiroptera and Microchiroptera. Among its more important characters, therefore, must have been these : — Skull. — In general shape probably essentially like that of a liouscttus or Pteropiis, with the following differences : the bony palate was not produced backward behind the tooth-rows (compare Insectivora and Jlicrochiroptera generally) ; the infraorbital canal long {cf. the Notopteris section of Megachiroptera, and somo Microchiroptera); the postorbital processes undeveloped (as in the majority of Microchiroptera"); tlie facial portion of the skull not, or only inconspicuously, deflected against the basicranial axis (as in nearly all genera of the Epomopliorus and Crjnopiei'us sections of Megachiroptera). n .-.• T^ ,-^e ^ t,ui ^' i" — ° P' — p^ p' m' m= nv\ Ventition. — Dental tormula probably .-^-^ *, I1I2I3CP1— P3P4m, ^2^3 this being the completest formula found in any bat (i' may have been ])resent, though lost in all known species) ; molar structure typically " Insectivorous," as in Microchiroptera with unmodified cheek-teeth. Fore-limbs and memhravcs. — Tuheratlum majus and miiuis of humerus relativel}' small, the former not articulating with scapula (cf. Megachiroptera and some primitive Microchiroptera) ; deltoid crest of humerus weak (cf. Megachiroptera). Ulna not more reduced than in Megachiroptera. Articular surfaces of the bones of the hand (earpo-mstacarpal, metacarpo-phalangeal, and interphalangeal joints) not more modified than in Megachiroptera ; trapezium large (cf. Megachiroptera) ; second digit rclalively independent of third, with three phalanges and a well-developed claw (cf. Archcmpteropus f and typical Pteropodida). Wing-i membranes from tlaiiks and inserted i)osteriorly on first toe. ' Tail. — Long and embedded in interfemoral (cf. ArcTiceopteropus, * On the liouiologies of the iiiissing premolars in Chiroptera, see Oldfielcl Thouias. Ann. & Maj;. K. II. (8) i. p. 3i'j (1908). On the missing upper incisor, K. Andersen, P. Z. S. 1908, p. 205. t On Aixhieopteropua traiisieus (Upper Oli^ocene, Monleviule, Italy) see IMesfhinelli, Atti R. 1st. Veneto, Ixii. p. 1329, pi. is. (1903). Note on il3 wine-struolm-e, infra, p. ssxvii. I,



I PLASTIC CHAKACTKRS. XIX a similar ridgo along the iuner side, the two ridges being separated by a median longitudinal depression ; in the sirailarlj* shaped lo'vrer niola.riform teeth the inner ridge is formed by cusps 1, 2, and 3, tlie outer by cusps 4 and 5 : (4) ig and m' are lost : (5) The second and, particularly, the third phalanx of the second finger are somewhat reduced in length. The principal plastic characters of the Megachiroptera are discussed in the next paragraph. II. Plastic characters of Megachiroptera. 1. Rostrum. Varies considerably in length, much less so in sliape. Length. — If measured from the front of the orbit to the extremitjof the nasal bones, the cranial rostrum is : — (1) Longest, i. e. from about f to nearly ^ of the skull, in Hi/psir/nathits and some species of Ejiomophorus, in both genera relatively longer in males than in females ; the male ffypsignafhus is the longest-faced of all Fruitbats : (2) Medium or somewhat shortened, i. e. from about g to |^ of the skull, in : — (a) the genera of the Bousettus section ; (b) all Macroglossince ; (c) within the EpomopiTiorus section in the genera Plerotes and Epomops, and in some (the smaller) species of Epomophorus; (d) within the C'ynopferus section only in Myonycteris and Spharias : (3) Shortest, i. e. about I— I of the skull, in :— («) the following four genera of the Epomoplwrtis section, Micro pteropus, Nanonycteri.'; (rostrum rather more than ^ of skull), Scotonycteris, and Casinycieris : (6) all genera of the Cynopterus group, except Myonycteris and SpTK^rias. Briefly summed up : a moderately long rostrum is characteristic of all Eonsettine and Macroglossine Fruit-bats, a very short of nearly all Cynopterine genera, while the Epomophorine is the only section showing any degree of variation in the length of the rostnira, from that of Plerotes (moderate), on the one hand to that of Ilypsrgnathus (longest known), on the other to that of Casinycieris and allied genera (shortest). No Fruit-bat with a very short rostrum (third stage above) has the full Megachiropteran number of cheek-teeth (the formula being eilher j, j, or A\ but this rule cannot be reversed, that is, a reduction in the number of teeth is by no means always, not even usually, associated with a shortening of the rostrum. Fruit-bats with a moderately long or very long rostrum (second and first stage above) exhibiting any variation of the dental formula from the highest number of cheek-teeth (|) to the lowest (0. Shape.— As a general rule the rostrum is conspicuously tapering 62


XX PLASTIC CnARACTERS. jinteriorly, and tlio aberrations from this typical (Insectivorouslike) sha[)e are, apart from Ilypsignatlms, but few and not very peculiar. The rostrum is unusually deep (truncate) in front in JSIeqarops and Nyctimene (both of the Cynoptevus section) ; in Niictimene the medial portion of the nasals is produced forward and downward as a triangular projection firmly united with the anterior edge of the mesothmoid cartilage (evidently acting as a support of the nasal tubes; c/. fig. 61, p. 684). In Fruit-bats with weak dentition (narrow cheek-teeth) the rostrum is lower and thinner than usual ; compare Stenonycteris (fig. 3, p. 49) with llousettus (fig. 2, p. 17), Pteropus scapulatas (fig. 18, p. 404) with the average Pteropus skull, Sphcprias (fig. 60, p. 672) with the ordinary Cyaopterine type of skull (fig. 48, p. 588), and all Macro(/lossince, except the more heavily-toothed Eonycleris. The only strikingly peculiar modification is seen in males of Hypsignathus {fig. 33, p. 502) 4 the rostrum is greatly increased in size and jjarticularl}' in depth, its dorsal and ventral profiles subparallel, its lateral surfaces concave (a large subcutaneous air-sac is present on either side of the rostrum), and a strong vertical crest runs from the nasals to the alveolus of the canine (supporting a cutaneous fold of the upper lip) ; in females of the same genus the rostrum is somewhat similarly, but much less excessively, modified. 2. Premaxillce. Vary in shape (breadth, degree of reduction), in direction, and in the mode of interconnection of the alveolar branches. The palatal branches of the premaxilliB are always absent. Breadth. — The three Macroglossine genera Mdonycteris, Ntsonycteris, and Notopteris exhibit what is probably the most primitive form of the premaxilke : the ascending branch is unreduced in breadth at its upper extremity, gradually narrowing interiorly, so that the breadth of the bone near the alveolar margin is only one-half or one-third of its breadth at the upper extremity (fig. 76, p. 791). The next stage is shown by genera in which the ascending branch is considerably reduced in breadth above, being quite or nearly as narrow above as near the alveolar margin (ex. Pteropus and allied genera ; Epumophorus ; Eonycttris ; Meijaloylossiis ; Macroqlossus; &c.) ; a still higher degree of reduction by genera in which the bone is tapering to a point above, its upper extremity often slightly curved forward (ex. nearly all geneia of the Cynopterus section) ; and finally, the whole of the ascending branch may become so thin as to be almost linear throughout (Dohsonia). The whole of the premaxillary region is increased in breadth in Bypsl(jnathus (owing to secondary modifications in the shape of tlie rostrum). In Nyctimene (fig. 61, p. 684) the alveolar branches are unusuall}' deep (vertically), the ascending branches so short as not to reach the nasal bones (extremity of rostrum modified, for support of the peculiar nasal tubes in this genus). Direction, — As a rule the ascending branches of the prcmaxillse are S4


TLASTIC CHARACTERS. ri carl}vertical oroulyslightlvproclivoiis. A somewhat, or even much, liighcr degree of proclivity is seen in Fhrotes, Epomops, SiJucrias, all Macrofjlossiiw; except Eomjcteris, and in Harpyioivjctens. Interconnection of branches. — In the large majority of Fruit-bats the right and left alveolar branch of the preraaxillte are in simple contact anteriorly, the suture between them being permanent. In certain genera, however, they become, at an early age of tho individuals, united by synostosis, i. e. firmly and solidly ankylosed together, without any trace in the adult (and, as a rule, not even in the semiadult) of an inter-premaxillary suture ; this is the case in Lissonycteris (subgenus of liousettus), Fteralopex, and Harpyioni/cteris, all of the Bouseitus section, in Hy psiynathus among the lipomophorine Fruit-bats, in Dyacopterus, C'hironn.v, and Ni/ctimene, of the Cynopterus section, and in AJeyaloyhssus, 2Iaeroylossus, Syconycteris, and Xotopteris, among the JIacroylossinee. The opposite extreme is represented by a few genera in which the premaxilloe are not even in contact anteriorly, but distinctly spaced, connected by fibrous tissue only (constantly 'in Eidolon and Boneia ; occasionally in Eonycteris and Mdonycteris). 3. Infraorbital canal. In Insectivora, as well as in primitive Mammalia genenilly, the infraorbital canal is long, its anterior aperture, the infraorbital foramen, situated a considerable distance in front of the orbital cavity. The only forms of Megachiroptera which in the length of the infraorbital canal approach to this primitive condition are the three genera Melonycteris (fig#75, p. 786), Nesonycteris (fig. 76, p. 7U1), and Notopteris (tig. 77, p. 795), all closely interrelated and belonging to the subfamily JIacroylossince. In all other Fruit-bats the canal is considerably shortened, its outer wall, as a rule, a narrow, often sublinear, bridge of bone, and the foramen situated nearly vertically below or only slightly in front of the orbital cavity. 4. Posiorbital processes and postorbilal foramina. Processes. — The postorbital processes are generally small or moderately strong, and if so there is scarcely any trace of corresponding lower processes from the zygoma. In a small number of forms, all with heavy dentition (chiefly species of Ftcropus and allied genera), they are stronger than usu'al, reacliing about halfway between frontal and zygoma, and the lower processes are more or less conspicuously developed. Earely (a few large species of Pteropus and Acerodon ; Pteralopcx) they are, at in aged individuals, so long as to join the lower processes and form a complete ring round the orbit. Foramina. — The base of the postorbital process is nearly always pierced by a relatively large foramen (no doubt homologous with the supraorbital notch or foramen of the human skull, for


XXU PLASTIC CHARACTERS. Nerviis and Vasa supraorhitalia). The oulj section of genera in which exceptions to this rule occur is the Cynopterus section ; the postorbital foramina are, within that group of Fruit-bats, normall}" developed only in Myonycteris, Cynopterus, Ptenochirus, and Aleycerops, minute (tending to disappear) in Dyacopterus, generally absent (or, it' present, minute or traceable only on one side of the skull) in Nyctiinene, always absent in Balionycteris, Chironax, Thoopterus, Penthetor, and Sphcerias. 5. Palate. Pteropus (fig. 6, p. 62) is probably one of the genera that shows the Megachiropteran palate in its least modified form. The interdental palate is only moderately broad, gradually narrowing in front (the tooth-rows distinctly converging anteriorly); the postdental palate about one-fourth of the total length of the palate, its lateral mai-gius forming almost straight lines converging posteriori)-. In Ho usetius and Boneia the interdental palate is relatively broader; an even greater increase of breadth is seen in Epomops, and in Plerotes this portion of the palate reaches its maximum of breadth (bearing a curious resemblance, in general outline, to that of the Carnivorous genus Proteles !). An opposite course of development is taken by Epomophorus, in which the palate is unusually narrow. If the lower canines are slanted outward, this necessitates a greater breadth of the palate anteriorly, between the upper canines, and therefore more nearly parallel tooth-rows (ex. Boneia, UypsignatJius, Macroglossus). The postdental palate is in the majority of genera essentially similar to that of Pteropus described above ; it may be broader or narrower, shorter or longer, more rapidly narrowing posteriorly or more nearly parallel-margined ; in Nyctiraene it is somewhat pandurate in outline, i. e. distinctly constricted at middle (owing to the unusually broad mesopterygoid fossa ; compare the presence of nasal tubes in this genus)But these variations are comparatively trivial as compared with the modifications exhibited by some genera of the Epomophorus section : — In Plerotes, Epomops, Hypsigiiathus, and Scotonycteris it is relatively simple in shape, though in the two former genera rather broader than usual ; but in Epomopliorxis (fig. 36, p. 515) it is deeply depressed posteriorly, and its posterior free margin high and prominent ; m Micro pteropus (tig. 38, p. 555) it is abruptly narrowed behind the roots of the zygomatic processes, the posterior depression is shallower, the free margin prominent ; in Nanonycteris (fig. 40, p. 560) it is short, but unusually broad, longitudinally depressed on either side, slightly convex between, and flattened at the extreme posterior extremity, the free edge not prominent ; and, finally, in Casinycteris (fig. 43, p. 569), a genus in other respects closely similar to Scotonycteris, the postdental palate has practically disappeared, the mesopterygoid fossa extending forward very nearly to the level of the posterior molar, a modification unique in Megachiroptera and more closely recalling the type of palate found in Microchiroptera,


PLASTIC CHAKACTEK9. XXUl 6. Tympanic bodies. Alwajs annular, but the ring varying a little In breadth (rather broader than usual in liousettus, Dobsonia, Eouyctey-is). The only peculiar modification is the development of a short bony auditory meatus in Eidolon (fig. 1, p. 3 ; cf. Miller, 'Families and Genera of Bats,' p. 55, fig. 7 B), a genus closely related to liousettus. 7. Facial axis. Normally the facial axis if projected backward is nearly parallel witli the basicrauial axis, i. e. the former forms an angle with the latter of not much less than 180 degrees, and the line of the alveolar margin of the upper tooth-row if continued backward passes through the lower edge of the occipital condyle or even somewhat below the condyle ; such is the case, with rare exceptions, in all genera of the Epomophorus and Cynopterus sections (ex. fig. 36, p. 515, and tig. 48, p. 588). In the Eousetto-Pteropine section generally the face is more distinctly, sometimes even very conspicuously detiected on the basicrauial axis, so that when the rostrum is kept horizontal the axis of the brain-case points obliquely backward-and-downward, and the posterior projection of the alveolar line passes through the middle or upper margin of the occipital condyle (ex. fig. 1, p. 3 ; fig. 6, p. 62) or even through the middle of the supraoccipital (fig. 3, p. 49 ; fig. 18, p. 404). The maximum of facial deflection is seen in the majority of Macroglossincn, in which the face is bent downward to such degree that the alveolar line if continued posteriorly would pass through the brain-case considerably above the lambdoid crest (fig. 70, j). 748). As a general rule the deflection of the face is greatest in genera or species with weak dentition ; thus it is considerably greater in the narrow-toothed Stenonycteris (fig. 3, p. 49) than in typical Rousettus (fig. 2, p. 17); in Pteropus the deflection is always noticeable, but least so in the heavy-toothed species (ex. fig. 15, p. 302, Ft. p)selaphon), and unusually great in the smalland narrow-toothed Ft. scapulatus (fig. 18, p. 404) and Ft. woodfordi (fig. 19, p. 408) ; the only Epomophorine genus with conspicuously deflected facial axis is the very weak-toothed Flerotes (fig. 28, p. 484), and the only Cynopterine genus with similarly deflected face is the weak-toothed 8p>ha;)-ias (fig. 60, p. 672) ; within the subfamily MacrocjlossincB the deflection is smallest in the relatively strongtoothed Eonycleris (fig. 66, p. 730), greater in Meyalaglossus (fig. 68, p. 739), which in the degeneration of the dentition is transitional between Eonycleris and Macroglossus, very great in all other genera, all of which are characterized by unusually narrow or small cheek-teeth. 8. Mandible. The heavier the dentition, the broader and deeper and more vertically ascending is the coronoid process of the mandible, the stronger its angular process, and the higher above the alveolar line its condyle ; the weaker the dentition, the narrower and lower and more backwardly sloping is the coronoid process, the feebler the


Xii.IV PLASTIC CnARACX'EliS. angular process, and the lower dowu the position of the condyle, Tliere is within the suborder every gradation from the unusually heavy mandible with nearly vertical coronoid process, deep and broadly rounded angular jjrocess, and the cond\le situated high above the alveolar line, of Pteratopex (fig. 21, p. 433j, Acerodon (fig. 20, p. 413), the heavy-toothed species of Pteropus (fig. 14, p. 281) ; fig. 15, p. 302), Dohsonia (fig. 24, p. 449), and most genera of the Cyuopterine section (fig. 48, p. 588 ; fig. 61, p. 684), through the intermediate conditions as exhibited in the Epouiophorinc section (fig. 30, p. 488 ; fig. 40, p. 560), to the mandible as shaped in the narrowest-toothed Macroi/Iossina; (fig. 70, p. 748), with its short, thin, and posteriorly directed coronoid process, weak angular process, and condyle situated considerably below the level of the alveolar line. The symphysis of the mandible is in the Pteropodince usually obliquely ascending, more rarely (ex. Dohsonia, Nyctimcne) subvertical ; in the Macroglossince as a rule more nearly horizontal. In some genera of Macro(jlossince {Eonijdcris, Mei/alor/Jossus, Macro(jlossus, Mdonycteris, Nesomjcteris) there is a more or less clearly pronounced tendency to a development of a longitudinal keel along the antero-inferior surface of the symphysis. 9. Incisors. Incisor formida'. — i^ is lost in all Chiroptcra ; ij is present in many Microchiroptera, but lost in all Mcgachiroptora (the incisors of Arclia'opteropus are unknown) ; no Fruit-bat, therefore, has more than % — 5 incisors, but some have less. The following five incisor formuliB occur : — iiMM 2 2 .-. = , the normal formula, found in all genera except '^ 'i 'i '-^ ^ , ^ those enumerated below. j2 i^ 1 1 . = , in one genus, Boneia (allied to lioitsetfus). \h\h ^' ^ 42 \\ p V9 2 . = , in one genus of the Ptcrojms subsection, viz. Sh/locfe7iium ; in four genera of the Cynopterus section, ptcnochirus, Megcurops, Balionyctcris, and Penthetor; and in two Macroglossine genera, I\esonycicris and JS'otoptcris (in the latter i^ is deciduous). ;2 i^ 1 1 = , in Dohsonia and Ilarpyionyctens^ both aberrant i., ij 1 '^ genera of the Pousethis section (on the homologies of the missing incisors in these two genera, see footnotes p. 452 and p. 801). i'i' 1 1 Z = , in Kiictimcnc onlv, a i)eculiar]v modified genus of the Cynopterus section.


I'LASTIC CnAEAClEES. In Pkropus i, is always conspicuouslj^ smaller than i„, and in some species {I't. lomhocensis group) so luuch reduced as to be almost rudimentary ; this foreshadows Stylodenium, in which ij is completely lost, and Dobsonia and Ilui-pyionijcteris, in which both ij and i' are lost. In those genera of the CynojJients section which have the normal number of incisors i, is Aveaker than i„, and i" iuintly shorter than i' ; this leads to those genera of the same bcclion, viz. I'tenochirus, Meijcerops, Balionijcteris, and I'eniheior, in which ij has disappeared and i" is shortened to about half the length of i' ; and to JVyctimene (lower canines moved forward to extremity of jaw), in which all lower incisors as well as i" have disappeared. In the Macroglossine genus MeJonyctens \ is smaller than i.^, in the closely related J^esomjcteris and Is'otoptivis it has disappeared. Differentiation. — In the majority of Fruit-bats the incisors are small, thin, terete (styliform) or subterete, the crown only slightly or even indistinctly differentiated from the shaft, the cutting-edge of the lower incisors often faintly bilobed. Normally the incisors above and below are placed approximately vertically, those in the upper jaA\" in contact with each other or narroAvly spaced, but with a relatively wide diastema on either side between the incisors and canines, those in the lower jaAV crowded and in contact with the canines. The principal variations, to be described below, from this predominant type of incisors are, briefly summed up, these : (1) the crown of some or all of the incisors may become peculiarly diiferentiated in shape, (2) conspicuous posterior ledges may be developed, or (y) some of the incisors may be enlarged or reduced; further, (4) the direction of the incisors may be altered, and (5) the spacing of the incisors may be conspicuously unequal. In Fteroinis (and the closely allied Acerodon) the upper incisors are larger than usual, the crown distinctly differentiated, posterior basal ledges conspicuously developed, sometimes so much so as to form a noticeable shelf (7^<. pscZo^^/ion group), i, always larger than ij, either owing chiefly to a reduction of ij, which may become nearly rudimentary (Pt. loinhocensis group), or chiefly to an increase of i^ {Ft. samoensis and iisda^ihon groups). Some of these tendencies are further developed in the related I'teralopex : the posterior ledge of the upper incisors is very large, shelf-like, rendering the antcro-posterior equal to or greater than the transverse diameter of the teeth, and i, is from twelve to fifteen times the bulk of i^, its cutting-edge unequally trifid (middle cusp much the broadest). In Splurrias {Cynopterus section) both the upper and lower incisors arc triangularly pointed. In Syconycteris {Macroijlossivcp) the upper incisors are larger than usual, the crown well-differentiated, narrowly chisel-shaped, i„ considerably higher and broader than i^, with obliquely triangular crown. In those genera (Dolsonict, Ea)pyionyci^ri.i, ]\''yctimenc) which have only [ — j or ^ — ^ incisors, and in which the lower canines are situated close together or in actual contact at the anterior extremity of the mandible, the single pair ol upper incisors is acted upon by


PLASTIC CHARACTERS. the tips of the lower caiiines and somewhat altered in shape, the crown being more or less obliquely bilobed. The normal vertical direction of the incisors is changed into a strong proclirity in SjjfKerias, Syconycteris, and Harpyionyctens. Aberrations from the normal crowded or equidistant arrangement of the incisors are seen in several genera of Maeroglossince : in Melonycteris i^— i^, in XesonycUris and Xoiopteris \^—i\ and in Macroglossus both i^— i' and i,— i^ are unusually widely spaced. In Hypsignathm {Epomophorm section) the lower incisors do not bite against but close some distance in front of the upper. Deciduous incisors.— \' (rudimentary, barely piercing gum) is deciduous in XotopteAs ; i^ (relatively well developed) otten deciduous in Epomops (for details see p. 489, footnote) ; i^ (the single lower incisor, almost functionless) sometimes deciduous in Dobsonia {D. viridis and crenulata), perhaps so in Harpyionyctens. 10. Canines. Apart from minor variations in length and bulk, the canines may be moditied : (1) by an enlargement of the cingulum, (2) by the development of secondary cusps, or (3) by the development of longitudinal grooves on the crown of the upper canines ; further, (4)°the direction of the canines may be changed (proclivity ;_ outward or outward-and-backward slant of lower canines), and (5) the position of the lower canines may be changed. Ciru/ulum.—As a general rule, in Fruit-bats with weak dentition the cingulum of the canines is thin or obsolescent ; on the other hand, the heavier the dentition, the thicker and more prominent the cingulum. In Pteropm (dentition, as a rule, unusually heavy) there is a pronounced tendency to an enlargement of the cingulum, particularly in the upper canines, and m certain species {Ft. samoeasis and pselaphoa groups) the prominent edge of the cin-ulum tends to break up into a number of small, rounded, more or less incompletely separated tubercles. The extreme of this tendency is seen in the related, very heavy-toothed FteraJopex, in which the edge of the broad cingulum of the upper canines is distinctly cuspidate. Secondary cusps.— ^aj be developed from the inner edge only (in both upper and lower canines), or from the outer edge only (in the upper canines), or from both edges (very rarely, and only in the lower canines): — _ From inner edge only :— In Cynopteitis (fig. 48, p. o88) and the closely allied Ptenochirus (fig. 51, p. 643) a distinct secondary cusp is developed near the middle of the inner edge of the upper and lower canines, produced by a prolongation of the cingulum. From outer edge only:— A small, well-marked secondary cusp from the outer edge of the upper canine, above the middle of the tooth, in Pteropus tubercidatus : a similar, but much larger cusp


PLASTIC CHARACTERS. XXVll halfway up the outer edge of the upper canine, in Fteralopex (fig. 21, p. 483); a similar, strong cusp in Harjiyionycteris (fig. 78, p. 800) ; often a more or less well-defined outer cusp in the upper canine of jYyctimene (fig. 61, p. 684). From hoth edges : — A strong secondarj' (cingulum) cusp at lower half of outer edge and a smaller one at middle of inner edge, in the lower canine of Harpyionycteris (fig. 78, p. 800). In this genus, it will be noticed, the upper canines are bicuspid, the lower tricuspid. Grooves. — The anterior (or antero-medial) surface of the upper canine is not infrequently marked with a deep and well-defined longitudinnl groove, reaching nearly from the base to the tip of the crown (and corresponding to the line followed by the tip of tho lower canine, along the crown of the upper canine, when the lower jaw is moved up and down) ; this groove is particularly conspicuous in Boneia (as a rule obsolescent or shallow in the related liousettus), Pteropus (rarely obsolete ; in the related Pteralopex shallower or rather indistinct), Styloctenhim, in all genera of the Cynopterus section except Myonycteris, Cynopterus, Meyarops, and Nyctimene, and in all Macroylossince. More rarely, and only in some genera of Macroylossin(x, there are one or two additional longitudinal grooves on the outer surface of the upper canine (barely traceable in Macroglossus, shallow in Syconycteris, as a rule better developed in Melonycteris, Nesonycteris, and 2iotopteris). Direction. — The lower canines are slanted conspicuously outward or both outward and backward in Boneia, PleroU-s, Epomops, Eypsiynathus (lower canines, like lower incisors, closing some distance in front of upper), Sphcerias, and all Maeroylossince. In Dolsonia the upper canines are a little proclivous ; in the related Harpyionycteris both upper and lower canines (as well as the incisors) are unusually proclivous, the lower canines crossing the upper almost at right angles. Position of lower canines. — In genera with a single pair of lower incisors the lower canines have, as a rule, moved a little closer together, toward the extremity of the jaw {Siyhctenium, PtenocMrns, Meycprops, Balionycteris, Penthetor; not in the two Macroglossine genera Nesonycteris and Notopteris). The extreme phase of this tendency is seen in three genera, Bohsonia, Harpyionycteris, and Nijctimene, in which the lower canines have moved forward to the very extremity of the jaw, so as to be quite or nearly in contact with each other (in Dohsonia and Harpyionycteris the single pair of lower incisors is rudimentary, wedged in between the canines, and at least in Dohsonia sometimes deciduous ; in Nyctimene all lower incisors are lost). 11. Premolars and molars. Formulct. — p^ and p^ are lost in all Chiroptera, m' in all Megachiroptera ; no Fruit-bat, therefore, has more than ^ postcaniue


xxvm pimstk; ciiakactkus. tfotli on ciUior side. The following seven formulae occur in tlic Buborder : — ji' p' p* m' m'' _ ''^ ii It n~T) nrm~ni ~ G' "ormal formula, found in all gcneni not enumerated helow. p' p' p^ m' m_ 5 . 11 i)~rrnr~m^^ ~ ^i' '" '^fll'-f^cteidani (related to I'icropus) ; Bal'ionjfckrts (Cy nop terns section); and one species of Macrojlossincn, Konj/cteris rosenberr/i (tlic other species of Eonycteris have the normal formula). — p' p' m' m^ _ 4 . ^ ' ^_^ m Dohsoma, an aVjcrrant trenus of the P. P3 Pi ™i °i. ™.-, '^ llouscllus section. p'p3p4mi_ _4 . ,„ -, in y lerotes (hnomop/iorus section). Pi p.. Pi f"i "'j I",-, ^ V i i / ])' p" p' m' — _ 4 ,, , „ , . , ^, ,, the usual lormula la the CynopUrus section, I'i Pi Pi "^j "' — " J J. 1 found in eight genera, viz. Cijno [items, PteiiocJiirus, Mcrj(erops, ChirorMX, Thoo/iterus, Penlhelor, Splurrias, and JVyctimene; further, in one sjjecies of Macro(flossime, iSj/ronT/cteris nains (the other s[iecio3 of Syconijcleris have the normal formula). — 1)^ p* m' m* 4 . = in Kotorderls (subfamily Macroqlossinoi) . — P3 Pi ™> "1. ^3 5' ^ ^ J J J — ~ — = -, the usual formula in the Eiiomoi/honis secPi P3 P4 "i. ™.— 5 tion, found in seven genera, viz. Epoinops, Jfiji'stf/iialhii8, Epomopliorus, Microplero/ms, Nanoni/cleris, Scotonycteris, and Casinycteris ; and in one genus of the Cynopterus section, JJyacoplerus. Formuhj: in iJie four 2->rimary sections of MeyacJiiroplcra. — All genera of the Itovseltus section Lave the normal number of premolars and molars (''), except Styloctcniwa, which has lost the small nij, and JJohsonia, which has lost the small p'. All genera of the Epomophoriis section have lost p', m'', and m^ (^ cheek-teeth), except Fhrotes, which has lost only m% and retained p' and m,, in a rudimentary state. All genera of the Cynopterus section have lost m* and m^ (* check-teeth), except Myonycteris, which has the full numher of check-teeth (but m^ and m^ are quite small), Bulionyetcris, which has lost only m^ (but m' is minute), and JJyacopterus, which has lost not only m^ and m^, but also the small p'. All Jilacrofjlosshue have the normal cheek-tooth formula (j!)^ except Eonycteris rosenherrji, which has lost m,, fin the other species of Eonycteris m^ is usually very small, and adult individuals occasionally occur in whicli it is absent at least on one side),


ri.ASTtO CnARACTEIiS. Si/con>/cte)-is iKiias, which lost in, and nr (hotli small in tlio other spt'cios of iho goims), aud JSotopta-ia, which lias lost p' and p (in the related MclonT/cteris and AcnoDi/citris p' is rudiiuoiitarv'). Modijications of molar sirwture. — Normally the structuro of the molaritbnn teeth iu Me?:achiroptera is this: a median lonjiitudinal groove flanked by a higher enter and lower inner ridge, eaeh ridge lising (or tending to rise) into a cnsp anteriorly. This typical structure is most clearly pronounced in p' and ni', p^, m,, and ni„. In p-' and p., the outer and inner ringo raises anteriorly into a higher, more narrowly pointed cnsp, and both cusps are either connected at base by a conspicuous commissuro or (very often) completely fused from base to tip, making these teeth iii i)rotile almost eaniniform. If present, ]>', p, and m, are nearly alwavs reduced iu size, and their surface structure more or less degenerated. The principal modilkations of the typical molar structuro are due to : (1) the development of a well-marked posterior basal ledge, (2) the development of an antero-internal basal ledge or cusj)! (13) a more or less complete splitting of the outer or inner ridge, or both, into two or more cusps, or (4) the development of surface cusps. If only one of these modifications becomes operative, the general ai>pearauco of the tooth is comparatively little altered; if several (/'/crw/o/),;.!-) or all {JJarpi/ioni/dena) take effect in the same tooth, its appearance is naturally profoundly modilied. In rta-opus there is a pronounced tendencv to the developnu-nt of a posterior basal ledge in p', p', p.,, and p„'often also in m' and nip not infrequently in m.^, i.e. the posterior portion of the tooth is more or less distinctly marked off, by a notch in the outer ridge, from the rest of the tooth (fig. 9 B, p. 08; llg. 101], p. (Jil); these posterior ledges are unusually strong in the rurojnis samoensis and pseliqtJwn groups, and in one species of the former group (7'/. anedanus) the ledge is continued forward, as a well-defined shelf, along the inner side of p^ ni„ and ni.^ (fig. 10 C, p. (;9 ; fig. 14* p. 289). In some species of the Ptcro}}iis lomhoccush, samocnsis, and psilajthon groups there arc traces of antero-internal basal tubercles in \)\ p^ p„ and p, or in some of these teeth, i.e. the antero-internal cingulum is some\\hat more ditVcrentiated thuu usual and tending to rise as a small ledge or tubercle. Tliese modifications aro further developed and more definitely fixed in Jcerodon (closely related to rUropi.s) : the jiosterior ledges aro always strong, in p^, m,, and m.^ the ledge always extends along the inner base of the teeth as a broad, sh.irply-defined shelf, and a well-marked antero-internal tubercle is always developed in' ])' and m', in some species also iu p' and p^, making these teeth distinctly tricuspid (fig. L'O, p. 413). The peculiar molar structure of PUraJojtcv (also related to rtcropux) represents only an extreme phase of the modifications developed in the sjiecies of the rtcropns jisclap/ion group. The teeth are shrM't and broad, the anteii(Uand post(-ri(>r ledges of }>",


XXX PLASTIC CHAKACTEES. p\ and in' prominently developed, slielf-like, with raised margin, the posterior ledges of the lower teeth heavy, and the cutting-edge of the outer ridge of p^ and m^ distinctly (though never deeply) bifid (fig. 22, p. 438). Again, in Dohsonia (an offshoot from the Rousetto-Pteropine section) some of the modifications seen in Pteropiis are taken up and further developed. In a single species {Dohsonia minor) the molar structure is practically unmodified Rousettine, hut in the more highly diff'erentiated forms of the genus there is a wellmarked posterior basal ledge in p^, p\ Pg, and p^, and a distinct antero-internal basal ledge or cusp in p' and p'', often also in p^, p^, m^, and m'; to these modifications are added, in the majority of species, a median surface ridge (or cusp) in m' and m^, sometimes also in p'' and m^, and a tendency to a splitting of both ridges of p*, m\ p^, and m^ (rarely p^ and ]}') into two or more cusps (fig. 25, p. 451). A distinct (but never very deep) splitting of the ridges of some of the cheek-teeth was noted above in Pteralopex and Dohsonia. The same tendency crops up in several other Fruit-bats : in Pteropus pselapJwn the inner ridge of m,, and in Pleropus leucopterus both ridges of m^ and m,^ tend to become bilobed ; in ffi/psignathus the outer ridge of p^ is more or less obscurely, that of m^ always distinctly trilobed or bilobed, that of m, bilobed (rarely, and only as an individual anomaly, an initial stage of a splitting of the outer ridge of m^, or both m^ and m', into two cusps is seen in the related EpomopJiorus ) ; in Nyctimene (Cynopterus section) the outer ridge of p^, p^, and m^ is sometimes more or less conspicuously bilobed ; and, finally, in Earpyionycteris (related to Dohsonia) the splitting of the ridges is more complete than in any other genus of the suborder. The occurrence of surface cusps (or ridges) in p'', m\ m^, and m^ of Dohsonia was mentioned above. Similar cusps are developed in the related Harpyionycteris, and in p, and m^ of four genera of the Cynopterus section, viz. Cynopterus (character not quite fixed), Ptenochirus, Dyacoptemis, and Thoopterus. The " multicuspidate " molar structure of Earpyionycteris is unquestionably the most peculiar in the suborder. In reality, however, it is eff'ected simply by a combination of all the four types of modification discussed above, viz. by the development of posteroexternal and antero-internal cusps or ledges and of surface cusps, and by a splitting of the ridges (see analytical description of the teeth of this genus, pp. 801-803, and fig. 79, p. 802). Deciduousness of pV — Even when reduced to a quite rudimentary (and therefore presumably almost functionless) condition p' is in most genera permanent. It is known to be deciduous in a few species of Ronsettus (seminudus, hrachyotis), in many species of Pteropus, and in Acerodon and Styloctenium (perhaps occasionally in Chironax). [It is permanently lost in Dohsonia, all Epomophorine


PLASTIC CHARACTERS. XXXI genera except Plerotes, in Dyacopterus (probably ; only one specimen is known), and in Notopteris. The latter genus is the only Fruit-bat that has lost both p' and p^.] Degeneration of chceh-teeth. — Probably owingto adaptation to a kind of food that requires little mastication, the cheek-teeth are imusually narrow (sublii)ear)iii >S'^6'no>M/c7is scapvlottis at least twelve per cent, will present anomalies in tlie dental formula, whereas in all other Fruit-bats taken together the percentage has been found to be only 0*9. The anomalies observed may conveniently be classed under the following four headings : — (1) Absence of teeth which are normally present in the species (it is hardly necessary to say that senile conditions have been left entirely out of consideration, and that a tooth has been considered " absent " only if there is no trace whatever that it has ever been present in the individual), this anomaly in some cases " foreshadowing "' the permanent disappearance of the same tooth in related forms : (2) Appearance of teeth which are normally absent in the species, but present in related forms of Itregachiroptera : (3) Appearance of teeth which are normally absent in all Megachiroptera, but (since present in some Microchiroptera) have probably been present in some ancestral form : (4) Accidental outgrowths. Whether a case ought to be classed under the fourth or third category is, as noted below, sometimes doubtful.


XXXU PLASTIC CHAR.VCTERS. Alisence of teeth that are normalli/ present in the species. — The moat frequent anomaly is the absence of the last lowei* molar (m, or ra^) or the last upper molar (ra'^}; more rarely p', pi, or i^ are undeveloped: — inj. — Absent on one side of the jaw, in four cases, viz. one Ptcropus vampi/r/f.i vampi/rus (British Museum, no. 9.1.,5.8(.)7), two Eonycteris spelaa (B.M.,, one Macroc/lossus minimus minimics (10.4.G.1.'»). Absent on both sides, in five cases, viz. one Pteropus qiqantcus (, four Pteropus soapwlatm (.')7. 10.24.1,, 8.8;8.'4, m,, is always greatly reduced in size in Pteropus and Eonycteris spelma ; in a gouus closely related to Pteropus, viz. Stylocteniam, the tooth has perniauently disappeared, and the same seems to be the case in one species of Eonycteris {E. rosenhcrgi). In Pteropus scapulatus and Eonycteris the cheek-teeth are tending to degenerate in size (breadtli), and in Macroglossus the degeneration is even more pronounced. m'. — Absent on one side, in one Pteropus scapulatus ( Absent on both sides, in one Macroglossus minimus minimus ( As noted above (under m^ both are forms with degenerated dentition, ni.,. — Absent on one side, in two cases, viz. one Cynopterus hrachyotis angulatus (, and one Cynopterus hrachyotis minutus (U.S. National Museum, 141243). In Cynop)terus and related genera the number of cheek-teeth is reduced to r, tliat is, nij and m^ are lost ; ui^ is therefore flie last lower molar, and it is always considerably reduced in size. p\ — Absent on both sides, in one Eonycteris spelaa ( In Eonycteris, as in many other Mogaehiroptera, p'^ is rudimentary ; in certain genera it is deciduous or lost. p,.— Absent on one side, in one Acerodon celehensis ( p^ is always small, but in no Fruit-bat deciduous, and only in one genus {Noiopteris, entirely unrelated to Acerodon) lost. ij. — Absent on one side, in three cases, viz. one Cynopterus hrachyotis angulatus (, one Cynopterus h. hrachyotis (, and one Cynopterus h.javanicus ( This anomaly, it will be noticed, has only been observed in Cynopterus, and ia several genera related to Cynopterus i^ is permanently lost. Occurrence of teeth that are normally absent in the species, hut present in related Jorms : — lUj. — Present on one side, in one Cynopterus hrachyotis javanicus ( The tooth is normally lost in Cynopterus, but present in all Fruit-bats with unmodified dental formula. Occurrence of teeth that are normally lost in all living Mcgachiroptera, but no doubt have been present in some extinct form : — jn3_ — Present on one side, in five cases, viz. one Eidolon helvum (B. M., unregistered, specimen c', p. 15), one Rousettus lea/;hi (., one Bousettus seminudus (unregistered, specimen a, p. 39), and two Macroglossus lagochilus lagochilus (, Present on both sides, in four cases, viz. one Eousettus agyptiacus (, one Pteropus gigantcus (, one Macroglossus minimus minimus (, and one Macroglossus lagochilus lagochilus (U.S. National Museum, 125316). \. n]3 is normally lost in all Megaohiroptera, but present in some Microcliir<)ptera. Of the nine cases recorded above of the occurrence of what seems to be an " m' " it is perhaps safest to eliminate the four observed in Macroglossus, owing to the nigh degree of degeneration of the dentition of that genus and the unusually frequent occurrence of supernumerary molars (compare the Marsupial genus Myrmecobius'.). There remain five cases in Eidolon, L'ousiitus, and Pteropus, and in ' view of the fact that all three genera occupy a low position in the suborder it is at least not unlikely that the anomaly is a reversion to a more primitive condition.


PLASTIC CHARACTERS. XXXIU Supermimerary teeth tliat arc certainly, or at least probably, merely accidental outgruwthi : — "13." — In one Pteropus vampynis skull ( a tooth is present, on both sides, behind i^ and leaning against the inner side of the canine. The possibility of this tooth being really an i^ (lost in all Megachiroptera, but preserved in some Microchiroptera) cannot of course be denied, but from the position and general shape of the tooth it appears more likely that it is an accidental outgrowth. i' reduplicated on both sides. — In one Maoroglossus mininuis sohrinus ( ireduplicated on one side. — In one Cynoptenishrachyotisangnlatus ( " P2," /. e. a tooth occupying the position of a ]i^. — Present on one side, in three eases, viz. one Pteropus scapidatus (, one Epomophorus gamhianiis (Berlin Museum, 10171), and one Eonycteris spelcsa (, Present on both sides, in one EpomopJiorus yamhianus ( It would be interesting if this supernumerary premolar really represented P2, a tooth lost in all Chiroptera, and, as in the case of "ia" (abovf), the possibility cannot be altogether denied. But it should be remembered that Pteropus scajndatus and Eonycteris are forms with somewhat degenerated teeth, therefore liable to accidental anomalies in the dentition, and the teeth of Epomophorus gamhianus^, though not exactly .degenerated, are remarkably small for the size of the bat ; further, that the diastema between p, and p^ is in all three forms considerably wider than elsewhere in the lower jaw, so that, if there is any latent tendency in the jaw to a development of a supernumerary tooth, this wide diastema so to speak " invites" it to crop u]) there ; and finally, that in two of the species, viz. Pt. fcopiilatus and E. gainliiairiis, cases are known (see below) of the appearance of a supernumerary premolar between p^ and p^ (compare a Eousettus angolensis with an abnormal tooth squeezed in between m' and m^). Tliis, to say the least, is a warning against too rash homologizations. Supernumerary tooth between p3 and p^. — On one side (tooth small) in one Epamophorus gamhianus ( On both sides (tooth well-developed), in one Pteropus scapulatus ( Supernumerary tooth between m' and m'. — On one side (tooth extremely narrow, abnormal in shape), in one Rousettiis angolensis ( " inj." — On one side, in four cases, viz. one Pteropus gigantens (]06.d), one Mctcror/lossus laqochilus nouns (, two Macroglossiis I. lagochilus (10.3i 23 ; L'.S. National Museum, 125316). On both sides, in two cases, viz. one Macroglossus mininuiS minimus (., one Macroglossus lagochilus nanus ( In no other geuus are anomalies in the cheek-tooth series so frequent as in Macroglossus (see p. 754). 13. Palate-ridges. Eight ridges forming regular curves from side to side, the interdental ridges almost equidistant, the postdental a little more narrowly spaced, and some of the latter slightly notched at the middle, as if tending to split into a right and left half, such is the simplest, and therefore presumably the most primitive, type of palate-ridges known in Megachiroptera (see fig. 28, p. 484, the Epomophorine geuus Pleroten). A similar, more or less slightly, but never profoundly, modified arrangement of the ridges is seen in the large majority of genera of the Rousettine, Cyuopterine. and Macroglossine sections ; the number of the ridges may be a little increased, and, if so, the increase generally takes place chiefly on tlic postdental portion of the palate, the spacing of the ridges may


XXXIV ri.ASTIC CHARACTERS. l)e a little less regular, and some of the posterior ridges mf\y be more distinctly divided at middle (for those details see the descriptions and figures under each genus). But the P^pomophorine is the only section of Fruit-bats in which more essential modifications of the system of ridges occur; in that section, and in that only, nearly every genus has its own peculiar form and arrangement of the palate-ridges, and in some cases {Epomops, Epomojilwrus) even the species may be identified from the characters of their soft palate (or, if this is removed, from the impressions left by some of the heavier ridges on the bony palate). Omitting all details, the modifications of the soft palate in the Epomophorine section of genera are briefly these : — The simple palate-ridges of Plerotes have already been referred to above. In two of the three known species of the related Epomops, viz. E. franqueti and huettil-oferi (figs. 31 A, B, p. 489), the general arrangement is still comparatively little modified, but the three anterior (interdental) ridges are become thick and prominent : in the third species, E. dohsoni (fig. .31 C, same page) a similar thickening of the three anterior ridges has taken place and, in addition to this, also the fourth and fifth ridges are modified, being heavy and triangularly prominent. The jialate of the closely allied Ni/psif/nathus (fig. 34 C, p. 504) differs only in less important details from that of Epomops franqueii. In Epomopiliorus (fig. 37, p. 516) all ridges (six in number, apart from a few thin and inconspicuous ridges at the extreme hinder edge of the palate) are thick and prominent. Micropteropus and Nanonijcteris exhibit the most peculiar modifications of the ridges in the whole suborder. Micropteropus (fig. 39, p. 556) in so far resembles Epomophonts (to which it is most probably closely related) as all the ridges (five) are thick and prominent, but the first ridge is typically hastate in form, with the point directed backward, the second to fifth divided by a deep groove extending along the median line of the palate, very broad in front and gradually n.irrowing posteriorly ; a glance at the palate of this animal is sufiicient to distinguish it from any other Fruit-bat. The latter remark would apply also to Nanonyctcrix (fig. 41, p. 561); the interdental ridges of this genus are rather similar to those of Epomops (probably one of its closest relatives), but the postdenfal ridges are increased in number, narrowly and very regularly spaced, thickened and elevated at middle (forming a j)rominent keel along the median line of the postdental palate), depressed and very thin laterall}; this is almost exactly the reverse of the type of palate-ridges found in Micropteropus, in which the soft palate is marked not with a prominent keel but Avith a dee]) and broad groove along the middle. Scotonyeteris and Casinycteris (fig. 45, p. 571) in so far recall Nanonycteris as the interdental ridges are thickened and the postdental ridges somewhat (Scotonyeteris) or much (Casinycteris) increased in number, but the latter are simple, thin, and serrate.


PLASTIC rnARACTKRS. XXXV 14. Tov'jue. Tlio principal morlifications of the surface structure of the tongue have been described and figured on pp. 723-728, fig. 65. 15. Wing-structaie. Detailed descriptions of the characters of tlie wing, in each genus of Megachiroptera, are given in the systematic part of this Catalogue. The present paragraph intends to give only a general survey of the variahilit}of some of these characters. [In all Fruit-bats there are three phalanges in the second finger, the terminal phalanx nearly always clawed, two phalanges in the third, fourth, and fifth.] ThirtJ, fourth, and fiftli metacarpals. — There is never any very great contrast in the lengths of the metacarpals of the three long fingers. As a rule, however, the third is distinctly the longest, in some genera the fifth, while in others again all three metacarpals are practically subequal. The variations in this respect, within the four primary groups of Fruit-bats, arc briefly these : — In RousettHS and its closest relatives (Eidolon, Boneia) the third metacarpal is as a rule slightly longer than the fourth, which is a little longer than or subequal to the fifth ; the " indices " of the metacarpals (/. e. their lengths for a supposed leugth of forearm of 1000) are in typical llousettus, respectively, 612, 5^5, and 586, in Eidolon 600, 668, and 641, in Boneia 671, 659, and 648. In Dohsonia (indices 621, 568, 586) and the related Harpi/ioni/cteris (697, 661, 673) the third has remained the longest, but the fourth tends to be a little shorter than the fifth. Finally, in Pteropus and its relatives, Acerodon, Pteralope.r, and Styloctenium, the fifth is slightly the longest, the third a little longer than or equal to the fourth (ex., Pteropus Jij/pomelanus with the indices 689, 670, 718, Pteralopex with 690, 659, 708, St>/locteninm with 723, 723, 739). In the Epomophorus section the third is nearly always slightly the longest, the fourth and fifth subequal (or the fifth tending to be the longer of the two) ; Epomophorus with the indices 680, 647, and 641, and Epomops with 728, 699, and 715, may serve as examples. In Scofont/ctcvis (682, 678, 688) and the related Casini/rtcris (692, 683, 692) all three metacarpals are subequal ; and iri Phrotes (679, 698, 689) the fourth tends to be slightly the longest. Again in the Ctjnopterns section the third is nearly always the longest, the fourth and fifth more or less subequal, though very often with a distinct tendency of the fourth to be the shortest, or the third and fifth may be subequal, the fourth slightly the shortest ; indices in Cijnopierus 640, 599, 625, in Bali any cterin 719, 697, 714, in yi/etimene 70S, 646, 678. Two genera of Macror/lossino' (and those two which also in skull and dentition are the least specialized in the subfamily), viz. Eon>/ctrris and Mer/aJoglossus, are nearlv Rousettine in the


XXXVl PLASTIC CHARACTEKS. proportionate length of the metacarpals : third longest, fourth intermediate, fifth shortest ; indices in Eom/cieris 675, 656, 608, in Megaloglossus 760, 716, 670. In all other Macroghsstnce these three metacarpals are either practically subequal in length (Alacror/lossvs and Sycom/cferis, both closely interrelated), or the fifth is distinctly the longest [Melonycteris, Nesonyeteris, Notopteris, all closely interrelated) ; examples, Macrocjlossus 726, 733, 740, Nesonyeteris 760, 755, 798. Phalanges of iJiird, fourth, and fifth fingers. — The second (terminal) phalanx of the third finger is always much longer than the first phalanx, but nearly always (for exceptions see below) decidedly shorter than the metacarpal of the same finger ; the two phalanges of the fourth finger are subequal, but with a very distinct tendency of the terminal phalanx to be the longer ; the two phalanges of the tlfth are subequal, the terminal phalanx being sometimes rather longer, but more often a little shorter than the proximal. In some forms, however, a conspicuous lengthening has taken place of the terminal phalanx of the third finger, making it subequal to or longer than its metacarpal ; this is the casein Eidolon, Slenonycteris (subgenus of liousettus), Boneia, Pleropus, Acerodon, Dohsonia, and Plerotes, and the three closely interrelated Macroglossine genera, Melonycteris, Nesonyeteris, and Notopteris. Subjoined is given in tabular form the absolute minima and maxima of the indices of the metacarpals and phalanges of the three long fingers, and for comparison the actual indices of one of tho shortest-winged (Rousettus) and one of the longest-winged (^Nesovycteris) Fruit-bats.


PLASTIC CUAKACTEUS. (relative length, for a forearm of 1000) of the whole of the third linger (metacarpals and phalanges combined). This index is between : — luOO-1000 : — In typical Eouscttus (i. e. li. amplexicaudatus and allied forms), the shortest-winged Fruit-bats. lUOO-1700: — In the following genera of the Cjjno-plerns section, viz. Cynopteras, Plenochirus, and FentJielor. 1700-1800: — (In the following genera of the liousettus section:) Eidolon (as a rule), Sttnonycteris, Bone'ui, and Dobsonia ; {Epomophorus section :) Eponiophorus, Micropteropus, iScotonycieris, and Casinycteris ; (Cijnopterits section :) Myonycteris, Meycvrops, Dyacopterus, Chiroiuuv, Thooptcrus, and Sphcerias ; {Jlucroylossince :) Eonycteris. 1800-1000 .—{liousettus section :) Lissonycteris, Pteralopex, and JIarpyionycieris ; {EpomopJiurus section :) Flerotes, Eponwps, llyps'ujnathus, and Nanonycteris ; (Cynopterus section:) BaUonycteris ; (Alacroglossince :) Megaloylossus and Macroylossus. 1900-2000 :—(A'ot«e^<((S section:) Pteropns (as a xxAo), Acerodoa (as a rule), and iStyloctenium ; {Cynopterns section :) Nyctimene ; {Macroglossince :) Syconycteris. L'()00-2100 : — In the three Macroglossino genera, Melonycteris, Nesonycteris, and JVotopteris. The table below shows the absolute minima and maxima of the indices of all live lingers, and for comparison the actual indices of one of the shortest-winged and one of the longest-winged Fruit-bats. let linger c. u.


PLASTIC CnAKACTERS. fingers. If hi^i identiiicalions M^ere correct, the indices of the third, fourth, and fifth fingers would be, respectively, 1486, 1450, and 1046, i.e. the third and fourth lingers would l>e very short and practically subequal, the fifth enormously lengthened, a condition unthinkable except on the supposition that the form of the wing of Archcpopteropus was fundamentally different from that of any other bat, living or extinct. It is obvious that what Meschinelli takes to be the fifth finger is the third, and what he considers tlie third is the fifth. With this correction the indices of the metacarpals and phalanges of the three long fingers (calculated from Meschinelli's measurements) are as indicated in the bottom line of the following table : —


PLASTIC CHARACTERS. is not found in any recent Fruit-bat (but tliere are more or less close parallels in ilicrochiroptera). A glance at the first table above shows that the third metacarpal was quite ordinary in length, and the discrepancy due to the greater length of the fourth and litth. The first phalanx of the third finger was normal ; the second, as usual, by tar the longest of all phalanges, and relatively even somewhat longer than usual in Fruit-bats. The two phalanges of the fourth finger were nearly normal, though the first rather shorter than usual. The first phalaux of the fifth finger w^as normal in length ; but there seems to have been two phalanges distally to this (hence Meschinelli"s statement that the "third" finger, which in reality is the fifth, had three phalanges), and, curiously enough, these two distal phalanges together are equal in length to the single distal phalanx of living Megachiroptera (one might almost be tempted tn think whether these " two " distal phalanges are not one broken into two pieces, but it must be admitted that this suggestion is not borne out by the published plate). As a rule in Megachiroptera the fifth finger is distinctly shorter than the fourth, though in a few genera (e. g. Epomops) practically equal to the lourth ; in Archceapteropus it was a trifie longer than the fourth (as in a few Microchiropteia). The general conclusions are : — In so far as the second finger had three phalanges, and its terminal piialanx was undoubtedly clawed, the hand of Archceopteropus was a genuine Megachiropteran hand ; and in so far as the second finger was less reduced in length, the hand of the fossil form may be said to be a little more primitive than that of any living bat : and in the features in which it differed, more or less slightly, from that of living Megachiroptera, it rather approached the hand of some Microchiroptera (except, of course, if it really had a third complete phalanx in the fifth finger). [The teeth of Archteopteropus are very little known, but the molar structure is said to have been cuspidate as in normal Microchiroptera, a statement that cannot be controlled with certainty from the published plate ; the tail was long as normally in Microchiroptera and in one genus of Megachiroptera, Noioj^teris.'] Claw of second fiiKjer. — The claw of the second finger is lost in one (aberrant) genus of the lionseltus section, viz. Do'>sonia, and in three genera of Macroglossinte, Eonj/cteris, Xesoivjcleris, and Aotopteris. Even if tlie claw is absent, the ungual phalanx is always present (rudimentary in Notopteris). Membranes. — The lateral membranes arise as a rule from the flanks or, rather higher up, from the sides of the dorsum. In Fteropm, Acerodon, and Stijloctenlum the line of origin lies generally somewhat nearer toward the spine, and in a few species of I'teropus {),ielunopo(jon, papuanus, ueohihet-nicus) the membranes arise verv close together, almost from the sides of the spinal tract.


Xl PLASTIC CHARACTERS. In Ptcralo^ex (related to Pleropus) they arise almost from the spinal line, or at any rate so close together that the interspace is scarcely appreciable. Finally, in two mutually entirely unrelated genera, Dobsonia {llousettus section) and Notopteris (Macroglossince) the naked membranes are perfectlj' continuous across the back and connected with the integument of the dorsum only along the spinal line. Posteriorly the membranes may be inserted on any toe, from the first to the fifth. The variations, within the primary sections, are these : — On first toe. — Itouseftns section : Eidolon, lioKsettus (pt.), Boneia, and Dobsonia (pt.), that is, in all genera except Pteropus and its closest relatives, and Harpyionycleris ; but in llousettus and Dobsonia the insertion varies between the first and second toe, and in Boneia the insertion is sometimes rather between the first and second toe than on the first. Epomophoms section : only in two genera, Scotonyderis and Casinycteris (in all other genera on second toe). Cynopterus " section : in Cynopterus, Ptenochirus, Meycerops, Balionycteris, PentJiefoi; and Spihcerias (in the other genera on second toe). Macroylossince : in Eonycteris either on first or second toe, or between. On second toe. — Rousettus section : in Pteropus, Acerodon, Pleralope.v (sometimes on first), Styloeteniam, and Harpy ionycter is (perhaps rather between first and second); on Rousettus ?i,x\A Dobsonia compare remarks under first toe. Epomophorus section : all genera, except Scotonycteris and tkisinycteris. Cynopterus section : Myonycteris, Dyacopterus, Chirona.v (?), Thoopierus, and Nyctimene ; in the latter the insertion varies between the second and third toe. 3IacroyJossince: Eonycteris (often on first), MegaloyJossus (sometimes on third), Notopteris (sometimes between first and second). On third toe. — Rousettus section : none. Epomophorus section : none. Cynopterus section : sometimes Nyctimene (see under second toe). JIacroylossime : MegaloyJossus (on second or third), Macroglossus (on third or fourth), Melonyctcris (third or fourth), and Nesonycteris (third or fourth). On fourth toe. — Only \n sovae Macroglossince : Macroglossus {Wnrdi or fourth), Syconycteris (fourth or fifth, rarely between ' third and fourth), Melonycteris (third or fourth), and Nesonycteris (third or fourth). On fifth toe. — In the Macrogiossino genus Syconycteris (fourth or fifth). n The membranes are marked with well-defined yellow spots in Balionycteris and Nyctimene (both of the Cynopterus section) ; more or less obscure spots are detectable in a few other genera, e.g. Eonycteris. The wings are said to be "bright orange" in life in Casinycteris {Epiomophorus section).


I'LASTIC CHAKACTEKS. xli 16. Tail. Varies ;is follows : — (1) Tailequul in length to forearm: — the oligocene ArchcFopterOjxis (8 free caudal vertebnc) ; one genus of living Megachiroptera, of the subfamily Macroglosshue, viz. Notopteris (" 10 vertebne,"' statement taken from literature, not verified by the writer). (2) Tail considerably reduced, subequal to or somewhat shorter than tibia, but longer than hind foot with claws : — one genus of the Cynopterus section, Xycthnene (7 free caudal vertebrae). Though the tail of Nyciimene is distinctly longer than in llonsettus, the number of vertebrae is the same as in Itousettus amplexiauidatus. (3) Tail further reduced in length, from about one-half to the full length of the foot with claws : — (i^owsc^^HS section:) Eidolon (4 free vertebrae), liouseltus (5-7), Boneia, Bobsonia; — {Cynoptenis section :) Myonycteris, Cynopterus (4), Ptenochinis, Dyacopterux, Fenthetor ; — {Macroylossimv :) Eonycferis (7). (4) Tail rudimentary, as a rule reduced to a small knob externally, more easily traceable by touch than by eye : — {Epomophorns section :) PJerotes (tail absent?), Epomops (2 free vertebrae), Epomophorus (2-3), Mkropteropus (3), Kanonycteris, Scotonycteris, Casinycteris; — (Cynopterus section:) Thoopteriis; — (Macroglosshia:) Megaloghssus (5 vcrtebrao, terminal two or three rudimentary), Macroglossus (3 or 2), Syconycteris. (5) External tail absent : — (Bouseti us section :) Pteropus (no free vertebrae), Acerodon, PteraJopex, SiyJoctenitim, Ilarpyionycteris ; — {Epomoplioriis section:) Hypsigmtthus (no free vertebrae); {Cynopterus section :) Megcerops, Balionycteris (no vertebrae), Chironax (no vertebrae), Spluerkis ; — {Macroglossinai :) Melonycteris, Xesonycteris. In the first, second, and third of the stages recorded above the basal portion of the tail is included in (connected by its dorsal integument with) the intcrfemoral, the tip freely projecting (whether or not this was the case also in Archceopteropits is unknown) ; in the fourth stage the tail rudiment is usually unconnected with the intcrfemoral. Two facts are evident from the above, first, that the tail is considerably reduced in length in all living Megachiroptera, with the single exception of JS'otopteris, second, that the degree to which it is reduced varies, as a rule even very conspicuously, within each of the primary sections of the suborder. The Bousettus section falls into three natural subsections, the Rousettus, Pteropus, and Dobsonia subsections ; in the first (Eidolon, Bousettus, Boneia) the tail is on the third stage of reduction, in the second (Pteropus, Acerodon, Pteralopex, Styloctenium) entirely absent, while in the third it is either on the third stage (Dobsonia) or absent ( Harpy ionycteris). In the whole of the Eponiophorus section the tail is rudimentary (fourth stage), except in HypsignatJius (and Plerotes'!), which has no tail. Within the Cynopterxis section, it is on the second stage in Nyetimene only, on (he third in Myonycteris. Cynopterus, Plenochirus, Dyacopterus, and Penthetor, on the fourth in Thoopleri'.!.


Xlii PLASTIC CHAUACiEUS. and absent in Mcyicrops, Balioni/<;tei-is, Chlroncu', and Sphcerias. Finally, in the Macrocjlossinii;, it is on the first stage in Notopteris onh', on the third in Eonifcteris, on the fourth in Alegaloglossus, Mucrojlossus, and SyfonycUris, and absent in Melonycteris and JS'esoni/cteris, these two genera being the closest relatives of the long-tailed JSotojiteris. 17. C'alcar. The calcar is rudimentary in Fhrotes (Ej)omo2^horus section) and Si/coni/cterls (subfamily Mucroghssitux), absent in Sphcerias [Cynoptenis section), the moditication being in all three cases due to a reduction of the lateral interfemoral ; in the two former genera the interfemoral is reduced to a narrow (Flerotes) or even sublinear (Syconycteris) rim along the tibia, in iSpha'rian it terminates at about the middle of the til)ia. 18. CoJour of fur. In the majority of Fruit-bats the prevailing tinges of the coloration of the fur are a dark brown, sometimes inclining toward dark hair-brown *, sometimes closely approaching to bistre, or (not infrcqueutl}') washed with dull olive, the underparts being usually distinctl}' paler than tlie back. To have a basis for comparison this colour type may be considered the " typical " Megachiroptcran coloration, and the principal modifications, exhibited by genera and species differing from this " typical '' colour, may then be referred to one or several of the following categories : — (1) The dark brown tinge may brighten to fawn-brown, cafe-au-lait, fawn-drab or related tinges; or to wood -brown, yellowish, buffy or even cream ; or by increasing admixture of (sprinkling with) greyish hairs to pale hair-brown, ashy drab, or even light silvery grey (for examples see the brief summary of the colour changes in the i)rimary sections of Megachiroptera, below) : (2) The dark brown tinge may darken into seal-brown or blackish, either on the dorsal surface only (many species of Pteropus, some Acerodon), or both on the dorsal and ventral surfaces (some s])ecies of FtcropHs), this latter modification leading in its extreme to complete melanism (Pteropus modigUaiiii., Pt. natalis, some races of Pt. vampyrus ; Pleralopex ; occasionally in Pteropus alecto and Pt. tytleri) ; or a darkening of the underparts may be combined with a lightening of the colour of tlie dorsal surface (some races of Pteropus hypomelanus ; Melonycteris): (3) The fur of the nape of the neck (the " mantle ") may become brighter-coloured, forming a more or less strikingly-coloured (chestnut, russet, tawny, buff, yellowish buff, cream-buff, whitish') "tippet" contrasting with the unmodified or nearly unmodified dark back (many Pteropus, Acerodon) ; or the development of a * Throughout the whole of this Catalogue the colours are named, as far as possible, in accordance with Ridgway's ' Nomenclature of Colore ' (1886).


PLASTIC CHARACTEP.S. xliii poculiurly colouied mantle maybe conibiiied witli any of the coluur changes indicated under (1), (2), and (4): (4) The head may be marked with sharply defined stripes or patches (Ftcrojjus caimtraiiis and personahis, )Stylocteninm, Scoionycteris, Casinycleris) : (5) A small tuft of hair at the anterior and posterior bases of the ear-conch may become pale-coloured, whitish or bufty (eeereingly a trivial modification, but in fact eminently characteristic of all genera of the K2}onwp]iorus section, except one) : (G) A dark spinal stripe may develop (only in Nyctimene, and not equally distinct in all species) : (7) Peculiar light or bright-coloured neck-tufts may develop, the brightening of the colour sometimes spreading across the foreneck and chest and backwaid along the fianks (in males of many species, rarely in females; see theparagraph "Secondary sexual characters," below). In each of the natural sections of Megachiroptera the more noteworthy modifications of the colour of the fur are, briefiy summarized, these: — liousettus section. — Junistftus, Bnnna. and liarjiyionycteris are essentially '-typical'' in colour. In Eidolon {helvum and sabmim) the colours are more or less conspicuously tinged with yellowish (and "more so in females than in males). In Dohsonia the dark brown (Kousettine) general colour is often more or less brightened by admixture of olive, raw-umber, or tawny-olive, these tinges often with an indefinable greenish hue. P^eVojJMS (together with the closely related Acerodon) shows greater colour variations than any other genus of Fruit-bats ; any of the modifications mentioned above under (1) to (4) occur, single or combined; a summary has been given in the systematic part of this Catalogue (pp. 74-75). The t\yo known species of Pteralopex are nearly {aneeps) or quite melanistic (airafa). The single si>ccies o{ Styloc'tniimn is unusually light-coloured (silvery greyish or silvery biiflfy, with sharjjly contrasting dark brown bases to the liairs), and with peculiar head markings, closely similar to those of Fteropus personatus, to which the genus is undoubtedly related. Epomophorus section.— Small whitish hair-lufts at the base of the ears anteriorly and posteriorly are present in all genera of this section, except iScotonyctcrin, but in no other Fruit-bats. The general colour of the fur is in Hypsir/nathus dull, dark ])lumbeous or slate, in all other genera as a rule more or less tending toward tho paler tinges of brown, fawn-brown, cafc-au-lait, brownish russet, brownish Isabella, or allied tinges. "White markings on the head, curiously analogous to those of Pteropus personatus and t'Stylocienium, are present in Scotonycteris and Casinycterix (the appearance of essentially analogous head markings in so widely separated forms as Pteropus personatus and Styhctenium on the one hand, Scofoiiyctei-is and Casinycteria on the other is difficult indeed to account for, except on the supposition that these arc cases of


Xliv PLASTIC CHARACTERS. reversion to a colour pattern of the head of some common ancestral form). Cynopterus section. — Brownish tinges with paler underparts are the rule, except in Nyclimene. Myonycteris is practically llousettine in general colour. In Gynopterus the colour of the back is often suffused with warmer (russet) tinges, sometimes Avith cinnamon, or a tinge between cinnamon and wood-brown, the foreneck, sides of chest, and flanks with chestnut, cinnamon-rufous, or deep hazel (but generally much more so in males than in females). Mego'vops is somewhat paler above, approaching cafe-au-lait, like many Eporaophori ; Sphan-ias nearly greyish hair-brown. Balionycter'is and C7itVo?irtct' are rather darker above than usual, with the head and nape of neck nearly blackish. A well-marked dark brown spinal stripe renders most species of Nyctimene easily distinguishable in colour from all other Fruit-bats ; the stripe is generally narrow (one-fifteenth to one-eighth of the breadth of the back), sometimes obsolescent, sometimes again (^V. ucllo) very broad (one-third of back) ; the general colour of the upperside is sometimes irregularly mottled all over with darker tips to the hairs, as a rule, however, paler than usual in Fruit-bats, fawn-brown, fawn-drab, ashy-drab, wood-brown, buff}', or even cream (the palest tinges seen in females of some species). Macrorilossince. — Eonyctcris is typically Eousettine in colour ; Megaloglossus and Notopteris dark. In Macroglossus and >Sycoiiycteris the colour of the upperside is lighter, varying from warm russet Front's brown to almost pure wood-brown. Melonycteris is approximately cinnamon above, with small white " epaulettes," nearly seal-brown beneath (an unusual contrast) ; the related Nesonycteris differing only by the pale underparts and the absence of " epaulettes." 19. Size. To show their differences in size the Avhole series of genera of Megachiroptera are arranged below according to the lengths of the forearm (that this list can be only approximately correct is a matter of course, some of the genera being as jet known only from one or a few specimens). The smallest Fruit-bats are but little larger than the very smallest Microchiroptera, the largest greatly surpass in size any form of Microchiroptera. The " expanse " of a Fruitbat is roughly about six times the length of the forearm (rather more than less). The extent of individual variation is in any Eat usually at least about ten, often twelve per cent., very rarely as much as fifteen to seventeen ; i. e. if in a sutficiently representative series of individuals, all belonging to one species or subspecies, the forearm of the smallest adult individual measures 50 mm., that of the largest will be about 55-56 or a little more, or if the minimum is 150, the maximum will as a rule not be more than 165, rarely as much as 170.


PLASTIC CPTAKACTERS. xlV 37_ 42 mm.: — BaUonycteris; ? Chironcnv, jVer/alor/losxvit ; Marroqlossiis ; Syconycteris. 4349 mm.: — Nanonycteris ; Scotonycteris ; Chirona.v; ?Sph(frias; Megdloglossus ; Macroglossus ; Syconycteris. 5059 mm. : — Plerotes ; Micropteropns : Nnnonycteris \ Scotonycteris ; ? Caninj/cieris ; Myonycteris ; Cynopterus; MeyoTops ; Penthetor; Sjihcerias ; Nyctlmene ; MeJovycteris ; Nesonycteris ; Notopteris. GO69 mm.: — Eousettux-^ Epomophorus •, Casinycteris ; Myonycteris; Cynnpterus; Penthetor; Nyctimene\ Eonyctcris ; Melonycteris ; Notopteris. 7079 mm. : — liotcscttus ; Dobsonia; EpomopJiorns:, C'ynojiteriis ; Dyacopterus ; llioopterus ; Nyctimene ; Eonycteris. 8089 mm. : — Eousettus; Pteropus; Dobsonia; TImyyionycteris; Epomops; Epiomopilwriis ; Cynoptervs; P/enocJiirus; Nyctimene; ? Eonycteris. 9099 mill. : — Pouseftus ; Boneia ; Pteropus ; Sfyloetemmn ; Epomops ; Epomopliorus ; Cynopierus. 100-109 mm.: — Pteropus; Dobsonia; Epomops. 110-119 mm. : — Eidolon; Pteropus; Dobsonia; ffypsiynatJius. [Archcvopteropus would belong here.] 120-129 mm. : — Eidolon; Pteropus; Dobsonia; Hypsiynathns. 130-139 mm. :—£'?c?o?on; Pteropus; Acerodon; Pteralope.r; Dobsonia ; Hypsignathus. 140-149 mm. : — Pteropus; Acerodon; Pteralope.r; Dobsonia. 150-159 mm. : — Pterojms ; Acerodon; Dobsonia. 160-205 mm.: — Pterojnts; Acerodon. 206-220 mm. -.—Pteropus. 20. Secondary sexual characters. In all or nearly all genera (in so far as both sexes are known) the canines average at least a little heavier and the zygomatic breadth of the skull at least a little greater in males than in females. In several Pteropodino' (some species of Pteropus; Pteralope.r; Penthetor) and the majority of Macroylossince (Macroglossus, Syconycteris, Melonycteris, Nesonycteris, Notopteris) this seems to be tlie only tangible secondary sexual differentiation, and even this difFerenco is in some cases so small as to be only appreciable in largo series of individuals. As a rule, however, the differentiation of the soxos is somewhat more conspicuous, the secondary sexual characters being referable to the following four categories : — (1) Males of a few genera average noticeably larger than females {Epomops, llypsignathus, and Epomophorus ; Eonycteris). (2) The general colour of the fur is in a few species conspicuously lighter in females than in males (some species of Eidolon and Nyctimene) ; or males average poraeNvhat richer in general colour than fomnles (Dobsonia riridis).


xlv PLASTIC CHARACTF.RS. (li) A tiift of hiiir on either side of the nock is in a large niimher of genera distinctly differentiated in the males, being moi'e rigid, unctuous, and brighter (or deeper or more saturated) in colour than the surrounding fur (development of " neck-tufts ") ; or the -whole of the fur across the foreneck and anterior portion of the chest may be more rigid and more saturated in colour in males than in females (development of a " ruff"), or the hair of the same region of males may differ, if not in rigidity, at least in its richer colour from that of females ; or (as is the case in some species of Pterojnis) there may hardly be any sexual difference in the fur of tlie foreneck, but that of the nape of the neck (the " mantle") may be distinctly more rigid in males, and in some of these species palecoloured from tip to base, whereas in females of the same species it is pale-coloured only at the exposed tip, dark-coloured at the concealed base {Fteropvs mariannus and conspic Hiatus groups). In some genera of the Epomopliorus section the " neck-tufts " of tlio males have, so to speak, moved higher np on the sides of the neck, being situated one on each " shoulder," i. e. the region of the sides of the neck immediately in front of the origin of the antebrachial membrane (" shoulder brushes" or "epaulettes,'' hence the name " Epomo-phorus," epaulette-bearer), at the bottom of a deep, pouchlike depression in the skin (" shoulder pouches), each shoulder brush being usually erectile and retractile at the will of the animnl ("when erected the tuft had a vibratory movement," collector's note on the label of a Micropteropus pnsilhis, British Museum). (4) Helatively rarely (Epnmops, Ht/psirpiaiJiiis, Epomopliorus) males differ from females by the possession of one or several pharyngeal air-sacs, a peculiarity very often, perhaps always, combined with an enlargement of the larynx of the males (the voice of Epomops and J/j/psignntJnis (both sexes? or males only?) is described as a loud croaking, that of the latter i-ecalling the croaking of the Ethiopian Giant Frog, liana occipitalis). In one of the same genera {Hi/psignatlms) the muzzle and cranial rostrum are much heavier in males, owing to the development in that sex of a subcutaneous air-sac on either side of the muzzle and large cutaneous folds on the extremity of the muzzle (the latter present, but much smaller, also in females). lleviewed in each of the four natural sections of Megachiroptera the secondary sexual characters are, briefly, these (diiferences in size of canines and in zygomatic breadth omitted): — Rousettus section. — EiJolon : males with richer-coloured necktufts (traceable, but smalhsr and less rich in colour, in females) ; females, at least in two species (7ie?rM)n and sa/>«?Hm), conspicuously V paler than males, liousettus : males as a rule with neck-tufts, the unctuous hair sometimes forming a " ruff " across the foreneck. Boneia: sexual characters unknown (females not on record). Pteropus: fur of mantle often (not always) more rigid and unctuous in males, softer and more spreading in females, in a few species


PLASTIC Cn.VRACTERS. xlvii darker at concealed base in females than in males ; neck-tufts (not conspic-uous, except on spreading the fur) in males of some species. jicerodoti : essentially as Fterojjvs. Pteralopex: no appreciable differentiatio]!. Styloctenium: sexual characters imperfect.l}' known. Dobsonia : as a rule scarcely any appreciable differentiation, in at least one species (D. viridis) males distinctly richer in general colour. Harpy ionycteris: known from one adult specimen (unsexed). Epomophorus section. — Generally characterized by the unusually high development of secondary sexual differences. Plerotes: differentiation unknown (one female). Epomops : males with large shoulder pouches and erectile shoulder brushes ; males with two pairs of pharyngeal sacs (p. 492) ; larynx enlarged in males ; males averaging considerably larger than females. Hypsir/nathits : no shoulder pouches or shoulder brushes ; males with pharyngeal sacs and enlarged larj'nx, as in Epomop>s, and with a pair of subcutaneous rostral air-sacs; cutaneous folds on extremity of muzzle much larger in males ; muzzle and cranial rostrum of males much enlarged and differing in shape from those of females ; malts averaging much larger. Ej^iomophorus : males with large shoulder pouches and erectile shoulder brushes, and with one small central pharyngeal sac ; males averaging larger (character much less pronounced in the smaller than in the larger species). Micropteropvs and Nanonycteris : deep shoulder pouches and erectile brushes in males (females often with small pouches, but no brushes); scarcely any appreciable sexual difference in size. Scoionycicris and Cashiyderis : sexual characters uncertain (adult males unknown). Cyiwpterus section. — Sexual differentiation, if developed, usually Rousettine in character (though often more pronounced). 2Iyo'iiycLeris: males with neck-tufts and " ruff " across foreneck (adult females unknown). Cynoj^terus and Pienochirus: males with nccklufts (small tufts sometimes traceable in females) and often with " ruff." Mi'i/a'rops and Dyaropierus : males unknot n. BaVtonycterix : hair of foreneck brighter in males. Chirona.v r.nd Tliooptirus: adult males unknown. Penlhetor : no sexual difiVieiitiation. Sphccrias: males not seen. Nyctimcne: in some species, males with fur of sides of neck, foreneck, chest, and flanks much brighter in colour ; in other species, scarcely any sexual differentiation in the colour of the underpaits, but females much paler above (cream-buff, cream-white) than males (fawn-brown, ashybrown, ashy-drab). Macroglossi7)a'. — Secondarjsexual differences in most genera undeveloped ; if present, essentially llousettine, thougli in the case of one genus combined with a conspicuous average difference in size. Eonyciervi : males with a well-defined "ruff" across the foreneck of generally deeper (more saturated) colour, and averaging noticeably larger. Megalcylossrts'. males with large (whitish) neck-tufts. Macroglossns, Syconycteri/i, Melonycteris, ycsonycferis, and Aofoptervi : no conspicuous sexual differentiation.


Xlviii INTERRELATIONS OF GENERA. III. Interrelations of the genera of Megachiroptera. It has been considered desirable to give here a summary of the mutual affinities and probable phylogen)' of the genera, " sections," and subfamilies of living Fruit-bats, omitting as far as possible all discussion of details. For these latter the reader is referred to the paragraph " Affinities " under each genus, in the systematic part of this Catalogue. Rousettus (in its typical form, as represented by the species of the subgenus l^ousdUn-) is one of the least specialized genera of living Megachiroptera. The rostrum is moderate in length and iu no respect peculiarly modified, the premaxillse in simple contact with each other in front (not ankylosed together, nor spaced), the lateral margins of the postdental palate forming straight lines converging backward, the postorbital processes are short, the postorbital foramina present, and the brain-case perfectly unmodified in general shajje. On the other hand, that all the most primitive cranial characters found in any living Fruit-bat should be united in the skull of this or any other living genus, is hardly to be expected; it is not difficult, therefore, to point out a few single skull characters in which Eouseitus stands a little higher than one or another genus of Megachiroptera : as in the large majority of Fruit-bats the infraorbital canal is quite short (considerably longer in MeJonijcteris, Nesonycteris, and Noto^^tens), and the ascending branches of the premaxilla3 as narrow at their upper extremities as near the alveolar margin (much less narrowed above in the three genera just mentioned); the palate is perhaps rather too broad to be considered quite unmodified in form, the tympanic bones, though typically annular, are rather broader than in most other genera, and the facial portion of the skull is distinctly, though not very strongly, deflected against the basicranial axis (more so than in the majority of Epomophorine and Cynopterine genera ; but it should be noticed that the deflection of the facial axis often varies to some extent in undoubtedly closely interrelated genera, and sometimes even in species of one genus : see anted, p, xxiii). The dentalformula is unmodified Megachiropteran (^ — ^ incisors, j. cheekteeth), the teeth in no way peculiar, except in so far as the inner cusp of p' and p^ is completely fused with the outer, and that of p^ uearly so, characters shared with a large number of other genera. The palate-ridges are arranged in regular and almost equidistant curves over the whole of the palate, some of the posterior ridges always distinctly interrupted iu the middle; the tongue papilUe essentially unmodified (there is a faint tendency to a lengthening of the conical papillae at the extremity of the tongue : compare Macroglossina). A short tail is present, the second finger clawed, the wing-membranes inserted on the preaxial side of the foot, the prevailing colours of the fur a dark tinge of hair-brown or brown, the secondary sexual differentiation inconspicuous (males as


IXTKRRELATIOXS OF GKNKRA. xlix a rule witli small, often more or less concealed, necktufts), and the general size moderate. The genus has divided into three branches (subgenera), Ronsetlu.', Sfenoiujclcris, and Lissom/cteris. The subgenus Eovsdhis (ten species) ranges over the Ethiopian and Oriental regions (extending from the former northward to Egypt, Palestine, and Cyprus) and the whole of the Austro-Malayan subregion, while SUnonycteris and Lissonjfcferis are confined to tropical Africa. In Stenonycteris (two species) the cheek-toeth are become narrow, m^ is reduced in size, and the facial axis more strongly deflected than in Eousettus s. str. Lissotij/cieris is somewhat more aberrant : the brain-case is peculiarly flattened posteriorly and the facial axis even less deflected than in Eousettvs, hoth characters giving the skull, viewed in profile, a rather striking resemblance to that of Epomops ; the prema.xillfe arc ankylosed together anteriorly, the postdental palate rejatively longer, the cheek-teeth shorter and broader (subsquarish) with the ridges more cusp-like (shorter antoro-posteriorly and higher vertically), those of p, separated, m' and p, reduced in size. Eidolon (three species, Ethiopian and jralagasy regions) has originated from a HouseUits-hke bat. p' is not quite so much reduced as in Roitseftus, but in other respects Eidolon is a little more peculiar : the premaxill?c are spaced in front, the postdental palate more expanded laterally, the tympanic bone elongated to form a short bony auditory meatus (a character unique among Bats),_ m, longer (antero-posteriorly) than usual, the sexual differentiation rather more pronounced (males always with neck-tufts, females often conspicuously paler than males), and the general size of the animals larger. Boneia (one species, Celebes) is closely related to liouKettu.t, but more specialized. The lower canines are slanted strongly outward, this necessitating a greater breadth of the palate between the upper canines, this again making the rostrum conspicuously broader in front; perhaps as a further consequence the premaxiila; are spaced anteriorly ; but in all other respects the skull is psscntinlly Rousettine in sha]ie. i' has disappeared, i^ is somewhat increased in size, the molariform teeth flatter, the outer and inner cusps of Pj separated and those of p' and p^ less completely fused than in liovseftus. The external characters are practically unmodified. Pteropus, the largest genus of Megachiroptera (eighty-five species, a hundred and three forms) and one of the most widely distributed, has originated from a bat essentially similar to Rousettus, but slightly more primitive than the living species of that genus, in so far as it must have had a narrower palate, the tympanic ring thinner, and the cusps of p^ and p^ distinctly separated. The dentition is in all typical sjiecies of the genus considerably heavier than in liovseifus, hence the skull much more heavily built, the crests stronger, the postorbital processes lono^er, d


1 IXTERREfATIOXS OF GENERA. tl>o coronoid process of the mandible broader and more steeply ascending, i^ is always distinctly smaller than i^ (and in some species, chieHy of the Fterojms lomhocensis group, nearly rudimentary), p^ usually deciduous, m' and ni^ somewhat reduced ; the occipital (sublambdoid) jiortion of tlie skull is more elongaie, vubtubular. The tail has entirely disapjiearcd, the colour's of the fur are mucli more varied (that of the nnpe of the neck, the " mantle," generally conspicuously brighter than the back), and the size of the animnls often greatly increased, tlio sjiecies varying in size from that of a .Fieldfare (or a medium-sized Jlousettus) to that of a Haven. Pteropus is distributed over nearly the whole area inhabited bj^ Megachiroptera, with the important exception of the African continent, a fact all the more remarkable inasmuch as the genus is relatively richly represented in the wliole of the Malagas}' region and occurring also in tiio island of Pemba, south of Zanzibar. The only claim of Acerodon (six species, nine forms: western Austro-Mitlaya and Philippine Islands) to stand as a genus distinct from Pteropus is the rather liigher specialization of its dentition : p'' and m' (sometimes also p^ and p.,) have develo])ed a well-defined antero-internal tubercle, and the lower molariform teeth (p^, m,, and m^) a sharply defined inner basal ledge; more or less distinct approximations to similar modifications are, however, seen in some specialized forms of Pteropus (see p. 415). The principal characters of Pteralopex (two species : Solomon Island^) are the excessively heavy dentition, the shortened, snbsipiarish form of the upjier molariform teeth, the prominent anterior and posterior basal ledges of the same teeth, the bicnspidato outer lidge of the lower molariform teeth, the thick upper canines with a heavy external secondary cusp and small inner basal cusps, and the very large outer and small inner lower incisor. As pointed out in detail elsewhere (p. 436) all these peculiarities represent, in fact, onl}^ the last phase of modifications exhibited, more or less in an initial stage, by certain species of the Pteropus ^^sflaphon group (2'>sel'jpJwn, pilosus, tuhercuJatu?, and leucopterus). That Pternlope.v lias developed from a bat closely related to the living species of that group, or is, in other words, the peculiarly modified Solomon Islands representative of that group, is scarcely open to doubt. From another branch of Pteropus has developed the genus Styloctenium (one species: Celebes): i, and m^ have disappeared, and the colour of the fur is pale with dark bases to the hairs and sharply defined head markings,— all characters so distinctly foreshadowed in, or closely apy)roached by, the living species of the Pteropus iemmincki and lomhocensis groups as to indicate, almost with certainty, tlie origin of Sti/locienium from a form closely allied to the living species of those groups of Pteropus (for a discussion of details, see p. 444). At some, probably not very remote, period of the growth of the Eousetto-Pteropine main branch of Megachiroptera a bat must


IXTKRKKr.MrOXS OF GJKN-ERA. j; have oxisfod which, though essentially similar tc a Pta-opu, i„ tlie Renenil appearance of the skull (palate narrower than in liou.ettn, occiput rather more » subtubular " than in nonsettu. but less so than in Ptcropus, tympanic ring thin) and with the whole series of cheek-teeth unmodified Eousettine or Pteropine in number ("_ •'*) nnd structure, differed in the following important dental characters'rtnnlT'l ''"'r' .r''' '^'1^"J' P'-°^'i^-°^i-^ the lower canines Mtnated close together at the extremity of the mandible, tl>e nner pair of incisors above and below fi> and i, ) had disappeared, the Mngl. pair of lower incisors (i^) was quite small, the sinc^e pair of upper incisors (i^) somewhat modified in shape (crown obliquely triangular or obliquely bilobed, owing to it beini actid ^.pon not by ,,, which was wedged in between the closely app'ox ! mated lower canines and much too small to reach i=, but by the tip ot the lower canines), and, as consequences of these modifi.'ations o the front teeth, the premaxilhx. were con iderablv reduced ,n breadth (sublinear) ; externally this bat must have beef, ^cry hke a liovsettvs: tail i>resent (at about the length of l.e foot), second finger clawed, wings from sides of back aiK n.scrted posteriorly on first or second toe, colour of fur some dark IS 1 iT^Vn" ^^"-^^«r"' ^'^-^ P'-obably as a small lionsett..,. If not smaller still. From this bat have developed in one direction mZ:^. ^^"^^'•°-^^^'«^-)' -^ --t^.or//.,,,,/j„,,.,,,.(^MS!n: ^.iL^t^Tt r^^^'K^ '^'"'''' ^''"^'"" ^«'-"^^^'> the following further modifications have taken place: the small p> (which i^ sometimes deciduous in Eouseltru., and usually so in Fteropm) ha, entirely disappeared (cheek-teeth I), the wings arise from the spinal lino of the back, which is therefore completely covered bv the naked membranes, and the claw of the second finger has been lost' T. the most primitive species ot the genus (D. vvuor, Xew Guinea) the molar structure is typically Eousettine, but in all other spec cs it^ IS more or less complicated, by the development of posterior basal ledges antero-internal basal ledges, median surfacl ridges or by a tendency of the outer and inner ridges to break up into two or more cusps (for details see pp. 450-452 and fi9.5) ^ tvne ffltnlT^^^r"^ ^™"' ^^' above-described su^p^sed protot pe Harpyionycteris (one species) has been modified as followstt.e upper and lower canines and upper ir.cisor are stron^lv proc ivous (upper and lower canines crossing each other at neaHy r ghfc angles), tlie premaxilla, solidly fused anteriorly with each other and laterally with the maxilla^ an outer secondary cusp ha developed m the upper and lower canines, and a small inner^.usn m the lower canines (which therefore are tricuspidate), the number of cheek-teeth is unchanged (^: p' present), but the tendency in n^Inf rr'\°^^^'^'^-^^^^ (•"«« ^'>*-^-''«'«^ above) toward a developme^nt of posterior and anterior basal ledges and of surface cusL and to a splitting of the outer and inne^ ridges into two or more


lii INTOKRELATIOXS OF G15XERA. cusps has been carried to an extreme, so as to render the molariform teeth raulticuspidate (for a discussion of the homologies of the cusps see pp. 801-803 and fig. 79). Externally, the onljnoteworthy modification is the loss of the tail (wing-membranes normal, claw of second finger present). The nine genera reviewed above constitute what may be termed the Ronsettine section of Megachiroptera, in contradistinction to the Epomophorine, Cynopterine, and Macroglossine sections. Harpyionycteris PCeralopsx Acerodon Dobsonia Fig. II. — Interrelations of the gcner.a of the Eousetti/s section. It has divided into three branches (.sub.sections); the Rousettine branch {Ronsettus, Eidolon, Boneia), characterized by the simple, unmodified cranial characters (rostrum never shortened, premaxillae not sublinear, occiput not elongated, &c.), by the full Megachiropteran dental formula (except for the loss of i' in Boneia, and the occasional deciduousness of p'), the simple form of the premolars and molars, and externally by the presence of a tail and by having nearly always the third metacarpal slightly but distinctly longer than the fourth and fifth ; the Pteropine branch {Pterojius, Acerodon, Pteralopex, Stylo cteniuni) : cranial characters Rousettine, except for the more " subtubular " occiput and relatively narrower


INTERRELAXIONS OF GENEEA. liii {jalute, dental formula unmodified (except for the loss of i, and m^ in ISti/lucteuinni), molar structure sometimes simple, but more often sliowing some, though rarely {Pieralope.c) a high, degree of specialization, tail absent, and fifth metacarpal uearl}' always slightly but distinctly longer than third and fourth ; and the Dobsonian branch {Dohsonia, Harpy ionycteris) : rostrum at least somewhat shortened, premaxillse reduced in breadth, lower canines situated close together at the extremily of the mandible, i' and i^ lost, molariform teeth with a pronounced tendency to a high degree of specialization, tail present or absent, of the three long metacarpals the third nearly always distinctly the longest, the fourth the shortest, the tifih intermediate. The Kousettine branch (eighteen species) has spread over the whole area inhabited by Megachiroptera, except Polynesia; the Pteropine branch (ninetj'-four species, a hundred and lifteen forms) covers the whole of the same area, including Polynesia, bat excluding the continent of Africa and the Eastern Medite:ranean countries ; while the Dobsonian branch (thirteen species, fourteen forms) is confined to Austro-ilalaya and the Philippines. The probable mutual affinities of the genera of the Rousettine section are expressed in the diagram (fig. II.) on the foregoing page. The Epomophorine genus Plerotes (one species : Ethiopian) probably originated from a primitive Uousettus-\\]\Q type. Except lor the less of m'its dental formula is typically Megachiropteran (iu all other Epomophorine bats not only m", but also p' and m^ are lost, and, as shown in fig. 28, p. 4S4, even in Plerotes these two teeth are so small as to be nearly functionless), and its remarkably simple palate-ridges are easily derived from those of Rousettus or rather from a form in this respect somewhat more primitive than the living representatives of that genus (see fig. 29, p. 485). But in other respects Plerotes is highly specialized : the palate is unusually broad, the cheek-teeth greatly reduced in breadth (almost Macroglossino), the molars and last premolar flattened, with scarcely a trace of the usual cusp-like elevations, the lower canines slanted outward, and the tail and calcar absent or rudimentary. Somewhat similar lines of development have been followed by the related genus Epomops (three species, four forms : Ethiopian). Like many, if not all, other Epomophori it subsists chiefly on soft juicy fruits, the contents of which it draws out rather by suction than by mastication, and its lips, pharynx, larynx, cranial rostrum, dentition, and palate-ridges have been modified accordingly. The lips are full, pendulous, and highly expansible; the pharynx long, wide, and greatly extensible, communicating with the oral cavity by a very restricted aperture, the larynx spacious, with ossified walls, and supported behind by the expanded hyoids, acting as an " exhauster" during the suction; the bony palate is broad (though not broadened to the same extent as in Plerotes), the interdental palate-ridges thick and prominent (fig. 31, p. 489), the canines and


liv INTEERKLATIOXS OF GEXEHA. ])' aud p, tliiii and .sharply pointed, adapted to piercing the rind of the fruit and keeping it in position, while it is squeezed between the jaws and pressed upward against the interdental paLite-ridges ; the other cheek-teeth are ndatively feeble, p\ m'% and m^ have disajipeared ; r is often deciduous. Hypsignatlius (one species: Ethiopian) is undoubtedly derived from a form very similar to Ejioinopx, but probably with the postdental pahite narrower. The rostrum is greatly enlarged, particularly in depth (and much more so in males than in females), the lower canines slanted more outward, p, more reduced, the ridges of the molariform teeth higher and their cusps more narrowly pointed, the outer ridges of some of the lower teeth more or less distinctly bior tricuspidate, the iiremaxillae aukylosed together in front, the muzzle thick aud truncate (' Hamuierheaded Bats"), the u])per lip with prominent integumentary folds, the palate-ridges (fig. 34, p. 504) with relatively slight moditicatiuns similar to those of The origin of JS'anonifctei-is, Scotoiii/cieris, and Casiiij/rteris (all monotvpic and Ethiopian) is perhajis not (luitc clear ; that they are rather closely allied to each other there cannot be much doubt, and the probability is that this small branch is an offshoot from a type related to, but less specialized than, Pleroies and Ejwnwps. In all three genera tlie rostrum is considerably shortened, in Scotoni/cferis and Cusinycteris almost to the same degree as in Cynoptems. and the dental formula is in all tl e same as in Epomops (clieek-teeth ^ : p', m-. and m, lost). They differ from each other chiefly in the form of the bony jialate. In Nanonycteris the postzygomatic palate is unusually short and broad (fig. 40, p. /iGO), in Scotonycteris of normal length, with the lateral margins converging backward in straight lines (fig. 42, p. 564), while Casinycteris exhibits the unique ])eculiarity of having no postdeutal palate, the mcsopterygoid fossa extending forward very nearly to the level of the last molar (fig. 43, p. 569). The palate-iidgcs, though specially modified in each genus (see p]). 561, 565, 571), are without difficulty derived from an Epomnps-MkQ pattern. Like Epomops, Epomophorus (eight species, nine forms : Ethiopian) feeds chiefly on soft fruits, but in having adapted themselves to this diet the two genera have followed to a certain extent different lines of development. In Epomops the rostrum and palate are broadened, in Epomophoms long and narrow ; in Epomops the postdental palate is broad and flat, i. e. essentially Eousettine in shape, in Epomophorus it is more or less deeply depressed posteriorly, with prominent palation rim ; in typical Epomops the anterior interdental palate-ridges are thick aud prominent, the postdental ridges simple (figs. 31 A, B, p. 48!J), in Epomophorus the postdental are practically similar to the interdental ridges, all being thick and prominent (fig. 37, p. 516). The dental formula and the characters of the teeth are the same in both genera.


i>:Tnr.RELATio>'s oi GI:^•EK.^. Iv Micropteropus (one species : Ethiopian) has jn-obabl) originated from a type related to, but riiore primitive than, tiie living species of Epomojjhoriis (wiliunit the lengthening and narrowing of the rostrum characteristic of these). As in Ejiomophorus the postdental palate is distinctly depressed posteriorly, with raised palation rim ; the palate-ridges are thick and prominent, but separated by a deep and broad groove along the median line of the palate (lig. iJ'J, p. 55G); the dental formula as in the majority of Epomophori ([>', m% and m^ lost) ; but the rostrum is as short and heavy as in Ci/nopterus, the interdental palate broad posteriorly, and the postdental palate abruptly narrowed behind the anterior root of the zygomatic arches (fig. 38, p. 555). The eight genera reviewed above (Plcrotcs, Epomops, Ihjpsi(jnathiis, JVanoni/cieris, Scotonijcte'ris, Caslinjcteris, Epohiopliorus, and Micropteropus) constitute the Epomophorine section of Eruit-bats. The principal differential characters, as compared with the llousettine section, may be briefly summed up as follows : — Dentition on the whole weak, p\ m'-^, and m^ lost in all genera except Plerotes, which has retained p' and m^ in a rudimentary condition ; molar structure perfectly simple, except for the nearly total degeneration of all surface structure in Plerotes and the splitting of some of the ridges in Ihipsignutlins ; number of incisors unmodified in all genera (i — "'), except for the deciduousness of i" in Epomops\ facial axis only very little deflected against the basicranial axis (except in Plerotes) ; brain-case distinctly flattened posteriorly (the same ]>eculiarity is, however, seen in one type of the Rouscttine section, Lissoni/cteris) ; form of postdental jjalate highly variable (in the whole series of Megachiro])tera this is the only section that shows any great variation in this portion of the skull) ; palate-ridges simple only in Plerotes, in all other genera more or less highly specialized (the only section of Megachiroptera showing any great modification of the surface structure of the soft palate, in fact the only one in which every genus may be identified with certainty only by an examination of the palate-ridges) ; tail eitlier reduced to an inconspicuous rudiment, more easily delectable by the touch than by the eye, and not connected with the interfemoral, or absent ; small tufts of white hair present at anterior and posterior bases of the earconch in all genera, except Scotoni/cteris ; secondary sexual characters often unusually highly developed : males with shoulder pouches and erectile light-coloured" shoulder tufts (" epaulettes '' ; exception : ]/;/psignathus), sometimes with pharyngeal sacs (Epomops, Ihipsii/iiathits, EpomnphorHs). and not infrequently averaging considerably larger than females (Epomops, IJifpsigna'tJms, several species of Epomophorvs). The range of the section is strictly confined to the Ethioj)ian region. Only Epomophorus is distributed over nearly the whole of this region ; "the other seven genera are inhabitants of the " West African Province " (approximately synonymous with the Great West African Eorest Tract), from the Guinea Coast east


Ivi IXTEKRKLATIONS OF GKNERA. to, or iu the case of one or a tew species a little be3'oiid, Victoria ]Syun/,a, south to Angola. Eeiiguela, and Damaralaud. The genera of this section fall into three natural groups (branches, subsections) : — (1) The Epomops branch, tlie genera Plerotes, Epomops, and Hiiptsignathus (five species, six forms) : rostrum long, palate broad, postdental palate simple, at least some of the postdental palate-ridges unmoditied (except in Epomops dobsoni) ; (2) the Nanonycteris branch, including JS'anom/cteris, ticofom/cteris, and C'f(sint/cleris (three species) : rostrum much shortened, postdental palate highly variable, though never as in the third branch, at least some of the postdental palate-ridges unmoaitied ; (3) the Epomophorus branch, the genera Epumophorus and Micropteropus (nine species, ten forms) : postdental i)alate depressed posteriorly, rostrum varying in length, all palate-ridges modified. The probable mutual affinities of the genera of the Eporaophoriue section are expressed in the subjoined diagram (fig. III.). Micropteropus Casinyctens Epomophorus Plerote ScoConycceris Fig. III. — Interrelations of the genera of the Epomophoms section. Myonycteris (four species: West African Province) has in many respects remained on the P^ousettine level of development, while in others it exhibits modifications approaching to those of Cynoptertis. The general external appearance, the dental formula, and the palate-


ISTEUKI LAI IONS OF OENEKA. Ivii ridges are (luitc or nearly as in Huuseltus, but tlie rostrum is conspicuously shortened, the facial axis less detiected, ni^ and m" (last lower and upper moLir) leduced almost to rudiments, the orbits larger, the nostrils more piomiuent, and the calcar weaker. One species, inluibiting the island of San Thome, Gnlf of Guinea, is a little peculiar in some details of its dentition and ought to stand as a distinct subgenus (F/n/(jelis, p. 57'.*, fig. 47). Cynopterus (six species, si.xteen forms: Oriental region, east to Celebes and Timor) is further modilied : the rostrum is still more shortened, the facial axis more nearly horizontal, m^ and m" have disappeared, more or less distinct surface cusps are often developed in pj and m, (see p. 589 and fig. 49), the upper and lower canines have develo])cd a small supplementary cusp on the inner edge, the papillie on the inner side of the lips are increased both in number and siiic, the palate-ridges more crowded, thicker, and more sharply lirojecling, the nostrils more })roniinent, the tail a little more reduced, the colours of the fur more inclining toward the brighter tinges of brown, and the neck-tuttsof the males more conspicuou&ly differentiated. Ptenochirus (one species : rhilii))iiiie Islands) is an only slightly modilied offshoot from the CipiojjiiVKs tj'pe (showing j)articularly close atfinities to the " J\'ia

Iviii IXTEKRELATIONS OF GliXERA. nulimciitary state : in all other Cynoptcriiie genera both m., atul m" sire lodt. lu the mere dental fornnila, therefore, Bnlioni/cteris has remained on a slightly more primitive stage than all other Cyuopterine bats, except Mi/onycferis, but in other respects it is peculiarly specialized: i^ is lost, i" shortened, m' and m, somewhat reduced, some of the lower teeth (p^ and m^) subscpiarish, p^ has developed an antero-external basal lobe, the ixjstorbital foramina liave disajipeared, the tail is absent, the wings longer than usual and marked with sharj)ly defined yellowish spots, and the general size of the animal is unusually small. Chironax (one species) is the Javan representative of the Bornean Balioin/cteris. The two genera are strikingly alike in nearh^ all characters of the skull, in the general aspect of the dentition, as well as in external aj)pearance, "nhile at the same time some of the peculiar characters of Chironax are too important to allow it to be included in Bulionycteris : — -Balioni/cteris has retained a small m", in Chironax this tooth is lost ; in Balionycteris i^ is lost and i" shortened, in Chironax i, is present and i'' normal in length ; Balionycteris has developed an antero-external lobe in p", in Chironax the lobe is represented by a well-defined cusp ; in Balionycteris the premaxilla? are in simple contact anteriorly, in Chironax ankylosed together ; and the wing-membranes of Chironax are probably unspotted, i'rom this it is not ditficult to suggest the characters of the common ancestor of these two genera ; it must, have been a bafc very similar to Chironax, but with a small m'' and the premaxillffi in simple contact; in Java it developed into Chironax (m^ lost, premaxillae solidly fused), in Borneo into Balionycteris {nf retained, i, lost, i" shortened, the antero-external cusp of p^ enlarged into a lobe, wings spotted). The two closely interrelated monotypic genera TJtoopterus and Fenthetor may be presumed to have originated from a bat similar to Cynopterus, with the same number (^ — -) of incisors, the same number of cheek-teeth (t : m^ and m" lost), the same length of the tail, and the wing-membranes inserted on the first toe ; but the raolariform teeth, particularly p^ and m,, have been unusually broad, quadrate in outline, and their inner ridge very low, the upper canines grooved, both upi)er and lower canines without secondary cusps, there was no trace of surface cusps in any cheek-tooth, and the postorbital foramina had disappeared. This form developed in Western Austro-Malaya into Thoopterus : incisors unmodified, jij and ra^ with large surface cusfis, tail rudimentary, wings from second toe; and in Indo-Malaya into Penthetor : i, suppressed, i* shortened, no surface cusps in any cheek-tooth, tail of normal length, wings from first toe. Sphserias (one species : Burma) is probably derived from a primitive " Cynopterus," i. e. a bat with the dental formula and other characters essentially as in C ynoj>ter us, hnt without secondary cusps in the canines and without surface cusps in the lower cheekteeth ; hut it is in certain respects more aberrant than any of the


IXXKRUELATiOXS 01' tiCXERA. lix foregoing Cynopteriiie genera. The premaxilla? are proclivons, tlie iiicitrobably as a further consequence of the extraordinarydevelopment of the anterior iiarcs, the posterior narial jiassage and mesopterygoid fossa became broader and deeper than usual (hence the slightly "pandurate" outline of the postdental palate). The wings are spotted with yellow (as in BaHoiii/cteris). The posterior circumvallate papilla is divided into two (fig. Go A, p. 725), and the cardiac portion of the stomach indistinctly differentiated (compare the alleged insectivorous habits of the genus). (Fur a more detailed discussion of the characters and afSuities of this genus see pp. 6yi-(i94.) The eleven foregoing genera constitute what may be called tho Cynopterins sectioa of Fruit-bats (thirtj'-one species, forty-one forms). The principal characters that link these genera together as a natural group are these : — ( 1) the rostrum is always conspicuously shortened, in all genera (except Mi/onj/cferis and Sphcerias) so much so that the distance from the orbit to the cxtrcmitv of the na?als is


iX INXEKKELATIONS OF GENEliA. con:>ideiaLly less than the breadtli of the rostrum across the lachrymal foramina ; parallels or close approximations are found only in certain genera of the Epomophorine section (Nanonyctens branch and MkfojiUropus) and in some species of Bobsonia : (2) the facial axis of the skull is in all genera (excejtt Myomjcteris and Sjjhcerius) only very slightly deflected ; a similar inconspicuous deflection of the face is seen in tlie majority of E])omojihori, whereas in the llousettiue and Macroglossine sections the lace is always somewhat, and often much, moi'e deflected : (3) in Myumjcteris the lust molar lielow and above (m^ and m") are reduced almost to rudiments, in Balioni/cteris m^ is lost, in all other genera both m., and m", reducing the cheek-tooth formula from the typical ilegachiropteraii J. to -, a formula very rarely found outside this section of Fruit-bnts (it occurs in one species of Sifconycteris, and in Notoptcris, but in the latter case owing not to the loss of m^ and mbut of p' and p,) : (4) only Mijonyderishaa preserved the Rousettine aspect of the soft palate, in all other genera (in so far as the palate-ridges arekno«n) the surface structure of the soft palate is remarkably uniform, the ridges being prominent and crowded ; an exactly similar aspect of the palate is not found outside this section : (5) in all genera (except j\Ji/oni/cteris and Sphcericis) the odontoid papllte on the inner side of the lips are more numerous, larger, and more crowded than in any other section. There is a marked tendency in this section to — (1) a siipjiression of the inner pair of lower incisors (i^, which if present is always distinctly smaller than i ) combined with a reduction of the outer •2 Jl -1 -2 pair of upper incisors (i") ; the incisor formula . — occurs iu a few other genera {Styloctenium, Ncsonycteris, Notoptens), but generally without a conspicuous reduction of i': (2) a develoj)ment of surface cusps in p^ and m^ [Oytwpterus, Ptenoilnrus, and Dyacopterus, but not in the related Meyarops ; in Thoopterus, but not in the related Fentlutor) ; a similar tendency is shown only bj' Bobsonia and Harpyionycteris : (3) to a disappearance of the postorbital foramen (present and well developed in Myonycter'is, Cynopterus, Ptcnochirus, and Meyarops ; rudimentary in Byacopterus ; absent in Balionycteris, Chironax, 2'hoopteriis, PentJietor, Sj'hcerias, and usually in Nyctimene) ; the foramina are present and welldeveloped in all other Megachiroptera witliout exception: (4) to a shortening or complete suppression of the tail. lixcept for Myonytteris, which is Ethiopian (West African), the section is confined to the Oriental region and Austro-Malaya, one genus [Syctimene) extending from the latter subregion to Australia. The question whether this section, so far as our present knowledge of the genera goes, is perfectly homogeneous, i. c. includes only genera which are more closely related to each other than to any genus outside the group, must, in the opinion of the writer, be answered in the affirmative, but it is quite possible that some systematists would be inclined to consider the status of Myonyctcris


INTEI!P-FT,ATIONS OF GTiXFRA. Ixi a little doTibtful. The fact is tliat, this genus has retained nian^characters of lioiiseUns, while in practically all the features in which it differs from liousetttis it more or less closely approaches to Cynopterus. Whether a genus exhibiting characters of this description ought, in a linear arrangement, to be classed at or near the end of the Rousettine section or as the "opening" genus of the Cynopterine section, must necessarily remain a matter of opinion. With regard to the principles for the arrangement of the genera, only a few remarks are required. Putting for a moment J/yonycteris and Nyctimene at one side, it would be easy to divide all the remaining nine genera of this section into two groups according to the number of incisors (|— | or ^— ^ ) ; but. as pointed out elsewhere (p. 649, footnote), an arrangement based primarily on Splisenas Nyctimene [Rousenus section ] Fig. IV.— Interrelations of the genera of the Cynopterus section. this character would be thoroughly artificial ; it would separate Ptenochirns and Megrn'ops from their closest relative, Cynopterus; Bdlionycteris from its Javan representative, Chironcuv ; and Penthttor from its eastern representative, Thoopterns; there is no doubt that the loss of i^ is a character developed independently in difl'erent branches of the section. The diagram above (fig. IV.) is based, therefore, on the following considerations : — .1///oh//W<'/-/

Ixii INTKI'.KKLATIONS OF G TN KIIA. may at once be se])arate(l, owiiij; to its less slioitcnorl roslruin and the presence of in,, and m" ; next the two closely interrelated genera Baliouj/cteyia and Cliironax, the former of which has f)rcserved m" ; as a tliird branch Nyctimene, and as a fourth Sphcpriax, on account of their peculiar specializations in diH'orent directions ; the remaining six tjenera may bo divided into two groups, those with {C'l/nopterus, Ptenochirus, 3Ier/a;rops, Dyacoptenis) and those without jiostorbital foramina (TJiooptcrus, PentJufor), Dyacoptenis being in this respect intermediate (foramina present but minute). All the genera reviewed above, the Housel tine, Eporaophorine, anil Cynopterine sections, form together a Subfamily ot Megachiroptera, the Pteropodinge. The characters of this subfamily, in contradistinction to tlie second primary subdivision of Megacliiroptera, the subfamily Macroglossinse, have been fully discussed in the systematic part of this Catalogue (i)p. 723-728, fig. 65), and it is sufficient here to emphasize the fact that the only real difference between the two subfamilies lies in the greater specialization of the tongue and the tongue papillie in the Macroglosshuv (adaptation to a diet consisting, at least partly, of pollen). Eonyctsris (three species : Indo-China, Indo-^falaya, Celebes) is on the whole the least specialized genus of JJacroylossinre. In the general characters of the skull, dentition, and palate-ridges, as well as in external appearance (except for the loss of the claw of the index), it is remarkably like lioHsetiihi, but the cheek-teeth are narrower than in typical liousettus, m., and m" somewhat reduced (m, absent in one species), the tongue is, of course, typicall}' Macroglossine, there is scarcely any trace of odontoid papillae on the inner side of the lips, and the difference in length l)etween the third (longest) and fifth metacarpals is greater than in llouseUus. Megaloglossus (one species: Ethiopian) is undoubted]}' derived from an Eonycteris-W^Q type, but is considerably more advanced in specialization. The molars are become low and linear, but p^, ])\ and p., are practically unreduced in height (fig. 71 B, p. 74P), the rostrum longer and slenderer, the premaxillai ])roclivous and ankylosed together in front, the extremity of the mandible elongated, the tongue somewhat lengthened, and the tail rudimentary (in some individuals hardly traceable externally). Macroglossus (two species, six forms : Indo-China, IndojVFalaya, Austro-iLalaya) is the eastern representative of the Ethiopian Meyalor/lossus. It has evidently originated from a typo essentially similar to lUer/aloijlosnHS, but with p\ m^, and m^ less reduced in size; and it has carried the specialization considerably fiirtlier than Mfr/aloylossus. All cheek-teeth (p*, p\ and p., not excepted) are become low and linear (fig. 71 C, p. 749), the premaxilla) are even more proclivous, the mandible even more lengthened anteriorly, the mandibular symphysis longer, the facial axis much more strongly deflected, the third metacarpal shortened and


INTF,Rlti:t,ATIONS OK GEXERA. Ixiii the fiitli lengthoiieJ (bo as to mako the thir.l, fourth, and liCth metacarpals ec^ual in length), and the caudal vertebra; further reduced in number. Syconycteris (three species, seven forms : Austro-Malaya and Australia) is ilosely related to Macrotjlossus, but all upj)er and the outer lower incisors are cons])icuously enlarged and proclivons, the posterior molar above and below reduced (absent in one species), the interfemoral unusually narrow, and the calcar rudimentary. In the foregoing four genera of Macror/lossiiia', which rna}' be called tlie Eonycterine section, as well as in all Ptero/)odin(v, the infraorbital canal is short (the outer wall a narrow bridge of bono and its anterior aperture, the infraorbital for;imen, situated vertically below or closely in front of the orbit), and the premaxillte are not or only very little broader above (at nasals) than below (near alveolar border). In the remaining three genera of Macrofflossinc^, the Notopterine section {Meloni/cferis, Ncsom/cten'^, and iVotopteris), the infraorbital canal is much loss reduced, the infraorljital foramen therefore situated a considerable distance in front of the orbit, and the premaxilla? are less narrowed in their upper halves, being about thrive or twice as broad above as below ; and the species of one of these genera (To^o^j^er/s) are the only living Fruit-bats in which the tail has remained long. Externally these two primary sections of Macroglossina' may easily be distinguished by an examination of the wings: in the Eonycterine section either the third metacarpal is distinctly longer than the fourth and fifth or these three metacarpals are subequal in length, and the terminal phalanx of the third finger is always conspicuously shorter than the third metacarpal ; in the Notopterine section the fifth metacarpal is the longest, and the terminal phalanx of the third finger subequal to or even longer than the third metacarpal. In point of dentition Melonycteris (one species: Xew Guinea RTid Bismarck Archipelago) is the least modified genus of the Notopterine section. The dental formula is typically Megachiropteran (incisors r, — ;„ cheek-teeth "!), all the cheek-teeth are much reduced in size and sublincar in form. The external characters are not peculiar, except in so far as the tail has disappeared (and the ventral surface of the body is much darker in colour than the dorsal). Nesouycteris (one species : Solomon Islands) is the slightly modified eastern representative of MeloiufcWris : i, is suppressed and the claw of the second finger is lost (the ventral surface of the body, as usual in Jlegachiroptera, paler than the dorsal). In having retained a long tail, much longer than in any other Fruit-bat, Notopteris (two sjiecies : Western Polynesia) is more primitive than Mrlonj/cterii and Nesoi^ijcliris, in other respects it has to a certain point followed similar lines of development, but is considerably more highly specialized, p' and p. arc lost (the latter


Ixir INTFRRF,T,ATI0N9 OF GKiNFRA. tooth ia in the two related genera quite small, the former nidimentary and almost functionless), p^is enlarged and moved forward flosely behind the canines, i^ is lost (as in Nesoyiyctcris), and i' is deciduous ; the extremity (premaxillary portion) of the rostrum is elongated, the corresponding portion of the lower jaw peculiarly hroadened, and the premaxillse solidly united in front ; the claw of the second finger has disappeared (as in Nesonycteris) and even the claw phalanx is reduced in size, and the wings arise from the spinal line of the back, which is therefore covered by the naked membranes. The Macror/lossimv (seven genera, with twelve sjiecies or twenty forms) are essentially Austroand Indo-Malayan in distribution ; one genus (Macror/lossvs) extends westward into the Indo-Chinese subregion, another {Notopterls) is peculiar to Western Polynesia, a third (Si^coin/cteris) extends southward to Australia ; and one genus {Megaloglossus) is confined to West Africa. The subjoined diagram (fig. V.) illustrates the probable interrelations of the genera. V.' o iB Macroglossus Syconyctsris u §. Megaloglossus Notopteris Fio. V. — Interrelations of the genera of tlie subfamily MacrogloasiiKP. As pointed out above, this subfamily falls into two natural sections, the Eonyctcrine and Notopterine. Tlie less shortened infraorbital canal and broad upper extremities of the premaxillas in all Notopterine genera, and the ]iresence of a long tail in one genus of the same section, all primitive characf.ers lost in all other Fruitbats, are evidence that the Macrof/Iossime as a whole, in spite of the often very high specialization in other directions of the living genera, are of slightly lower origin than the Pteropodincp (see fig. VI., next page).


GEOGEAPHICAL DISTRIBUTION. Ixv Macroglossms Pieropodidse Fig. VI. — General view of the interrelations of the s

Ixvi GKOGEAPIIICAL DISTP.IBtTTION. Rouse fills section. Eidolon (3) Kousettiis (14) ] 7 Boneia (1) Pteropus (85, 103) Aoerodon (tj, 9) ... Pteralopex (2) Stvlocteniiiin (1) ... Dobsonia(12, 13)... Harpyionjcteris (1) Epo'inaphorus section. Plei-otes(l) Eponiops (3, 4) Hypsigimthiis (1) Epomopborus (8, 9) Micropteropus (1) .... Nanouycteris (1) Scotonycteris (1) Casinycteris (1) Cynopterns section. Myonyeteris (4) Cynopterns ((i, 16) .. PtenoeliiruB (1) Megserop.s (1) Dyacoptenis ( 1 ) Balionycteris (I) Ohironax (1) Thoopterus (1) Penthetor(l) Splijerias (1) Nyctiuiene (13) MacroghssiiKS. Eonycteris (3) Megaloglossus (1) Macroglossus (2, fi) Syconyctoris (3, 7) Melonycteris (1) ... Nesonycteris (1) ... Notopteris (2)


GEOGRAPHICAL DISXRIBUTIOX. Ixvii The number of genera, species, and forms represented in each region and subregiou is as follows : — E. Mediterranean subregion ... Etiiiopian region Malagasy rejiioii Iiiflian & Ceyloiiese siibregions , ludo-Cliinese subregiion.-itro.''i(s (Gambia to Monbuttu). J'jiij/iiiijjhonis wahfhet-gi halih'miuu* (Canieroojis to G. & B. East Africa) ; (;n/?(A;Vrw?^< (Senegal to Sennaar and Abyssinia); angulciufis* (Benguela, Damaraland); puuaarffuefi * (Upper Shari). Mici-op/erupiis * ptiHllus (Gambia to Victoria Nyanza). yeroons). Mi/o'ii/cfcris* wrimt/h/oni (Welle district); lepfodon (Sierra I^eone, Lil)eria) ; turquata (Lower Congo, Angola) ; braelii/cejtkala (Sau Thome). Mi'tjaloglossus * woermanni (Congo to Liberia). Eastern side of Continent : — Eidolon hclviiiu (Ethiopian region generally). liouscitus leachi* (B. East Africa to Cape Colony) ; rP(7.w7V?V?f«(Erylhrra, south-west to Gaboon and Loanda, norlli t(> Egypt, Syria, &c.) ; Kcrnpi * (Shoa, B. East Africa).


Ixviii GEOGRAPHICAL DISTRIBUTION. Epomaphorws waklbergi wahlbergi * (B. East Africa to Cape Colony) ; lahiatus * (Sennaar, Abyssinia, Shoa) ; minor * (Shoa to G. East Africa) ; anurm * (Abyssinia aud Erytlirea to Tanganyika) ; ci-ypturus * (Nyasa to Transvaal); yambianus (Sennaar and Abyssinia, west to Guinea Coast). South Arabia : — Eidolon sahcsiMii *. Uoiisetfus arahicus (also Karachi). Peuiba Island : — Ptcropui ooeltzkuwi *. Malagasy Eegion. Madagascar ; — Eidolon dupreannm *. Pteropus rufus* nifus (northern and central) ; riifiis priiiceps (southern). Comoro Islands : — Pterojms comorensis * ; liningdnnei *. Aldabra : — Ptcrojius aldabrensis *. Seychelles : — Pterojms seychettensis *. Mauritius, Reunion : — Pteropus suhiiger * ; niger * ; ? roilricensis. Rodriguez : — Pteroptis rodricensis * (perhaps also Mauritius). Indian and Ceylonese Subrkgioks. Maldives : — Pteropus ariel*. Ceylon : — Rousettus seminudus *. Pteropus giganteus giga7}teiis (also Indian Peninsula). Cynopterus sphinx sphinx (also Indian Peninsula); brachyofis ceylonensis*. Indian Peninsula : — Rousettus arabicus (Karachi ; also S. Arabia) ; leschenaulti (also IndoChina). Pteropus gigantetis giganteus (also Ceylsn). Cynopterus sphinx sphinx {^. &E. provinces; also Ceylon, Assam, N. Burma, N. Siam) ; sphinx gangeticus (N.W. & C. Provinces). Indo-Chinese Scbregion. Himalayas (Kooloo, Nepal, Darjeeling, Assam, Cachar, Manipur) : — Rousettus leschenaulti (also Indian Peninsula, Burma, &c.). Pteropus giganteus leucocephalus *. Cynopterus sphinx sphinx (Assam ; also Indian Peninsula, N. Buraia, N. Siam) ; brachyotis angulatus (also Burma, Siam, &c.). 'i Macroglossm minimus sobrinus ("Darjeeling"; also Malay Peninsula, &c. ; perhaps Siam, Burma, Tenasserim). Burma : — Rousettus lescheimidti (also Indian Peninsula, Siam, &c.). Cyno'pterus sphinx sj>hinx (N.Burma; also Indian Peninsula, N. Siam); brachyotis angulatus (also Assam, Siam, &c.). Spharias * blanfordi. Eonycieris spelaa (also Tenasserim, Siam, Malay Peninsula,

GEOGRAPHICAL DISTEIBUTION. Ixix Mergui Archipelago ; — Ptcropus ki/pomelanus geMinorum *. South China (Amoy) : — Boiisettus Icschenaulti (also Siain, Burma, Nepal, Indian Peninsula). South Liu-kiu Islands : — Piernpus dasymalhts * ; loochoensis *. Formosa : — Pteropus formosics *. Siani, Cambodja, Saigon, Pulo Condor: — Buuscf/us ksvhenaulti (Slam ; also S. China, Burma, Nepal, Inijiau Peninsula) ; ainplcccicaudiitus (Cambodja ; also Indo-Malaya). Pteropus hypomelamis condorensis * (Siam, P. Condor) ; It/lei* . Ci/nopteruA sphinx sphinx (also N. Burma, Indian Peninsula) ; hrachyofis angulatus (also Burma, Assam, Malay Peninsula, &c.). Eonyctcris spelcea (also Burma, Teuasseriw, Malay Peninsula, &c.). ? Macroglossus minimus sobrinus (also Malay Peninsula ; perhaps Buiuia, Tenasserim, Darjeeling). InDO-MaLAYAN SUBREOION. Andamaus : — Pteropus safyrns* (Naroondam); tytleri*. Cynoptcrus brachyotis irruchysoma *. Kicobars: — Pteropus faunulus * ; melanotus *. Cynopterus brachyotis scherzeri *. Lower Siam : — 1 Bouset/us amplexicaudafus (Indo-Malaya generally, Cambodja). ? Pteropus intermedius (also Tenasserim). Cynopterus brachyotis angulatus (also Siam, Burma, Malay Peninsula, &c.) ; brachyotis brachyotis (also Malay Peninsula, Borneo, Sumatra, &c.). Eonycteris speltea (also Siam, Burma, Malay Peninsula, itc). 1 Macroglossus minimus sobrinus (also Malay Peninsula, perhaps Siam). Malay Peninsula : — t Eousettus amplexicaudatvs (Indo-Malaya generally, Cambodja). Ptcropus va/iipyrus malaccennis (also Sumatra). Cynopterus brachyotis angulatus (also Lower Siam, Borneo, Sumatra, &c.) ; brachyotis brachyotis (also Lower Siam, Borneo, Sumatra, &c.) ; harpax *. Megarops ecaudatus (also Sumatra, Borneo). Penthetor lucasi (also Ehio Archipelago, Borneo). Eonycteris spelcea (also Siam, Burma, Sumatra, &c.). Macroglossus minimus minimus (?) (also Java, perhaps Sumatra); minimus sobriniis (also Sumatra and Java; perhaps Lower Siam, &c.). Sembilan Islands (off west coast of Malay Peninsula) : — Pteropus hypomelanus robinsoni *. Sumatra (incl. Rhio-Linga Archipelago. Banka): — h'ouseltus amplcxicaudatus (Indo-Malaya generally). Pteropus vampyrus malaccinsis (also IMulay Peninsula). Cyivrpterus sphinx titthcecheilus (also Java, Lombok) ; Irrachyotis angulatus (also Malay Peninsula, Sinialu, Mentawei) ; brachyotis brachyotis (also Malay Peninsula, Borneo, &e.); horsfieldi lyoni*. Mcgierops ecaudatus (also Malay Peninsula, Borneo). Penthetor lucasi (Khio Archipelago, so far not recorded from Sumatra ; also in Malay Peninsula, Borneo). Eonycteris spelcea(also Malay Peninsula, Siam, Java, &c.). Macroglossus minimus minimus (?) (also Java, perhaps Malay Peninsula) ; minimus sobrinus (also Malay Peninsula, Java). Simalu Islands ; — Cynopterus brachyotis angulatus (also Mentawei, Sumatra, &c.).


IXX GEOGRAPHICAI. DISTRIBUTION. Nias ;— Pteropiig iriddieics *. Ci/!iopf(ri(s hruchyo is miiinlus * ; major * ; princejK *. MacnH/loasus minimus sobrinus (also yuniatra, Java). Mentawei Iftlixncls : — ? Plerapiis ht/pomehnics oiffwius (;ilso Engatio). Cyiiopfcrus hrachi/u/is anyidaius (also 8iiiialu, Sumatra, &c.). Engano : — Bousei/us amplexicttiidafiis (Indo-M:ilaya geiTerallr). Pierojms fiypomelaiius eiiyanns (*) (pei-liii)i3 also Mentawei) ; modiz/Hcrnii *. C'lii'istnuis Island (S. of Java) ; — P/cropus yiatalis*. Norib Nattmas : — Pferopns hypcmelunus cnniix * ; reinipi/riis natunee (also Borneo). ? Cynoptcriis brackyotis brachyotis (ulso Malay Peninsula, Borneo, Sumatra, &c.). South Natimas ; — Pteropus hypomelanics annccfcns *. [Pteropus vampynis natuucs, not yet recorded.] Tambelan Islands, Pnlo Aor, Tiomari, Lantingii, Great Redang, Perbentiau : — Ptcropns hypomelaiiits lepidus*. Borneo (incl. Lnbiian, Mengalim, Sibufii, Lamboyan Islands) : — Jiouseiins amjilcricandafus (Cambodja, IndoMalaya generally)Pteropus hypomelamis fomesi* ; speciosus (Sibutu I. ; also Malanipa I., ofT Zamboanga) ; vampyrus naiuniE (also N. Natunas). Cynopta-us brnchyotis braehyotis (also Malay Peninsula, Sumatra, Philippines, CelKbes). HJeqcsrops efav.datns (also Malay Peninsula, Sumatra). ByacirphTits * spcidicens. llalionycti:ris * lunculaia. Peiithefor lucid (also Malay Peninsula, Rliio Archipelago). Eonycteris major *. MacroyloSfJis higvchilus lagochilus (also Philippines, Celebes, &e.). Mata Siri l.slands : — Cynapienis bracliyoiis ivsuhtruiii (also ICangean Islands). Java (inol. Madura) : — lioitscitiis aJwrtridgei * ; minor *. Pleropus vampyriis vampyrus*. Cyiiopierus sphinx titihcechciius (also Sumatra, Louibok) ; brachyo/is javaniciis * ; hori^fieldi liorsficldi *. Chironax * mdauocephali(s. Eonycteris speUea (also Sumatra, Malay Peninsula, Siam, Burmn). 31acrog/,o<'sus miHimus minimus {nhn Kangean Is.; perhaps Sumatra and Malay Peninsula); minimus sobrimts (also Sumatra, Malay Peiiiusula ; perhaps Indo-China). Eawean Islands: — Ptcropiis atcrrivms (also Kangean Is.). Kangean Islands :^ Pteropus aierrimus (also Bawean Is.). Cyuoptertis brachyotis insulariim ^als(> Mata Siri Is.). Macroglosstis minitnus minimus (also Java, &c.). B;ili : — Pteropm vampyrus pi utnn (also Lombok). Piiilippines (incl. Cagayan Sulu and Palmas Is.): — Jioiiseiius amphxicaudaius (Indo-Malaya generally), Pteropus hypomelanus caqai/avvs * ; sp?ciosus (Malanipa I., off Zamboanga ; also Sibutu I., off N. Borneo); w/?««s (also Celebes) ; puiuilus* (Palmas Is.); leucopferus* (Luzon); vampyrus lancnsis* . '' Acerodoii jubotus* jubatus (Luzon, Leyt^, Negros, Dlnagat, ? Panay) ; jubatus mindanensis {'M\\\i\a\VM-); Incifrr* (Paimy).


GF.ofiRArnicAi, DrsTRinuxroN. Ixxi Hai-pf/ioiii/ctci-L'i * whiteheadi (Jlindanao) Ci/nopterus braahi/ofis hrackyolis (also Borneo, Celebes, kc). J fci.'ocliirus *jnguri. Macrof/lossus lugochiliis (also Borneo, Celebes, &c.). ,, , , AuSTKO-MaI.AVAN SuilKEGlON. ( elebcs : — Riiusettits celebensis (also Sangliir). Buiieia * hidens. Pteroptis hypomelanua i)iaca.i$nricus (also Sanghir, Talaut) ; huiiihs (also Piiilippines) ; doiisuni'^ ; pfCAW/rf/z/s (also Gilolo and Amboina groups) ; alccto (also Salayer. LomSok). Accrodon cclehcnsis (also ijalayer, Sula Is.). Styloctcniam * w lUarci. Dolisonia cxolcta *. Ci/nopterus hrachyoih hrachynlis (also Borneo, riiilippiiies, kc). TliooptiTiis iiifjresccns (also Gilolo group). yyctimetie minutus* ; cephaloles (also Auiboina group, Timor Laut, Timor). fhnycteris rosenhcrgi *. Macrog/iissiiJi lagochilits lagochilus (also Borneo, riiilippines, Sanghir, Aniboina group). Sangliir Islands : — Ji'ousefti/s celchensis (also Celebes). r/rropiis /i.ypome/anit.f macassaricus (also Celebes, Talaut) ; rankeps (also Gilolo group, Sula Is.); ? mdanopttgon (also Amboina group, Banda Is., Timor Laut) ; ? chrysoprocius (also Amboina group). Macroghfsits lagochilus lagocliUus (also Celebes, Philippines, Avnboina group, &c.). Talaut Islands (Lirong) : — P/eropiis hypomclanus macassar tens (also Sanghii-, Celebes). Accrodon htimilis *. Bonerato, Dyampea, Salayer : — I'Icropiui griscus (also Timor) ; alecfo (also Celebes, Lombok). Accrodon celchensis (also Celebes, Sula Is.). liOmbok : — Picrnpits Inmhoccnsis (also Flores); vampi/rus pfiilon (also Bali); nlccfo (also Salayer, Celebes). Cynopterus sphinx titthmcheilitii (also Java. Sumatra). Flores : — Kouseltus amplexieauddfus (Indo-Malaya generally). P/eropiis lomhocensis (also Lombok). AcerodoH iiirm/clo/ijlore/sii *. Dohsoniu pcroiti (also Timor, Alor, Wetter). Sumba : — Ptcropits morio (also SavuJi Accrodon gilrus *. Dijhsonia ssuuihana *. Savn : — Rouftttuf. amplcxicaiidatiis (also Flores, Alor, Timor, Inilo-AIalaya). Pteropiis vampyrus edulis (also Timor) ; morio (also Siuuba). Timor (incl. Wetter): — Jiouscttits amplcxicaudutus (also Flores, Saru, Alor, Iiido-AIalaya^. Picropux gri.-tciis (also Bonerato, Dyampea); tenimiihcki (also Amboina group) ; vampyrns edulis (also Savu). Acrodon mnckloli inacklofi *. Djbsonia moluccensis (?) (also Amboina group, Aru Is.) ; pcroni (also Flores , Alor).


Ixxii GEOGRAPHICAL DISTRIBUTION. ? Cyiioptenis sphinx tiiihacheilus (also Lombok, Jaxa, Sumatra). Nyctime'iie cephaloies (also Celebes, Aiuboina group, Timor Laut). Alor (Ombay) : — Eousettus amplexicaudatus (also Flores, Savu, Timor, Indo-Malaya). Pieropus soli/arius *. Acerodon mackloti alorensis *. Dobsonia peroni (also Flores, Wetter, Timor). Gilolo group (Morotai, Gilolo, Ternate, Batchian): — Pieropus hyponielanus hypomelanus * ; caniceps (also Snla Is., Sangliir) ; persmuitus (also Celebes, Amboiua group) ; chiysauchen (also Gliebi, Salawati, Mysol, N.W. New Guinea). Dobsonia crentdata *. 2'hooptenis nigrescens (also Celebes). Nyciimene albiventer *. Sula Islands : — Pieropus canice-ps (also Gilolo group, Sangliir). Acerodon celebensia (also Celebes, Salayer). Amboina group (Buru, Amboina, Cerani, Goram, Manawolba, Watubella) : — Eouseiius hrachyotis (also New Guinea, &o.). Pieropus Hops*; "i argentaius* ; melanopogon (also Banda Is., Timor Laut, probably Sanghir) ; chrysoproctus {*) (perhaps also Sanghir) ; temmincJci (also Tiuior) ; personatus (also Celebes, Gilolo group) ; ocularis *. Dobsonia moluccensis (also Aru Is., perLaps Timor); viridis umbrosa (also Banda Is.). Nyciime^ie varius * ; cephafotes (also Celebes, Timor Laut, Timor). Macroglossus lagochihis lagocJiilus (also Banda Is., Celebes, Philippines, &c.). Syconycteris crassa major *, Banda Islands : — Pieropus pallidus * ; melanopogon (also Amboina group, Timor Laut, probably Sanghir). Dobsonia viridis umbrosa (also Amboina group). Macroglossus lagochilus logochilus (also Amboina group, Sanghir, Celebes, &c.). Timor Laut: — Pieropus melanopogon (also Amboina group, Banda Is., &c.). Kyctimetie cephaloies (also Timor, Amboina group, Celebes). Key Islands ; — Pieropus keyensis *. Dobsonia viridis viridis *. [D. moluccensis not recorded, but possibly Occurring ; known from Amboiua group and Aru Is.] Nyciimene pajmanus (also New Guiuea, &c.). [Macroglossus lagochihis names not recorded, but possibly occurring ; known from Aru Is., New Guinea, &c.] Syconycteris crassa keyensis *. Aru Islands : — Pieropus aruensis * ; wacroiis *. Dobsonia moluccensis (also Amboina group). Macroglossus lagochilus nanus (also New Guinea, &c.). Syconycieris crassa papuana (also New Guinea). Ghebi, Salawati, Mysol, Waigeou : — Piei-opus chrysnuchen (also Gilolo group, N.W. New Guipea) ; 1 papuunus (also New Guinea). Dobsonia magna (also New Guinea). Nyciimene a'ello (also New Guinea). Macroglossus lagochilus nanus (also New Guinea, &c.).


GKOGKAPHICAL DISTKIBUTION. IxXlii New Guinea : — Bousettus bra<;hyofis (also Amboina group, Bismarck Arch., Solomon Is.). Ptero/ms hi/pomekinus luteus (also islands off S.E. New Guinea) ; chrysauchen (N.W. New Guinea; also Mysol, Salawati, Ghebi, Gilolo group); conapicilkifus (S.E. New Guinea with satellite islands, N. Australia) ; papuanus(*) (perhaps also Ghebi, Mvsol) ; epularius*. Dohsonia minor* ; magna (also My.sol, Waigeou). Nt/ctimene papuanus (also Admiralty Is., Key Is., Cape Yort) ; cyclotis * (Arfak Mts.) ; ccrfans* ; yeminus (also Trobriand and D'Entrecasteaux Is.) ; aello (also Mysol). Macroglossns lagochilus nanus (also Mysol, Bismarck Arch., Aru Is.). Syeonycteris crassa papuana (also Aru Is.). Alelonycieris melanops (also Bismarck Arch.). Conflict, Trobriand, D'Entrecasteaux, Alcester Is., Louisiades: — Pieropus kypomclainis luteus (also New Guinea) ; conspicillaius (also S.E. New Guinea, N. Australia); ? poliocephalus (also Australia). Dobsonia panuietensis *. Nyctinicne gemmus (also New Guinea). Syeonycteris crassa crassa *. Woodlark Island ; — Pteropus hypomelamis Inteus (also Trobriand and Conflict Is., New Guinea) ; conspicillatus (also Alcester and Trobriand Is., New Guinea, N. Australia). Nyctimene lullulce *. Syeonycteris naias *. Murray Islands (Torres Straits) : — Macroglossns lagochilus pygmaus*. Admiralty Islands : — Pteropus admiralitatum *. Nyctimcne papuanus (also New Guinea, &c.). Bismarck Archipelago : — Botisettus hrachyotis (also New Guinea, &c.). Pteropus capistrafus * ; neoliihernicus* . Dobsonia pradatrix *. Nyctimene major *. Macroglossus lagochilus nanus (also New Guinea, &c.). Syeonycteris crassa finschi *. Melonycteris melanwps (also New Guinea). Solomon Islands (whole Megachiroptera fauna) : — Bousettus brachyotis (also Guadaloanar, Bismarck Arcb., New Guinea). Ptercrpus colmius * ; solomonis * : cognafus * ; rayneri * ; ruhianus * ; lavellanus * ; grandis * ; ivoodfordi *. Pteralopex * anceps ; atrata. Dohsonia. inermis * ; nesea *. Nyctimene scitulus *. Macroglossns lagochilus yniavtus*. Nesonycteris * woodfordi. Known distribution in the separate groups of Soloiuon Islands; — Bougainrille, Shortland, Fauro : — Bousettus brachyotis (also Guadalcanar). Pteropus coloiius * ; grandis*. pteralopex aiiceps*. Dohsonia n^sea (also Eubiana). Nyctimene scitulus (also New Georgia, Florida, Guadalcanar). Nesonycteris woodfordi (also Guadalcanar). Vella I^vella : — Pteropus lavellanus*.


Ixxiv GEOGKAnnCAL DlSTRinUTION. Gliizo, Riibiana, New Georgia : — Fteropus iolomoHis* ; rubianus * ; woorfjunh (also Giiachilcaiiar). Bohsunia iiesea (also Sliortlaiid). Nyctimene scitulus (also Sliortlancl, Florida, Giiadalcauar). Florida, Giiadalcauar: — Emisettits hmchyotis (also Fauro). I'teropus rayneri * ; uoodfordi (also New Georgia). I'teralopex atrata *. yyctimene scitulus (also Shortland, New Georgia). Macroglossus layachilus microtiis '^. Nesonycteris woodfurdi (also 8horlland, Fauro). San Christoyal, Ugi : — Fteropus coynutus *. iJohsonia incrmis*. Australian Continent. ricrajiiis briinncns* ; youldi* ; conspiciUatus (also S.E. New Guinea and adjacent islands) ; i)olwccplialus (*) (perhaps also Trobriand Is.) ; scapula! us *. JS'yciimene pctpuanus (also New Guinea, &(;.) ; robbtsoni *. ,»7' t<)jconijctcrU australis*. Polynesian Subregios. Bonin and Volcano Islands : — Fteropus pselaphon *. Mariannes ; — Fteropus mariaunus*. I'elew Islands : — Fteropus pelcwcnsis * ; pilosus *. Carolines (all islands togetlier) : — Fteropus yapeiisis * ; ualanns * ; moloss/nus * ; insnlaris * ; ph *. 1 Nutoptcris macdomddl (also New Hebrides and Fijis). Records from the separate islands : — Yap, Mackenzie : — Fteropus yapensis *. Kuck Atoll : — Fteropus insularis *. Mortlock : — Fteropus molossinus (also Ponape) ; pluEoccplialua *. Ponape : — Fteropus molossinus (also Mortlock). TJalan (Kushai) : — Fteropus ualemus *. Sta. Cruz Islands (Vanikoro) : — Fteropus vanikoreusis * : ? tuherculatus * (there is Some uncertainty about the liabitat of this species). New Caledonia; — Fteropus ornatus * ; ycddici (also New Hebrides). Xotojjtcris neocaledonica *. Loyalty Islands : — Fteropus auratus *. New Hebrides (Aneiteuni) : — Fteropus ycddici (also New Caledonia) ; anetiauui^*. Notoptcris macdonaldi (also Fijis). Fijis: — Fteropus tovyanus (also Tontras, Sanioas) ; nanairusis *. Notoptcris macdonaldi (also New Hebrides). Tongas ; — Fteropus ioiiyanus (also Fijis, Sanioas). Saiuoas; — Ffcmpus ionyaiuis (also Fijis, Tongai-) ; famohiMS*.


GEOGRArillCAL DISTKlBtTTIOX. IXXV 3. Jiemarls on the geoi/rcqjlucal distrihation. The northern limits of the area known to be inhabited by Megachiroptcra are Seneg:ambia, Egypt, Cyprus, Syria, South Arabia, Karachi (one unverified record of I'teropus girjanteus from Kelat), Himalayas, Amoy, Formosa, South Liu-kiu Islands, Bonins, Pelew Islands, and Mariannes ; in I'olynesia they occur as far east as the Samoa Islands. No Pruit-bat is known from North Africa west of Egypt and north of Senegal, nor from Asia Minor, Persia, Japan proper (records in literature of Pteropus dosi/mallus from Kiushiu and Hondo probably refer to captive specimens), Hawaii, New Zealand, and Tasmania (one ])robahly erroneous record of riero/ms poJiocephalus). The only fossil Fruit-bat thus far described is Archcopteroptis irmisiens, from the Upper Oligocene of Italy C' llousettus" (/aillardi, from the Middle Miocene of France, was, judging from 'the published figure of its humerus, a species of Microchiro])tera). Of the four primary sections of Jfegachiroptera, one, the Epomo]i7iorHS section (8 genera, 17 species, 19 forms), is strictly confined to the Ethiopian region ; a second, the Cjinopterus sectiou (11 genera, 31 species, 41 forms), extends from India and Ceylon in the west to the Solomon Islands and Australia in the east, and is represented bv one genus {Mijonyctevls, -4 species) in West Africa ; a third, the Sfacroqlossina' (7 genera, 13 species, iil forms), ranges from'Indo-China eastward to the Fiji Islands, and reoccurs, in a single genus and species (Mer/ahr/lossvs), in West Africa ; the fourth, the liouseltus section (9 genera, V2o species, 1-17 forms), in the number of species more than twice as large as the three other sections together, covers the whole of the area inhabited by Megachiroptera, from West Africa east to the Samoa Islands. A few genera have a very wide distribution : Ihnsettits (l-i species) from West Africa to the Solomon Islands (but not to Australia), Flernpus (So species, 103 forms, the largest of all genera, in the number of species nearly equal to all the other genera together) from the island of Pemba (south of Zanzibar), through the Malagasy, Oriental, and Australian regions to the Samoa Islands; others have a moderately wide range, e. g. Macroghssus from IndoChina to the Solomon Islands (not to Australia), Chinopteras from India and Ceylon to Celebes. But for the mnjority of genera the geographical "limits are much narrower, and not a few (chiefly monotypic genera) are so far known only from one place, island, or group of islands : Spharias from Burma only, Dtiacopterus and Balioni/clfiris from Borneo, Chironax from Java, Uarpi/ioni/cteris and Pienochints from the Philippines, Boncia and ^Slglocieniiim from Celebes, Pteralopex and Nesoivjrteris from the Solomon Islands : all I'lpomophorine genera, c:s.ce\>th: porno phorus, as well as Mi/onifderis and Megnhglossvs are practically confined to the whole or part of the great West African Forest Tract. The range of the species is, of course, as a rule restricted within much narrower limits, or if a species is widely distributed it has usuallv difl'erentiated into a number of local forms; Pteiopvs hi/po-


Ixxvi GEOGRAPHICATDISTKIBUTION. vtelavus, for instance, covers practically the whole area from IndoChina to New Guinea and Woodlark Island, but is broken up into no less than twelve subspecies ; Fteropus vampyrus ranges over the whole of the Indo-Mala3an subregion, but divided into six races ; IJjiomoj^horvs wuhlbergi is distributed over the greater part of the Ethiopian region (except the Guinea coast west of Canieroons), but falls into two races — a western and eastern ; Cynopterus bmchyotis is known from tho whole of Indo-Malaya, extending eastward beyond this subregion to Celebes, westward to Kurma and Assam, and reappearing farther west in Ceylon, but is broken up into eight local forms ; MacroyJossus Ingocliilus extends from Borneo in an unbroken range eastward to the Solomon Islands, but has differentiated into four subspecies ; Syconycteris crassa ranges from tho Moluccas to the islands south-east of New Guinea, but has split into five races. There are, however, a few noticeable exceptions from the general rule of the relatively narrowly limited range of the species and subspecies; Eidolon hdinmi is generally distributed, without any appreciable change of characters, over nearly the •whole of the Ethiopian region (South Arabia excepted) ; Itousetius ampJexicaudatus occurs in Cambodja, through the Philippines, Uorneo, Sumatra (not Java), east to Flores and Timor, apparently without splitting into local forms ; Bousettus brctcliyotis ranges, seemingly unchanged, from the Amboina group to the Solomon Islands ; Cynopterus bracJiyotisbrachyotis from Lower Siam, through the Malay Peninsula to Sumatra, Eorneo, the Philippines, and Celebes ; Mncroylossus layochilus lagochilus over Borneo, the Philippines, Celebes, and the Amboina group. Whether these exceptions are really much more frequent and much more striking among Megachiroptera (and Chiroptera in general) than among non-flying Mammalia, is perhaps doubtful. In any case, against these few instances of an exceptionally wide distribution of one apparently unchanged form may be placed a large number of species known only from, and in many cases probably really restricted to, one island or group of islands. The evidence afibrded by the geographical distribution of Bats has generally been considered of doubtful value ; hence they have either been entirely excluded from the material worked out by zoogeographers or at least treated with pronounced suspicion, as likely to be more or less unreliable documents of evidence. This unwillingness or hesitation to place Bats on an equal zoogeographical footing with non-flying Mammalia would seem to be due, partly to the preconceived idea that owing to their power of flight Bats must evidently have been able easily to spread across barriers which, in ordinary circumstances, are insuperable for wingless Mammalia ; partljf to the fact that hitherto very often whole series of distinct forms have been concealed under one technical name. So long as (to mention only three cases among many) " Macroglosstis minimus" was believed to range unchanged from the Himalayas to New Guinea, Australia, and the Solomon Islands (now two distinct genera, thirteen recognizable forms), or " Cynoi-iUrvs margiiuilus "


GEOGRAPHICAL DISTRIBUTION. IxXvii over India, Ceylon, Indo-China, and Indo-Malaya (now six species, fourteen forms), or " Rhhiolopfms ferrum-equiniim" uniformly over Europe, Asia, and Africa (now numerous distinct forms), they were undoubtedly of questionable value as zoogeographical material, liut these and similar anomalies invariably disappear as soon as modern methods of discrimination apijlied on vastly increased material render it possible to draw the lines of separation between the species (and their local modifications) somewhat more closely in accordance with the lines drawn by Nature. The second argument referred to above, that the spreading of Bats from one locality to another must obviously have been greatly facilitated by their possession of wings, may in theory appear plausible enough, but when tested on the actual distribution of the species and subspecies it proves to be of much less importance than commonly supposed ; it rests, in reality, on a confusion of two different things : the j)Ower of flight no doubt would enable a Bat to spread over a much larger area than non-fiying Mammalia, but, as a matter of fact, only in very few cases is there any reason to believe that it has caused it to do so. The following pages, in which the distribution of the Alegachiroptera within each zoogeographical region or subregion is discussed in some detail, will give ample evidence to this effect, but a few of the more striking examples may be mentioned here : a species of Pleroj)us inhabits the island of Pemba, south of Zanzibar, but although this island is separated from Africa by a channel only o5-40 miles wide, not this particular species only but the whole genus is unknown from any part of the adjacent continent ; although absent from Africa the genus Pteropus is distributed all over the Malagasy region, and each group of islands (Madagascar, Comoros, Aldabra, Seychelles, Mascarenes) has its own peculiar species, intermigration between the groups of islands is unknown ; the Epomophorine section of Fruit-bats is distributed over the whole of the Ethiopian region (eight genera, nineteen forms), but not a single form has spread to any island of the Malagasy region ; the Fteropus melanotiis group of species is distributed over the Andamaus, Nicobars, Nias, Engano, and Christmas Island (south of Java), and the whole group is confined to this chain of islands, no form having spread to the neighbouring Malay Peninsula or Sumatra; Fteropus hi/pomeJanus is represented by a local form in Engano, off Western Sumatra, but the species, though widely distributed elsewhere in Indo-Malaya, is unknown in Sumatra and Java ; the Megachiropteran fauna of Ceylon shows, of course, very close affinities to that of the Indian Peninsula, but the Indian llouseittis lesclienanlti is replaced by a distinct species {seminudus), and the Indo-Chinese and Indo-Malayan Cynopterus hrachi/otis is represcntea by a local form, though the species is unknown in the Peninsula ; the Eruit-bat faunas of the Malay Peninsula, Sumatra, and Borneo are closely interrelated, like their Mammalian faunas in general, but each has some distinct autochthonous forms of Eruit-bats (Borneo even two autochthonous genera), as it has of other ifammalia ; the Javan Mammalian fauna in


Ixxviii GEOGUAPHICAI. DISTEIDUTIOX. general is more peculiar, both by the absence of some of the forms found in Sumatra and the Malay Peninsula and by the greater percentage of autochthonous forms, and this is again borne out by the Megachiropteran fauna of the island (absence of two Sumutran genera, presence of one autochthonous genus, among ten forms six autochthonous); the Ftero^nts rui/neri group is represented probably all over the Solomon Islands, but it has ditferentiated into five distinct species, one in the Bougainville group, a second on Yella ].avella, a third in the New Georgia group, a fourth on Guadalcanar, and the fifth on San Christoval. This (and a series of similar examples might easily be adduced) tends to show that the present distribution of the Megachiroptera has not been influenced to any great, and as a rule not even to any appreciable, extent by their power of flight ; if it had, the Fruit-bat fauna of one group of islands could not, so commonly as is actnally the case, difl'er from that of a neighbouring group or continent, and the tendency to differentiation of insular species or forms would have been neutralized by the free intercourse between neighbouring faunas. Etldojiian Herjion. 13 genera, 32 si^ecies (3-1: forms). Of the four primary sections of Megachiroptera, one, the Bpomopltorus section (8 genera, 17 species, 19 forms), is entirely Ethiopian ; of the liousettus section (9 genera, 125 species, 147 forms) three genera are represented in this region, Eidolon by two species, liousettus by seven, and Pteropus by one ; the Cr/nopterus section (11 genera, 31 species, 41 forms) by one peculiar genus, Myonycle)-is (four species) ; the Macroglossince (7 genera, 13 species, 21 forms) by one peculiar genus, Megalo'ihssus (one species). Of the thirteen genera ten are peculiar, viz. all except Eidolon (outside the Ethiopian region occurring only in Madagascar), llmis^ttus, and Fterojms. Of the tliirty-two species thirty are peculiar, viz. all except Rm'settus cef/ifptiacus, which extends to the eastern Mediterranean subregion, and liousettus arahiciis, which extends to Karachi. A subdivision of the Ethiopian region, on the basis of its Megachiropt«ran fauna, is very simple indeed. The region falls decidedly into two well-defined ])rovinces, viz. (1) the Wtst African Province, i. e. the whole of the Guinea coast and Congo basin, east to the western bank of Victoria Nyanza, south to Benguela and Damaraland, thus approximately synonymous with the Great West .\frican Forest llegion, though at least in the south-west extending beyond the limits of the forest belt, and (2) the East African Province, i. e. the eastern side of the continent from Erythrea and Abyssinia in the north to the Cnpc Colony in the south. So sharp is the contrast between the faunas of these two provinces that the large majority of genera and species occurring in the former are unknown in the latter, only a few species extending beyond its limits to part of East Africa. A single species, Eidolon Jiehnnn, seems to be thoroughlv common to both, ranging from Senegambia,


GKOfiKAPinCAL UISTRIBUriON. Ixxix Seminar, and Somaliluiid in the north, to Namaqualanrl, Mashona, and Nyosalaud in the south. Besides these two provinces, .South Arabia and the island of Peinba re(inire special remarks. — Matsehie's attempt to subdivide the Ethiopian region into a large number of zoogeographical districts ("Gebiete"), each with its own distinct Mammalian fauna, breaks completely down when tested on the distribution of the Ethiopian Fruit-bats. ^\^est African provi)ice.— 12 genera, 23 species C24 forms). Pterojms is the only genus which, though represented in the Ethiopian region (the island of Pcraba), is absent from W. Africa. Of the twelve genera no less than nine are peculiar, viz. Flerotrs, Epomops, Ifi/psi(jnafhi(S, Micropteropus, Kaiioiii/ctens, Scofoni/cteris, and Casini/cteris (all of the Epnmciphorus section), Mi/o»>/cteris (Ci/noptertis section), and MegaJoglossiis {31'(crofjJossincr) ; expressed in other words : all genera of the Epomophorine section of Eruitbats are contined to this province, except Ej)omophorus itself, which is common to W. and E. Africa, and this province (" Afro-Malaya," as it was, perhaps a little jokingly, called by the late Dr. 11. Bowdler Sharpe) is the only part of Africa in which the otherwise essentaallj' Oriental and Austro-Malayan Cynopterine and Macroglossino sections are represented. The three non-autochthonous genera are Eidolon, lloasettxs, and Epomopihorns. The genus liousettus falls into three subgenera, liouseUns, Slenon}jcteriK, and Lissonycteris ; of these the first is widely distributed in the Ethiopian and Oriental regions and Austro-Malaya, while the two latter are confined to the Ethiopian region ; Steaoiii/cterin is common to the West and East African Provinces (one species in each), Lissoni/cteris (two sjieries) is peculiar to West Africa. Of the twenty-four "West African forms no less than twenty-one are peculiar, {. e. all forms except Eidolon helvum (generally distributed in the Ethiopian region), lioiisettan cvijfipliacus (Eoanda and Congo to Egypt, Palestine, &c.), and Epomophorus (jamhiivms (Guinea coast to Sennaar and Abyssinia); only one of the twenty -one peculiar forms, viz. Epomophorus wahlbergi haldemani, extends eastward a little beyond the borders of the province, into German and British East Africa. Suggestions as to a possible subdivision of the province would be ])remature ; in too many cases the range of the genera and species is only imperfectly known. Some species (Ili/psignathus moiistrosus, Micropteropus pusillus) have a continuous distributit)n from Gambia, along the Guinea coast, through the Congo ]5asin to Victoria Xyanza ; in other cases the area is divided between several species of one genus : Epomops hurttilofcri in Liberia and Sierra Leone, E. fritnq\teti from the Gold Coast, east to Victoria Xyanza, south to Loanda, E.dohsoni in Bongucla and Katanga; Mi/oni/ctcris leptodon in Liberia and Sierra Leone, J/. wroiKjhtoni in the \\'e]lc district of the Congo Basin, M. torquafa in the Lower Congo district; and Angola. Islands in Gnlf of Ginnfn. — The followin

IXXX GEOGRAPHICAL DISTRIBUTION. and Annobon : — Fernando Po, Eidolon helvum, Hypsignathns monstrostis, Scotom/cteris zenlceri : Principe, Eidolon helvum : San Thome, Eidolon helvum, Myoni/cteris hrachycephcda : Annobon, Eidolon helvum. Myonycteris brachycejdiala is known only from San Thome, and the sole representative of the subgenus Phyyeiis ; all the other three species are common to the islands and the adjacent coasts of the continent. East African province. — 3 genera {Eidolon, Rousettus, EpomopJiorus), 10 species. This province is incomparabl)' poorer than W. Africa, both in genera and species and in the number of really peculiar forms. None of the genera is autochthonous. Eidolon helvum, Rousettus ^gyptiacus, and Epomophorus gambianus are common to this and the foregoing province ; the other seven forms (two Rousettus, five Ejjomojihorus) are entirely East African. Rousettus leaclii ranges from the Cape Colony north to British East Africa, and is in Egypt and part of West Africa replaced by R. cegyptiacus ; Epomophorus wahlbergi ivahlbergi likewise from the Cape Colony to British East Africa, and is replaced in West Africa by E. IV. haldemani ; Epomophorus crypturus extends from Transvaal to Nyasaland, and is replaced northward, from Tanganyika to Erythrea, by E. anurus. It will be noticed that the southern forms have as a rule an unbroken distribution from the Cape Colony or Transvaal north to British East Africa or at least to Nyasaland. South Arabia. — 2 genera, 2 species. Affinities decidedly African, J^i(fo?o)isa^f»!M/rt (autochthonous) being closely related to the widelydistributed African E. helvum, Rousettus arabicus to the East African R. leachi. R. arabicus extends as far east as Karachi, N.W. India. Pemba Island. — Pi.emarkable by being inhabited by a distinct species of Pt/ropms, although the genus is absent from the whole continent of Africa. The Pemba species {Pt. voeltzkowi) is closely related to a Malagasy form (Pt. comorensis). Malagasy Region. 2 genera, 9 species (10 forms: Eidolon one, Pterojnts nine); neither of the genera is confined to this region, but all the species are peculiar. The single species of Eidolon (E. dupreanum, closely related to the Ethiopian E. helvum) is apparently restricted to the island of Madagascar, while Pterop)us is distributed over the whole of the Malagasy region. The affinities of the Malagasy Pteropus fauna have been discussed elsewhere (pp. 79-80). Indian and Ceylonese subregions. 3 genera, 7 species (8 forms), viz. Rousettus (three species), Pteropus (two species), Cynoptcrus (two species, three forms). Rousettus arabicus is a South Arabian species which only enters the


UKOGKAl'IirCAr, DISTRIurTION. hxxi north-west corner of the Indian Peninsula ; li. le.Juncndti occur. all over the Teninsnla as well as in Lulo-Gluna, but i« in Ceyloa rep aced by a very closely related peculiar fori, H. semJdus. 27nL^M7 %^^" ^"^''" ""^ Himalayan representative of tHvi 1 df M 7^> ' r"^'^''"' (^'""' ^^^'''^^ it only dm-ers in trivial details; Peninsular specimens (subspecies ^fuanieus) are nidistinguishable from Ceylonese, but slightly different from ilimalayau (subspecies leucocej^halns) a.d :\raldive specimens (««,?) Cpioptcrus spJnu.v has in continental South Asia differentiated into two subspecies, the one (sphin.v) ranging over the southern ar-d eastern provinces of the Indian Peninsula! south to Ceylon ortheast nortiern Indo-China, the other („eticus) known ojy rom the central and north-western provinces of India. In addition ioJtoMus ssminudns Ceylon possesses a second peculiar Pruitbat, Cynopten.s brachi/olis ceylonensis, to which there is no equivalent in the Indian Peninsula, all the other forms of C. bracZ otis being Indo-ilalayan or Indo-Chinese. ^ ladoChinese suhnyion. 6 genera, 14 species (15 forms), viz. llonsetu,s (two species) l^eropus (seve.i species, eight forms), Cynopterus two ^"c e.) >Sp]ueruts (one), Eouycteris (one), Macroylossus (one). Continental Indo.China.-Roasettus JeschenauUi and Cynonteru, splnn. .pAuuare invaders from west, the Indian Penmsuk h same may be said of the Himalayan Pteropus gigar.teus ulcocei^alns, but it is subspecitically distinct from the Indian form Evidence of the affinities of the present area with Indo-ialaTa are, first the direct invaders from the latter; second, those forms closelj related to Iiido-ilalayan forms. To the first cate^orv beongi?o«.... a.n,ae.icaud[at.s, Eonycteris spekeJ^titZl rjlossus nunnnus. To the second, I'terojn^s inieJediusWenasserim) a representative of the Indo-Mala/an Ft. van>pyrL S Si (Siam and Sa,gon), a peculiar species of the slme group and fi-hpomelanus condorensis (Siam, Cambodja, Palo Condor) a local race o a widey-distnbutcd Indoand Austro-Malayan spedes Cynopterus brachyoUs anyulatus, rather than being an invader from wTrd'thr^::':^;'''^^ "' ^"^^-^^Z-^-^ -^ch h°as spread Luh.fonh 8 a highly peculiar autochthonous genus and species of the Cynopterus group so far known only from Burma. Formosa.-T\x& single species recorded, Ptevopus formosvs, has ts closest relatives to the south in the Amboina groip (Pt. lops) to the north in the South Liu-kiu islands (Pt. dasymallus). ^ ^' Jl^^i/^-^^^-^-j^^i^lcmds.-Tbe two species of Pteropus known from these islands belong to different groups of the genus. Pt cZ ForL"an/y ,°^«"t-"-l -^--; ^'^ '^-^ repre^sentative of tte^. fonnosue; Pt. loochoensis a species of the otherwise entirely Polynesian Pt, rnariannus group. omerwise /


Ixxxii CEOGUArnicAL msTEiBuriorf. Indo-Malayan subrcgion . 13 genera, JiG species (55 forms). Of the genera seven (all nionotypic) are peculiar, yiz. one of the Rousettus section [Dobsonia liranch), Hn>-j)yionyrtens, and six of the Gijnoptervs section, Ptcnochiriis, Megteropx, Di/acopteru$, Balionycteris, Ch'rona.v, and Penthelor. Twenty-seven species (forty-six forms) are peculiar. Only one or two forms have entered the present subregion from the west (Indo-China), viz. Cijnnplerus sphinx (here differentiated as a distinct subspecies, titiJuec/ieiJvs) and, perhaps, Cynojitenis brachyoiis angjilatus; but four species have spread in the opposite direction, from Indo-Malaj'a into Indo-China, viz. Rousettus amplexicaudattis, Fieroptis hypomelanus, Eonycteris spelcva, and Macroglossus minimus. Macroylo^sus lar/ochilus is the only species which is widely distributed both in Indoand Austro-Malaya, but four other Indo-Malayan species extend into the extreme western or south-western Austro-Malaya, viz. Pteropus mimus and Cynopferus brachyotis to Celebes, Rousettus amjjlcxicautlatus and Fierojms vampyrus to Timor. Malay Peninsula, Sumatra, and Borneo. — Taken as a whole this " province " is characterized, as against any other part of IndoMalaj'a, by the presence of four peculiar genera, Megcerops (ecaudatits), Dyacoptenis {spadicevs), Balionycteris (mactdaia), and Pentheior (liicasi). It must be mentioned, however, that Balionycteris has its "equivalent" in Java {CJtirona.v), and Penthetor a close relative in western Austro-Malaya {Thoopterus). The status of the Malay Peninsula, Sumatra, and Borneo inter se is, judged by their Megachiroptera, practically the same as if judged by any other large group of Mammalia, •/. e. they are undoubtedlj' closely connected zoogeographically, while at the same time each has certain distinctive features, these latter decidedly more pronounced in the Eorneanthan in the Peninsular and Sumatran faunas. Borneo has two peculiar genera {Dyacnpterus, Balionycteris) and four peculiar forms, the Malay Peninsula and Sumatra have no peculiar genera, but each one peculiar form. The following forms arc common to all three areas, viz. Rousettus ample.cicaudatus, Cynop^terus brachyotis brachyotis, Meya'rops ecaudatus, and Penthctor lucasi (not actually recorded from Sumatra itself, but from the E,hio Archipelago) : of these the two latter are the most important, since found nowhere outside this province ; the two former have a wider distribution in Indo-Malaya. The following are common to the Malay Peninsula and Sumatra, but unknown from Borneo, viz. Pteropus vampyrus malaccensis (in Borneo replaced by Pt. v. natuna:), Cynop)terus brachyotis angidatus, Eonycteris spelcea (in Borneo replaced by E. major), and Macroglossus minimus (in Borneo replaced by a form of M. lagocMlus). No form is common to Sumatra and Borneo but absent in the Malay Peninsula, and none is common to the Malay Peninsula and Borneo but absent in Sumatra.

PAGE 87 distribution. Ixxxiii Andamans.~On\y Pteropus (t^ro species) and Ci,mpterus (a local form of C. hraclnjotis) l,avc found their way to these island lu hree orms are peculiar. Of the two s^Lies of Pter^^s, one (PL teller) belongs to the Pt. melanotus group, a group en irelv confined to the Andaman, Nicobar, Nias, Engano, and Chris Lv Ks and chain; the other (P. ...^..s-) to the Z lUufponrdanus group. C;faopten.s brachyotis is a common IndoChinese and Indo-l[alayan type. Nicohars.~L% in the Andaraans, one species of the Pteroom melanotas^ron^ i^Pt. ^nelanot.s), one of the Pt. k.ponulannZZ ifuaaidus) an^ one local form of Canopterm hraduioth, all throe torms peculiar, though closely related to those of the Andamans. Nias.—MacrngJossus minimus sohriaus (also in Sumatra^ • a J.ecuhar species of the Pteropus melanotus group (Pt nuulLJ rfhepf^^t'l '^ ''' ""''-'^^ ^^'^^'?^ ^' "^^^ I4^trt u : of the It. melanotus group is al,scnt in Sumatra); no less than three pecnliar forms of C.noj.terus, vi. one local «ubs^^ ies o C. hraIn,ot,s (a different subspecies in Sumatra), one species ;h«/o.) allied to C. bracJnjotis anrpUatus (Indo-China Euloninsula, Sumatra), and one unusually well-differentiated snecies &r?;'j.t;.;^;sir""-"°° " "'""""'• <-*--''^^" the widely-distributed Pousatus an,ple.ncaudalus. The differences from the fauna of the neighbouring Sumatra are strikit tha many Sumatran types are absent in tins outlying island, wa^ only o be expected; but the Pt. Jujponuhmus ^ud^melanous groups though represented m Engano, are unknown in Sumatra. ^ ' Christmas Island (S of Java).-One peculiar species oi Pteropus belonging to a group of the genus {Pt. melanotus) represented £ where only in Engano, Xias, the Alcobars, and Andamans. J-«,« The Fruit-bat fauna of Java, like ils Mammalian fauna n %nZk '' ^oma^-kable (1) by the absence of certain types foS Intorb , ^'°'"'"\ (') ^^' ^^' ''^^''''^^y 1-rge number of autochthonous forms, and (3) by its complicated affinities these poniting partly toward tlie neighbouring islands, part y w th exc usion of these latter toward Indo-ChinI To takJ thes^ thJee categories of faunistic characters separately :—(i) The followip<. iorms present both in Sumatra (and the Malav Peninsula) and Borneo are absent from Java, viz. Eousettusample.cicaJ(aC Cynopta-us hracujot.s hrach;,otis, Me.,a>rops (ccaudaL), and P.t' ITJT'^ . (2) Of ten forms known from Java six ar. pecu i one of these (C%u-o,m.. mdanocepludus) being a pecnliar genus (.5 a) Poruis affiuities with Sumatra, or Borneo, or boll /2


Ixxsiv GEOGRAPHICAL DISXRIBmOX. viz. Bousettus 7ninoj\ the Javaii representative of B. cimple.vieauJalus; Pieropus vnmpurus vojnpyrus, replaced in Sumatra by Ft. V. malaccensis, in Borneo by Ft. v. natunce ; Cj/nopterus hruchyotis javanicus, a local form of the widely-distributed C. b. hracliyotis ; Cunopterus horsjieldi horsjieldi, replaced in Sumatra by G. h. lyoni, while the species is absent in Borneo ; Chironax mclanocej)halus, represented in Borneo by BaJionycteris macidata, botli genera having no equivalent elsewhere; Eonyctcris speJcea, also in Sumatra, but in Borneo a distinct species {major) ; Macroghs^ns minimus, also in Sumatra, but in Borneo a distinct species (lagocTiilus) : (3 b) Evidence of affinities with Indo-China, with exclusion of Sumatra and Borneo, Bousettus shortridyei, closely allied to the Indo-Chinese B. leschenaulti, a bat which has no rei)rescntative elsewhere in Indo-Malaya; probably also Cinmpterus spJiiny tittJuvcheihis (though this form is common to Java and Sumatra), Cynopterus sphin.v being decidedly not an Indo-ilalayan but an Indian and Indo-Chinese type, which, probably, in Java has ditferentiated into a distinct race, C. s. titthecheilus, which again has spread westward to Sumatra, eastward to Lombok, perhaps as far as Timor. Philippines. — 7 genera, 13 species (14 forms). The general characters of the Fruit-bat fauna (which is, of course, still imperfectly known) would seem to be these : the autochthonous element is strongly developed (tv/o genera, one of which is so peculiarly modified that it has recently been proposed to separate it in a distinct subfamily, six species, nine forms), the affinities of the autochthonous forms are partly Indo-Malayan, partly Austro-Malayan, while one species has its closest known relatives in the Bonin and JPolew Islands; in addition, some direct invasion has taken place both from Indo-Malaya (Borneo) and Austro-Malaya (Celebes), perhaps also, though if so to a much less extent, movements in the opposite direction. The details are: — Indigenous genera: Harpyyionycteris (whiteheadi), a highly peculiar genus, with no nearer relative than the Austro-Malayan Dobsonia ; Ptenochirus {jwjori), closely allied to Cynopterus, and most closely to its " Niadius '"' section, which is known only from the Malay Peninsula, Sumatra, Nias, and Java. (2) Indigenous representatives of (essentially) Indo-Malayan species: Pteropus hypomelanus cagayanus; Pt. vampijnis lanensis. (3) Indigenous representatives of Austro-Malayan species: Pteropus pumdus, closely allied to Pt. griseus (Bonerato, Dyampea, Timor) ; Acerodon juhatus and lucifer, all other forms of Aeerodon being Austro-Malayan. (4) Indigenous representative of an otherwise exclusively north-west Polynesian group : Pteropus leucopterus, allied to Pt.pselaphon (Bonin Islands) and Pt. pelewensis (Pelew Islands). (5) Probable invaders from Indo-Malaya (Borneo) : Bousettus amplexicaudatus ; Pteropus speciosus (?) ; Cynopterus bracliyotis bracJiyotis ; Macroglossus lagochihis lagocldJus. (6) Invader from Celebes (or from the Philippines into Celebes) : Pteropus mimus.


GEOGRAPHICAL PISTRirtriTOy. IxXXV Austro-Mahtyati Suhrecjion. If) genera, 80 species (93 forms), thus much richer in species than any other subregion, the number of species being more than twice that of Indo-Malaya ; 43 per cent, of all known genera and exactly the same percentage of all known species are represented in this subregion. Of the fifteen genera seven (with altogether nineteen species) are peculiar, viz. four of the liousettus section, Boneia (one species), Ptcralopex (two), Sti/hctenium (one), and Dohsonia (twelve) ; one of the Ci/nopterus section, Thoopterus (one) ; and two Macroglossincp, Melonyctcris (one) and 2\esoni/ctens (one). The affinities of these seven autochthonous genera are as follow s : — Boneia is closely related to the widely-spread llousetius ; Fteralopeo' and Sti/Iocienium are offshoots from the even more widelydi.stributed Pteropns ; Dohsonia is an unusually peculiarly modified g( iius with only one near relative, the Philippine Harpyionycteris, and is both by its comparatively large number of species, all strictly confined to this subregion, by its many aberrant characters, and ])y its distribution over practically the whole area from Celebes to the Solomon Islands, zoogeographically perhaps the most important of the autochthonous genera ; Thoopterus is the Austro-Malayan representative of the Indo-Malayan Fentlietor x while Midonycteris and Nesonycteris have no closer relative than the Polynesian Notopieris. The non-autochthonous genera are. liousettus, Pteropus, Acerodon (found elsewhere only in the Philijipines), Cynoptents (only entering the extreme western part of Austro-Malaya), Nyctimene (entirely Austro-Malayan, except for one Australian species, and one common to New Guinea and Australia), Eonycteris (ranging from IndoMalaya only to Celebes), Macroyhssus (from Indo-China to the Solomon Islands), and Syconycteris (entirely Austro-Malayan, except for one Australian species). If Nyctimene and Syconyctei-is were added to the number of autochthonous genera, and they could evidently be so without much error, the number of such genera would be no less than nine (out of fifteen), with altogether thirtythree species, so that the number of peculiar species belonging to peculiar genera would be 41 per cent, of the total number of species known from the subregion. Seventy-one of the eighty species (89 per cent.) are entirelyconfined to Austro-Malaya. The nine species that are not wholly restricted to this subregion are : — (1) such as extend from IndoMalaya only to the Lesser Sunda Islands, viz. Rousettus amplexicnudatus, Pteropus vampyr\is, and Cynopterus spJiin.v (subsp. titthcecheilus) : (2) one extending from Indo-Malaya only to Celebes, Cynopterus bracJn/otis (suhsi^. brachyotis): (3) such as extend from Ihdo-ilalaya over a greater part of Austro-Malaya, but here differentiated into a number of subspecies, viz. Pteropus hypomelanus and Macroylossus layochilus : (4) one common only to the Philippines and Celebes, Pteropus mimus: (5) such as are common to Xew Guinea and Australia, viz. Pteropns ronspiicilJaius and Nyctimene pa2^uanus.


ixxxvi geogra; nicAL w.siraisuTioN. Celebes and Sanr/Jiir Islands. — 11 genera, 17 species (excluding a few doubtful records from literature). Two genera, both monotypic, are peculiar, Boneia, closely allied to liouscttus, and Stijloctenium, closely allied to I'tero/nis and more particularly to the entirely Austro-Malayan I'L temminclci group, one species of Avhich {Ft. ]^iersoncUus) occurs also in Celebes. 80 far as the records go (for there is no doubt still much to be added to our knowledge of the Fruit-bats of Celebes and Austro-Malaya in general) seven species seem to be confined to Celebes, with the Sanghir islands, viz. liousettus celebensis, Boneia bidens, Pteropus dobsoni, Stylocteniiim wallacei, Dohsonia exoleta, Nyctimene minutus, and Eonycieris rosenhergi ; and in connection with these may be mentioned Pteropus liypomelanus mucassnricus, Pteropus alecio, and Acerodon celebensis, though the first is known to extend north to the Talaut islands, the second south through Salaycr to Lombok, and the third south to Salayer and east to the Sula Islands. An aiiah'sis of the relationships of the fauna gives this result: — (1) Indigenous species pointing, geographically, in uncertain direction, liouseifus celebensis, a rather peculiar, narrow-toothed species of doubtful affinities, and Boneia bidens: (2) Indigenous species of an otherwise entirely Indo-Malayan genus, Eonycteris rosenberrji -. (3) Forms common to Celebes and some part of Indo-Malaya (Borneo and Philippines), Pteropus mi^nits, Cynopterus brachyotis brachyoiis, and Macroylossus layocMlus lar/ocJiilus : (4) Distinct form Avith quite close relatives both ill Indoand Austro-Malaj'a, Pteropus I/ypomelanvs macassaricus : (5) Distinctforms with clearly pronounced Austro-Malayan affinities, Pteropus dobsoni, Pt. alecto, Acerodon celebensis, Styloctenium ivallacei, Dobsonia exoleta, and Nyctimene minutus : (6) Forms common to the Celebes and Moluccas (Gilolo group, or Amboina group, or both), Pteropus caniceps, Pt. personatns, Thoopterus niyresceiis, z^nd A'yciiinene ccplcalotes. The general conclusion is that, while the Indo-Malayan element is by no means inconspicuous, the Austro-Malayan affinities of the Fruit-bat fauna are decidedlj' predominant. Amboina group (Buru, Amboina, Ceram, and smaller islands) and Banda Islands. — genera, 14 species. Of the Celebean genera, Boneia, Acerodon, Sfj/locfenium, Cynopterus, Thoopterus, and Eonycteris are absent, but Syconycteris has been added. No genus is autochthonous. Five species are (so far as known) restricted to this group of islands, viz. Pteropus pallidus (closely allied species in Celebes and Timor), Pt. Hops (no very close relative until in Formosa), Pt. cltrysoproctus (]K'rhaps also in Sanghir islands; curiously enough no closely related species until in the Solomon islands, where the gronj) is widely distributed), Pt. ocularis (closely allied species in the (iilolo group and New Guinea), and Nyctimene varius (a closely allied species in Celebes) ; to these may be added two peculiar subspecies, Dobsonia viridis umbrosa (another subspecies in the Key Islands) and Syconycteris crassa, major (other subspecies in the Papuan section of Austro-Malaya). The non-


Gi:onK\ri[ic.\r. DisTr.iP.nioN. Isxxvii ])Oculiar elemeui points towards any of the siirroundinf,' islands, the (jtikilo group. Celebes, South West Islands (Timor), .South East Islands (Timor Laut, Key, Aru), and New Guinea, it being in most cases mere conjecture whether the movements have taken place from the Amboina group into the neighbouring group or iu the o[)posite diicction. G'dolo ijroup (Morotai, Gilolo, Ternatc, Batjan, &c.). — 4 genera, 7 species; but tlie fauna is no doubt more imperfectly known than tliat of the Amboina groiij). Tlie faunistic leanings are moro decidedly toward Celebes than to the Amboina grou[) and New Ciuinea, but a large number of Celtbean genera are absent from the list, viz. lloiisi'ttiis, Boneia, Acerodon, tSti/Ioctenium, C'l/nojiterns, Konijcteris, and Mcici'Of/lonsits; the absence of Iiousettus and Macroijlossiis, both generally distributed elsewhere in Indoand AustroMalaya, may be due to incompleteness of the I'ecords. As in the case of the Amboina group, no genus is autochthonous. Two species seem to be confined to the (iilolo group, Duhsonia cremtlata (a closely allied species in the Amboina group, viz. I), viridis) and jXiictimcne aJhiventcv (its closest relative iu New Guinea); in addition to tliese, one autochthonous subspecies of Pteropus hiqiOrtnlamis (other subspecies in CelebcK and New Guinea, but not in the Amboina group). Specimens common with Celebes and Sanghir Islands are, Fieropus caniceps, Ft. 2>erso»a(us, and Thoojiierus nigresccns (genus confined to Celebes and the jjresent group). One species is common with N.W. New Guinea, Fieropus cliryscmcJien, New Guinea (including Ghebi, Salawati, Mysol, and Waigeou, but excluding the satellite islands to the north, east, and south).— 7 genera, lO species. The genera are tlie same as tliose occurring in the Amboina group, with the single addition of the monotypic j\lel('»^icteris; but all the species, except one, are different. No genus is strictly confined to New Guinea, the nearest a[i])roach being Jleloin/cteris, wliich is known onl}from t!.is island and the ])ismarck Archipelago. No zoogeograidiical jirovince has so rich a Nyclimcne fauna, all four natural groups of this genus being represented, and two of the groups (the cijdotis and aello groups) found nowhere else. Six species are autochthonous, viz. Pteropnx cpvlarivs (closely allied to Ft.macroiis from the .\ru Islands), Dobsonia minor (no close relative elsewhere), D. ma

IxXXViii GEOGIIAPIIICAL DISTIUIiUTION. confined to New Guinea is : (1) occurring both west (Amboiua group) and east of New Guinea (Solomon Islands), Hoiisetius hrachi/otis : (2) common with the Gilolo group, Pteropus chrysauchen: (^3) common with the Key and Aru Islands, hut not extending to Australia, Macrofjlossus hujochilKS nanus: (4) common with Australia and also with either Key or Aru Islands (if not both), JSyctimene papuanus and Suconiicteris crassa papuana : (5) direct invader from New Guinea through the Torres Straits islands into Australia, Pteropus conspicillaius : (6) common with the Bismarck Archipelago only, Melonycleris melanops. A faunistic contrast between the North-west and South-east of New Guinea is, so far, only indicated by Pieropus chri/saachtn of the former area (extending to the Gilolo group) being replaced in the latter by its geminate species. Pi. conspiciUatits (extending to N. Australia) ; but more definite suggestions for a subdivision of New Guinea into faunistic districts may be derived from the distribution of the species of Nyctimene when better known. Key Islands. — Four genera have been recorded, Pteropus, Dohsonia, Nyctimene, and Syconycteris, each (so far as known with certainty) represented by one form ; in addition to these, Macroglossiis probably occurs in the islands. Of the four forms three are ncculiar, viz. Pteropus keyensis, a species allied to Pt. melanopogoii from the Amboina group, Banda Islands, and Timor Laut, but with no close relative in New Guinea; Dohsonia viridis viridis, also with Amboiuan affinities (no ecjuivalent in New Guinea); and Syconycteris crcssa Jceyensis, which is more intimately connected with the New Guinean than with the Australian race of the species. The fourth form, JSyciiinene p)apuanus,\s common to the Key Islands and New Guinea. As evident from this, the Fruit-bat fauna shows both New Guinean and Amboinan affinities, and the former are in so far more intimate than the latter, as the two Amboinan forms have differentiated the one into a distinct species, the other into a subspecies, whereas of the two New Guinean forms the one is unchanged, the other distinguishable as a subspecies. Arw Islands. — Nyctimene has not been recorded from these islands, otherwise the genera are the same as in the Key Islands, but the relative distinctness of the Aru fauna is shown by the fact that no form is known with certainty to be common to both groups (Dohsonia inoluccensis and Macroylossus layochilus nanus may occur in both). The affinities point partly toward the Key Islands and the Amboina group (Pteropus aruensis, a peculiar species, related to Pt. keyensis and to the Amboinan Pt. melanopogon ; and Dohsonia violuccensis, indistinguishable from the Amboinan form), partly and more decidedljtoward New Guinea (two common forms, Macroylossus lagochilus nanus and Syconycteris craesa papuana ; one distinct species, Pteropus macrotis, closely allied to the New Guinean Pt. ('pularius).


GEOaRAPHICAL DISTRIBUTION. Ixxxix Bismarck Arddpelago. — 7 genera, 8 species. Four (or, if Plerop us Jieohibc miens is distinguishable from Ft. papuanus, five) forms are peculiar. Naturally the fauna is chiefiy influenced by that of the neighbouring New Guinea ; the genera are the same, and of tlie species and subspecies liousetlus hracluiotis, Macroglossus Inyochilus nanv.s, Melomicteris melutiops., and (probably) I'leropns ntohihemicus are direct invaders, Niictiinene major and Syconycteris crassd finschi only slightly altered iudigenous representatives of JS'ew (iuinean forms. But at the same time there seems to be a distinct Moluccau (non-Papuan) element, represented by Pteropus cupistratus and JJobsonia pnedairi.v, both of which, though welldiHerentiated indigenous species, have their closest relatives in the Gilolo and Amboiua groups, but (so far as known) none in New Guinea. Sohinon Islands. — 7 genera, 16 species. Whether the genera are compared with those of New Guinea or with those of the Bismarck Archipelago, the result is the same : Syconycieris is absent, Mdonyctcris is replaced by the closelj' related Nesonycteris, and Fteralopeais added, otherwise there is no change. But the amount of autochtlionous forms is enormous, all beiug peculiar, except lionsettus hracliyoiis, and of the seven genera two (with altogether three species) are jjeculiar, viz. those just referred to, Fteralope.c and Nesonycteris. The sixteen species may be classed according to their probable faunistic and natural athnities as follows : — (1) Direct invader from west, Ilouseitns hracJn/olis, ranging from Amboina, through New Guinea and the Bismarck Archipelago, to these islands : (2) Indigenous representatives of New Guinean types, three, viz. Nyctimene scitalus, most closely allied to N.geminusivom. New Guinea and N, major from the Bismarck Archipelago ; Macrof/losstis layochilus uiicrotus, the most closely allied races of which are found in tlie Bismarck Archipelago and New Guinea ; Nesonycteris u'oodfordi, as mentioned above the Solomon Islands representative of Melonycteris mr.lanops from New Guinea and the Bismarck Archipelago : (3) Indigenous species having their closest known relatives in the Moluccas only or both in the Moluccas and Bismarck Archipelago, but apparently none in New Guinea (compare remarks on the Moluccan element in the fauna of fclie Bismarck Archipelago) ; here belong, first, five species of Fteropus, viz. Ft. grandis, laveUanvs, rnbianus, rayneri, and coyuaius, all representatives of the Ft. rayneri group, the only other known species of which inhabits the Amboina group ; second, the two species of Dohsonia, D. inermis and MfSOT, representing a group of the genus inhabiting the Amboina grou]), Gilolo group, and Bismarck Archipelago, but not known to occur on the continent of New Guinea : (4) Indigenous species having their closest relatives in Australia, three, viz. Fteropus tvoodfordi, a species strikingly similar in all essential characters to the peculiar Australian Ft. scapulalu.t, only much smaller; Fteropus culoHus and ^olomonis. two species of the Ft. hj/ponielanus group and both perhaps most closely related to the Australian Ft. brmineusi


XC CEOGRArUrCAL DiSTRIBUnOX. (5) Indigenous representatives of a chiefly Polynesian type, th'o peculiar genus FleraIo2'>ex (two species), allied to the I'teropiis psehiplion group, tlic members of which are North-west Polj-nesian in distribution, with the exception of one species occurring in the rhilippines. A discussion of the distribution of the various species throughout the Solomon Archipelago would to a great extent be prematv;re ; the Fruit-bat fauna of northern chain of islands (Ctioiseul, Isabel, Malayta) is as yet entirely unknown, that of the extreme eastern islands (San Christovnl group) very imperfectly explored, and there is probably not a little to be added to our knowledge of the Eruitbats of the other islands as well. But two important facts are already now sufficiently well established to call for some comment here. First, that some species (liouseiiiis brachyotis, Niictimene scihdus, Nesonycttris woo'lfortli, and probably Pteropus ivoudforili) are evidently uniformly distributed from the Bougainville group in the west eastward through the southern chain at least as far as Guadalcauar. Second, that in spite of this spreading of a few species over the greater part (if not the whole) of the Archipelago, there is unmistakable evidence that the Fruit-bat fauna of the Solomon Islands consists in reality of a series of more or less distinctly separated " fauuula\" This is most clearly shown by the distribution of the si)ecie8 of the Pteropus rayneri group : Pt. yrandis inhabits the Bougainville group, is in Vella Lavella replaced bjPt. lavellanuK, in the New Georgia group by Pt. ruliiamis, in the Guadalcanar group by Pt. rayneri, and in the San Christoval group by Pt. coynatns. Other, though much less complete, evidence to the same effect is afforded by the two species of the Pteropus JiypomeJanus group {Pt. colonus in the Bougainville group, replaced by Pt. sohmonis in tlie New Georgia groui>), the two species of Pteralope.v {anceps in the Bougainville group, replaced by atraia in the Guadalcanar group), and the two Dobsonia (nesea in the Bougainville and New Georgia groups, replaced by inermis in the San Christoval group) [compare the distribution of the three known Solomon Islands species of the Microchiropteran genus Hippiosiderus, IJ. diiwps in the New Georgia group, H. oceanitis in Guadalcanar, and //. deinissus in San Christoval]. The faunistic areas of the vSolomon Archipelago indicated by the distribution of the five species of the Pteropms raj/neri group are very nearly the same as those lately recognized by ornithologists. Australian subregion (Continent of Australia). o genera (none peculiar), 8 species (six peculiar). The only genera which have reached the Australian continent are Pteroptis (five sjiecies), Nyctimene (two), and Syconycteris (one); all of these occur also in New Guinea, which however is inhabited, in addition, by four genera which have not spread to Australia, viz. liottsettus, Pohsonia, MacroyJossus (extends to the Torres Straits islands, but has thus far not been recorded from Australia), and Mdonycteris.


GEOGBAnilCAL DISTKlBCTIoX. XCl As might be expected New Guinean affinities are predominant, no less than five of the eight 6])ccies pointing in their origin, eitlier nndouhtcdiy or at leiist ])robably, toward the great neighbouring island ; PUrojms cons/ncillaivg and Nyctlmene pajmunKs are direct iinehan.i^ed inva lers from New Guinea ; Plerojnis polioccpludas is probably the Australian representative of Ft. ejinlarius (New Guinea) and Pt. macroiis (Aru Islands) ; Nyct'imene robhisonl, a ratlicr peculiar species, is perhaps most closely related to A', ganinus (New Guinea) and N. hdluJo' (Woodlark Island) ; and iiyconyderis auslralis differs only in -trivial characters from S. crassa, a species which, differentiated into several local forms, ranges over New Guinea and its satellites, west to the Amboina group. Two Australian species of Pkropus, scajndatus and hvunneus, have, so far as known, no equivalent in New Guinea, but related forms inhabit the Solomon Islands {luoodfonU; cohnus and nolomonis). And one species, Pteropus goiddi, points toward the Lesser Sunda Islands and Celebes, being very closely allied to Ft. cdecto. No Fruit -bat has been recorded from AVestern Australia, south of tlie Kimberley Division, or from Southern Australia, west of Melbourne ; and the single record (by Temminck, but contradicted by Gould) of a Fruit-bat from Tasmania {Pteropus ]JolioccpTudus) remains unsupported by recent evidence. Polynesian sulreicept for one species inhabiting the Pl\ili])pincs ; farther south and east in Polynesia (New Hebrides, Fijis, Samoas) it is replaced by the related and purely Polynesian Pt. samoensis group (three species). Each of the eighteen species is (so far as knowu) confined to one group of islands, except Pt. tjeddiei, which is common to the New Hebrides and the neighbouring New Caledonia, and Pt. ionymnis, which is common to the Fijis, Tongas, and Samoas. The autochthonous genus Notoj^teris has its closest relatives in the Solomon Islands (Nesonyctcris) and the Eismarck Archipelago and Ncm' Guinea ( Melon ycteris) : it has thus far been recorded from the Carolines in the north, and New Caledonia, New Hebrides, and Fiji Islands in the south.


XCU SYNOPSIS OF GEUEKA. Indications are not wanting that the Megachiropteran fauna of the Carolines, spread as it is over many islands covering a large area, may fall into at least two sections. Thus, the Pteropus marianyius group is in the extreme west represented by one species (yapensis), in the extreme east by a clearly distinct species (ualanns) ; iihe Pte7'opus j^selaphon gronp has so far been recorded only from two places in the Carolines, the Ituck Atoll and Mortlock, but specimens from the former {insularis) seem to differ from the single available specimen from the latter (phceocephalus). v. sruopsis of the sttbfamilies akd genera of Megachieopteka. I. Molariform teeth not multicuspidate ; lower canines not proclivous. A. Tongue simple : tixed to floor of mouth by posterior half, and without uufriuged filiform papillse at tip (figs. 65 A-H, [p. 1. p. 725) I. PTEROPODIN^, a. Facial axis of skull conspicuously deflected against basicranial axis : alveolar line, if projected backward, passing through middle or upper edge of occipital condyle or even through some point of supraoccipital [Rousetti;s section.] «'. Incisors never j — ^, lower canines normal in position, cheek-teeth ^ or (in Styloctenium only) ^ ; cranial rostrum not shortened, premaxilla? not more reduced than usual ; second finger clawed. a'. Occipital (sublambdoid) region of skull not elongate ; a short tail (third metacarpal nearly always slightly but distinctly longer than fourth and fifth). [ROUSETTUS SUBSECTION.] «'. Incisors | — -. «*. Tympanic forming a short bony auditory meatus, premaxillsa spaced in front ; m, equal in length to mj and m, combined. [3 species: Ethiopian and Malagasy regions.] 1. Eidolon, p. '2. h*. No bony auditory meatus, premaxillse in contact or co-ossified in front; m, shorter than mj and mj combined. [14 species : Ethiopian, E. Mediterranean, Oriental, Austro-Malayan.] 2. Rousettus, p 10. 6\ Incisors -^ — ^ (i' lost), premaxillse spaced in front. "1 species: Celebes.] 3, Eoneia,^. bb.


SYNOPSIS OF GENEUi. XClll l'. Occipital (sul)laiubdoid) region of skull elongate, subtubular; no tail (tifth metacai-pal nearly always slijxlitly but distinctly longer than third and fourth. [Pteropus subsection.] c^. Incibors | — ^, cheek-teeth g. c\ rrenia.xilliB in simple contact in frout ; io once and a half to .six times the built of i,, upper canines without secondary cusp (e.\cept in I'feiopus tithe) culadts). a'. No well-ditt'ereutiated antero-internal tubercle in p' and m', no sharply delined inner basal ledge in p^, ni|, and m^ (^except in Pt. imetianits). [85 species, 103 forms: Malagasy, Oi-iental, Austro-Maliiyan, Australian, Polynesian ; one Ethiopian species.] 4, Pteropus, p. 61. l>\ A well-differentiated antero-internal tubercle in p* and m' (sometimes also in p^ aiid -p^), a sharply defined inner basal ledge in p^, m„ and m„. [6 species, 9 forms : Philippines, W. Austro-Malaya.] o. Acerodun, p, 41:.'. dK Premaxilla? co-ossified in front (in adults) ; i, twelve to fifteen times the bulk of i,, upper canines with large external secondary cusp, upper niolariform teeth with prominent anterior and posterior basal ledges, external main cusp of p^, m,, and m^ distinctly bilobed. [2 species : Solomon Islands.] .... 6. Pferalope.v, p. 432. 2 2 -,5 (P. Incisors j — ^ (i, lost), cheek-teeth j (nij lost). [1 species: Celebes.] .. 7. Styloctenium, -p. AA2, U. Incisors j — j (i' and i, lost), lower canines at extremity of mandible, cheekteeth y (p^ lost); cranial rostrum at least slightly, often considerably shortened, premaxillae sublinear ; second finger without claw, a short tail, wings from spinal line, back seemingly naked (the furred back being covered by the naked wingmembranes). [12 species, 13 forms: Austro-Malayan.] 8. Bobsonia, p. 448. b. Facial axis very little deflected against basicranial axis : alveolar line, if projected backward, passing through lower edge of occipital condyle or even some distance below condyle (exceptions : Plerotes and 82)h

STSOrsIS OF GKXEUA. /. Biaiii-case peculiarly flattened pn3tcriorly ; a small whitish hair-tuft at base of ear-couch anteriorly and posteriorly (except in Scotomjct.eris). [All genera Ethiopian.] [Epomophoeus C-. Cheek-teeth \ (ur absent), molariforai .siicxio.vf.] teeth sublineav, with flattened crowns ; palate-ridges simple (tig. 29 J3, p. 48-3). [1 .species.] P. TkrMs, p. 483. d-. Cheek-teeth g fp^ m", and m.^ aVsent) ; some or all of the palateridges peculiarly modified, e^. Cranial rostrum long : orbit to tip of nasals much more than lachrymal breadth, e*. Rostrum long and broad, postdental palate flattened posteriorly. <'. Koslrum not deeper than usual, premaxilla; in simple contact antei-iorly ; lower incisors and canines biting against corresponding upp'^r teeth, outer ridge oV lower molars simple ; muzzle without cutaneous leaves, tail rudimenfary (two free caudal vertebrtie) ; males with shoulder pouches and erectile shoulder brushes. [3 species, 4 forms.] . . 10. Eiuy.nop:', p. 487. d"^. Rostrum considerably increased in depth, premaxiilje ankylosed together anteriorly ; lower incisors and canines closing some distance in front of upper, outer ridge of lower molars bilobed or trilobed; upper lip with cutaneous leaves, tail absent (no free caudal vertebraj); no shoulder pouches or [p. 501. brushes in either sex. [1 species.] 11. Ihjpsiynathus, /•'. Rostrum long and narrow, postdental palate deeply depressed [p. oil. posteriorly. [8 species, 9 forms.] 12. Epomophorm, f^. Cranial rostrum short: orbit to tip of nasals equal to or less than lachrymal breadth. ^'. I'ostzygomatic palate not (or scarcely) narrower posteriorly than anteriorly ; bead %%-itliout white markings (except for the usu.'il Epomophorine whitish tufts at base of ear-conch anteriorly and posteriorly). * e\ Postzygomatic palate at least as long" as broad ; maxillary toothrow extending backward very nearly to ventral margin of orbital


synopsis OK fJENKRA. XCV cavity; soft palate wiih a deep median ^-oove, narrowiujj puste[p•^•jA. riorly. [1 species.] ,13. M'uropteropics, /'. Postzygomatic palate almost twice as broad as loi;g ; maxillary toothrow not nearly reacliiug ventral margin of orbital cavity ; soft palate with a prominent median keel. [1 species.] 14. Xaiio/ii/rt>ris, p. 559. /('. rostzygomatic palate either much uarrow-er posteriorly than anteriorly or entirely absent ; head with poculifir white markings. [/''. Postzygomatic palate long, its lateral margins forming straight lines converging posteriorly; no whitish tufts at base of ear-conch. [1 specie^-.] 15. Scotoui/cferis, p. 563. k'. N postzygomatic palate, the mesopterygoid fossa continued forward nearly to level of posterior molar ; small whitish hair-tufts at biise of ear-concli anteriorly and posteriorly. [1 species.] . .' 16. Cti.uiii/clen's, p. 568.
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xcri SYNOPSIS OF GENEKA. L\ Incisors j— j (ii lost), i' much shortened ; upper canines with distinct vertical groove on antevo-medial siu-tace. [I species: rhilippines.] 19Ptenchirus, p. 643. t". Cranial rostrum as deep in front (at canine) as at p'' ; no secondary cusp in upper canines; tail absent. [1 species :Indo-Malaya.] 20. Mer/^rojjs, \x bib. j\ Tostoibital foramen extremely small ; prenia.\^illae ankyloeed together in front; cheek-teeth = (also p' lost); membranes from second toe, tail present. [1 species : Indo-Malaya.] 21. Dyacoptems, p. 6ol. J*. Postorbital foramen absent. k". Incisors simple (terete, crown not peculiarly differentiated), subvertical, cheek-teeth not narrow ; interfemoral and calcar unmodified. c". p^ with autero-exterual basal lube or cusp, c" Pren)axilhe in simple contact 2 2 anteriorly ; incisors ^— •[ (i, lost), i' much shortened, cheekteeth \ (only m3 lost), p' with lar^e antero external basal . , lobe; wings spotted. C^ ^ _ ,. , . (.-, I ' ' species: Borneo.] 22, Bahonycten., p. 6-4. (V. Premaxillre solidly united au2 2., 1 teriorly; incisors^— 2,iiiorm at, cheek-teeth \ (m^ and m'* lost), p^ with well-defiued antero: external basal cusp; wings (probably) not spotted. [I ^ ^, . p^^ species : Java.] . ^3. Clwonax, p. 0o8. d". p' unmodified. e'. Incisors |— g, i'^ normal, p^ and nil "^ith surface cusp: tail rudimentai-y,wing3 from second toe. [1 species: W. AustroMalaya.] 24. Thooptems, p. 662. f\ Incisors \—\ {h lost), i* much sbortened, no surface cusp in any cheek-tooth ; tail about half as long as foot with claws, wings from first toe. [1 species: Indo-Malaya.] ._^. • • • 25. renthetor, p. 665. l\ Upper and lower incisors (^— ^) proclivous, croM-ns highly differ. ' . entiated, triangularly pointed^ ^^ , ^^^^

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SYNOPSIS OF GKNKRA. clioiilc-ttPtli (-) iiniisiinlly narrow; iiit<'rrt'iiioral imt fxteiulinr!bevdiul miildlc cil' tibia, calcar and tail al)sent. [1 .«pecies : Inclo(Jiiina.j A^ Incisors „— ,, (i-, i,, i., lost), clieelctf^eth j (ni-, m., lost), lower canines close tog-etlier at extremity of mandibh ; nostrils elongated into cylindrical tubes, tcnignu with four circuin vallate pa]iill!c, tail not much shorter than tibia, winjis spottei]. [13 spcjies : Anstro-Malaya, Anstralia.] I. Tonn-ne more extensible, fixed to floor of mouth by its postei ior third, its terminal fourth or fiftli coveied above with iinf'rinuvd hlifornr papilhie (figs. 60 I-L, V.7-jr,.) f. Premaxillm not broader in upper than in lower half', infraorbital canal slun'ter; third metacarpal longest, or third. f'liuilh. and ti'th siibequal, terminal plialanx of third fing'er always mnr-li sjiorter than tliird metacarj)al. fp' and )), often unnsnally larj;e, nevtr alisent.) e'. I'remaxilhe in simple eontak-teetli much more reduced ; second finger clawed, t.iil rudimentarv or absent. //-. rrpper incisors minute, crowns not peculiarly diHevenliatfcd, lower incisors subequal in size ; interfemoral and calcar unmodified. (IncisorsF, — ", cheek-teeth '^, but individual anomalies in number of cheek-teeth of fVeipient occurrence in Macrnt/losK>i.<.) P. ]'' axis much lesj deflected, premaxilho less pi-oclivous ; p' (second premolar) much higher than ]>', l()wer incisors bilobate, subeqnaliy spaced ; fifth metacarpal nuu-li shorter than third, fourth inri'rmediate; males ivith 1-irge necktufts. [1 species: I'Uhlopiuu.] . . . . 2(j. S/i/ur-r/ds, p. G71. Nijcfimnif, p. 6yl. MAC^iWHU.OS[SIN/K, p. 723. oxvrTiaus siu'Tion. '2i>. JSo7n;c'i'i/\. p. 7i.'8. 29. Me.^a'o(ilossus, [p. 738.

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SYNOPSIS OF 6EXJ5RA. /'. Facliil axis stroiifrly deflecled, premaxillre more proclivous ; p' more reduced, little hijjbev than pS lower incisors simple, i, — i^ widely spaced; third, fourth, and tiftli n?etacnrpals subequal ; males without neck-tufts. | "2 species, 6 forms : Indo-Cliina to Solomon fslandf.] . . 30. Macroyhssm, p. 746. h-, UjSper incisors large, crown well dilierentiated, narrowly chisel shaped, i,, mucli larsjer than i, and with obliquely triangular crown ; lateral interfemoral unusually narrow, calcar rudimentary. (Incisors ^ — 5, cheek-teeth '^ or, by suppression of small nij and ni^, ..) [3 species, 7 forms : Austro-Malaya, A.ustralia.1 31. iSi/co77i/c/cn's, \^. 771. (J. PremaxilliB twice or thrice as broad in upper as in lower half, infrairbital canal long-; fifth metacarpal lonirest, terminal phalanx of third finger subequal to or longer than third metacarpal, (p' and p, minute or absent.) . [Notoptf.kis skction.] [/' . Premaxillre in simple contact in front or slightly spaced, anirular process of mandible large ; p' and pi present (cheek-teeth '^), p, smaller than m^ and m^ : tail absent, membranes from flanks and third or fourth toe, back furred, tibia much less than half the length of forearnj. {-. Incisors | — 7,; second finger clawed. (Underparts much darker than lack.) [1 species: New Guinea, liisuiarck Archipelago.] 32. Melonyderis j-. Incisors y — ^ ('i ^o^^) ' f^econd finger without claw. (Underparts paler than back.) [1 species: Solomon Islands. | //. Premaxillae coossified in front, angular process of mandible small ; p' and pi absent v'cheek-teeth -^), incisors — " (or in adults, i' being deciduous, — ), p^ the largest lower cheektooth ; tail very long, snbeqnal to forearm, membranes from spinal line and first and second toe, back seemingly naked at middle (the furred back being covered by the naked membranes), second finger without claw, tibia longer than usual (half as • long as forearm). [2 species : Polynesia.] 34. Nofopferh. p. 793. p. 78.') 33. Kesoin/cferis, p. 790.

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KEY TO GliNEHA,. XCIX II. Molarifonii teeth inulticus|iidfite, loTrer canines strongly proclivous (crossingupper canines at nearly rifjht angles) III. IIARPYIOI'remaxillaj!ietl in front ; incisors j — ; IL il'^lilA-r., p. / .*• . (i' and i, lost), lov.'ev pair rudiuseutary, uppei' canines bieuspidatej lower canines tricuspidate and situ-ited close together at extremity of nuibdible, cheekteeth ^ ; second finger clawed, tail absent, tibia uuusiially short. [1 species: Philip[p. 7i).). pines.] , .'{5. Harpi/imii/rferis *, " Knj/ " to the (jenera, husrd cniirelij on dcntid atul cmnidl characters. Ill the subjoined artificial "key" to the genera such characters of the skull ami dentition have been selected as are easily observable auil readily understood b\anybody witb an elementary knowledge of Maunuuiian osteology and odontoloy,y. liut the siniplitication of the key bas liad to be l)Ouabt by sacrifice of the seriid ariangenient of the genera adopted in this Catalogue. [A complete key to the genera of Megachiroptera based entirely or chiefly upon their external characters is an impossibility ; a large number of genera, even though they may belong to widely distinct groups of tlie (Suborder, cannot lie discriininatud with certainty from eacli other without an examination of tlie skull uiul dentition.] a. No lower incisors JS'jc/imeiu; p. 081. ii. One or two pairs of lower incisors. a'. One pair of lower incisors. «-'. One pair of upper incisors. <('. Cheek-teeth ".on each side llitrpi/iuiii/cfcris, p. 799, h^. Cheek-teeth i on each side /J.///,soj/./(/, ii. 118. ' * t-. Two pairs of upper incisors (middle pjiir deciduous in Notopteris). c'K Cheek-teeth '. on each side iWsoHmVici.s-, p. 790. (/•'. Cheek-teeth less than j, on earb side. <(•' Cheek-teeth .. o «\ Length of rostrum (orbit to tip of im.sals) greater than lachrymal breadth (across lower edges of hicbrymal foramina); 111' (penultimate upper cheek-tooth) equal in size to yi^ (iintepenultiinate) UtijloctcuiKiii, p. 112. b''. Length of rosliiim much less than Ih( lirymal breadth : ni' much smaller than p' Balitiii/iftcrit:, \'. iibi. b'. Cheek-teeth t. o ('. Hostruni long, strongly deflected: lowir incisors widely spaced ; reduced luimber of cheek-teeth due to loss of anterior premolar above and below (p' and p,) . . Xolapteris, p. 793. '!''. Ivostrum short, scarcely deliected ; lower incisors close together; ledured number of cheek-teeth due to loss of posterior molar above and below (mand iii^). <(''. I'ostorbita' foramen (through base of postorbital process) large. * Ou the affinities of IIar/)i/iniii,rttr>f. sec p. SUiJ.

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KEY TO OtKEKA. a'. SurCace cusps in in and ui] (third and fourth lower cheek-tooth; ; upjier canines grooved I'tciK/vhlrus, p. 613. b^. Ko surlaco cusps in check-'tteth ; upper canines without groove Meyarops, p. 154(3. t". Post orbital foraiuen absent J:'entl(etvr, i>. Wo, b'. Two pairs of lower incisors. C-. ihm pair of upper incisors Boneia, p. 55. rf-. Two pairs of upper incisors (outer pair often deciduous in Mpuniops). £••*. o upper cheek-teeth on each side (anterior tootii otteu deciduous in i'tempus and Acerodon). c^. A short bony auditory meatus Eiilvlvii, p. -. d''. No bony auditory uieatus. e^. Occiput elongate, subtubular. c^. Outer lower incisor not more (but g."nprally less) than six times the bulk of inner. c'. No well-defined an tero internal tubercle in p'' and in' (third and fourth upper cheek'tootli) Fttni^nis,}'. t'A, d'. A well -defined antero internal tubercle in p^and m' Arcrodmi, y, 112. d''. Outer lower incisor twelve to fifteen times the bulk of inner I'fcralDpe.v., \u Sni'2, f''. Occiput not elongate. «'". Length of rostrum (orbit to tip yf nasals) subequal to lachrymal breadth (across lower edges of lachrymal foramina) M,ijouijCtel-is, \^. o~G, f'\ Length of rostrum conspicuously greattr than lachrymal breadth. €•. Outer lower incisor much higher than ; p. 771. inner .Svco/(//;- ,,. rt", Clieek-teetli not linear •< -,, , -^ -.,^, 6». Cheek-teeth linear. ^ di..M,jjru ,!>> , p. /.s. «". Premaxilla' much broader in upper than in lower half IiUhnij/vtcris, p. 7tio. 6''. Premaxilhe not broader above than below, a'". Lower incisors subeiiually spaced ?Jt;/'iln'//iiti!ii(s, p. 738. b'". Inner pair of lower iiici?ors widely spaced Maci-Kiilosiua. p. 7J<'. ,/"''. \jes> than o upper cheek-teeth on each side. (,-'. 4 upper cheek-teeth on each side. ij". t) lower cheek-teeth on each side I'Urtitci-, ji. 483. h^. o lower cheek-teeth on each side. /)''. Postorbital foramen (through base of postorbital process) large. J'. Rostrum short, scarcely deflected ; inner secondary cusp in u)iper canines ('j/uujjffru.':. \i, oS<^. h'. liostrum long, strongly deflected ; no secondary cusp in canines Hijconi/ctci-is (jit.). p. 771. Ifi. Postorbital foramen absent. /''. Incisors proclivous, highly differentiated, triangularly pointed Sphferias, p. 671. * As pointed out elsewhere (p. 733), it would be difficult to give any reliable (and practically useful) cranial or dental character by w hich Euui/ctefis spelcea and iiicijnr (K. ronfiiherrfi differs by the loss of m:i) can be discriminated from cnij/ s])ecies uf lioHsetlns. although these two genera belong tc difleicnt .subhimilies. Externally they are easily distinguished {Suusetlus with. Eonj/clcris without, claw in second finecr).

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KKY TO GKNERA. CI J', Incisors subverticiil, simple. k'*. Surface cusp in third ami fouitli lower clieek-tooth (p4 ami iiiv) . T!ioiiplerit.t, \> 002. (l\ No surface cusp iu any clieektooth Cliirututj:, p. (j58. f^s 3 upper [and 5 lower! clieek-teeth on each side, (•'i Length of rostrum (orbit to tip of nasals) much griater than lachrymal breadth (across lower edges of lachrymal loramina). t''. Post dental palate deeply depressed posteriorly Epiiimiji/iui-iWi \K oil: /•>. Postdental palate flattened posteriorly. X''. Outer ridj^e of lower molars simple ... Eputimps, \t. IS7. IJ. Outer ridge of lower molars bilobtd or trilobed UjjpsiyiwthaSi \). 'y)l. J'', Length of rostrum less than or subequal to lachrymal bre.idtli, jI^ Palate not extending beyond tootlirows ^choanie at level of la^t molar) ... Cuaiiij/'.-lci'i.s, p. 5(i8. /'', Palate extending behind tootli-ro«s. m''. Premaxilla; fused anteriorly Dj/acvjitenis, \i. Hoi. ii/. l'r( inaxilla; separated by suture anteriorly. c'*. I'ostzygoniatic palate much narrower [losteriorly than anteriorly . Sr(itiiii,i/(ieris, p. o''3. y*. Post/.,\gon!atic jialate subequal in breadtli throughout. c^. Upper tooth rows extending l)ractically to ventral margin of orbital cavity MirrojiternpiiSy p. ooi. d'^. Upper toolh-rows not nearly reaching ventral margin of orbital cavity l^'unuiiyclcvie, p. .1.59.

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Order CHIROPTERA. Suboider I. MEGACHIROPTERA. Differential characters, as compared with tlie suborder Microoliiroptera. — Second digit: retaining an evident degree of independence from third, its ungual phalanx present, though the claw is sometimes wanting. Deltoid crest of humerus low ; tuhercidani mnjits and t. minus small, the former never articulating with the scapula. Facial portion of skull vnriable in length, hut never specialized in form ; cochlea small, basioccipital and basisplieuoid not narrower than usual ; glenoid fossa of sqiramosal unmodified ; angular process of mandible broad and low, or practically absent. External ear not specialized, tragus never present, margin of ear forming a complete ring. Range. — Tropical and subtropical portions of Old "World, including the whole of the Malagasy region, and extending to Australia and Polynesia, as far east as the Carolines and Samoa, but not to Xew Zealand. Fnniily PTEROPODID.i:. Sul)faiMily I. PTEROPODIN.E. Differi'ittiiil cliaractfrs. Tongue not highly extensible. liioi.-o;j tioriual. Ramie. — The same as tliat of the Megachiroptcru [lupra).

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1. EIDOLON, liojin. Ci/non)/cicris (pt.), Dobson, Cat. Chir. B. M. p. 70. Type. 1815. Eidolon, Rafinesque, Anah/se de la Nature, p. 54 . . E. lielvum. 1861. Pterocyon, Peters, MB. Akad. Berlin, p. 423 E. helvum. 1881. Leipouyx, Jentink, Notes Leyden Mies. iii. p. 60 (Jan. 1881) \_nec Liponyx, Vieillot,18\Q, a genus of birds]. E. helvum. 1882. Liponyx, Forbes, Zool. Record, xviii. (lor 1881), Mamin. p. 13 (nom. emend.). Diaf/nosis.— Basicranial axis di.stinctlv deflected ; occiput not elongated and tubular ; a short bony auditory meatus ; palate much broader posteriorly than between canines ; length of rostrum much greater than lachrymal width ; front of orbit vertically above middle or posterior halt of m' ; premaxillaries separated in front. Incisors^ — ^; cheek-teeth ^ ; p' in cross section much larger than an upper incisor ; m^ equal in length to m, and m^ combined. Second digit clawed ; membranes from sides of back ; a short tail. Forearm 114-132 mm. Skull (fig. 1).— Brain-case moderately deflected : alveolar line projected backward passing very nearly through bases of posttympanic and paroccipital processes and upper margin of occipital condyle. Occiput not (as in Pierojms) produced backward and downward into a conspicuous tube. Tympanic elongated externally into a short tubular bony auditory meatus (a peculiarity unique among Bats). Palate broadest between m'^-m^, breadth at palation border much greater than between canines, and about equal to breadth between inner sides of p* (cf. Pteropvs). Rostrum long : length from front of orbit to tip of nasals much greater than lachrymal width of skull, subequal to (a little longer or shorter than) length of maxillary tooth-row ; front margin of orbit vertically above middle or posterior half of m^ (cf. Myonycteris). Tip of nasals nearly vertically above front of premaxillaries (cf. Pteropiis). Premaxillaries slightly, but distincti) and constantly, separated in front (cf. Boiisfttus, Myonycteris, Pteropus). Width of frontals at interorbital constriction slightly less than (or at most equal to) width at postorbital constriction. Postorbital processes in aged specimens reaching about halfway between frontal and zygoma. Sagittal crest short and low, the temporal ridges sometimes remaining .separate throughout the life of the individual ; larabdoid crest strong, projecting backward considerably beyond the plane of the supraoccipital. Posttympanic processes longer than paroccipital processes. Ectopterygoid processes small. i* i" c p^ p'' p* m^ m' Dentition (fig. 1). — r-. ^-^--^ X 2 = 34. p' and m% p, i,i^cp,p3p,m^m^_m3 ^ 'i' and m^ reduced. One skull of E. helvum in the collection (ad., teeth slightly worn ; no history ; specimen c' in the list — infrc'c p. 15) has a minute m"' on one side.

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EIDOLON. Teeth without special modifications; no secondary cusps in canines or cheek-teeth. General structure of molars: a median longitudinal groove, flanked by a higher outer and lower inner ridge each ndge rising, or tending to rise, into a cusp in front of middle ot tooth ; in the upper molars the outer ridge representa Fig. I.— Eidolon helium, cT. Bilelipi, Fernando Po. No. \. cusps 4 and 5 in the typical molar in Insectivorous Bats, the inner ridge cusps 6 and 7 ; in the lower molavs, the inner ridge cusps 1-3, the outer ridge cusps 4 and 5. Upper incisors small, terete, very nearly equal in size (the outer incisors, if anything, faintly smaller in cross section) ; crown indistinctly differentiated from shaft, subcircular in cross section, E 2

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4 EJDuLOX. cuttino;-e(lge bluut ; i'-i' widely separated (chiefly owing to separation of premaxillaries), the interspace being nearly equal to diastema i'^-c ; i'-i^ rather closely approximated. Canines simple ; cingjlum not sharply pronounced, no secondary cusps ; surface of crown as a rule marked by three shallow vertical grooves, one anterior, one internal, and one posterior. All postcanine teeth more or less separated; interspace c-p' much greater than diastema i"-c. p' small, but much less reduced than in Rousetlus, being in cross section at base of crown from three to six times the size of an upper incisor, and nearly equal to m^ ; as a rule situated nearer to the canine than to \)^ : as clearly seen on comparison with p' and p'', tlie crown of yj' is formed by the completely fused cusps 4 and 6, a slight f sometimes obsolete) depression on its posterior face representing the remnants of the median groove, p' almost caniniform, its princi])al cusp (formed by the anteriorly completely fused cusps 4 and 6) from one half to two thirds the height of the canine, acutely triangular, sharply pointed ; a vertical furrow on its posterior face corresponding to the longitudinal groove in the posterior cheek-teeth, p"" molariform, longer (antero-posteriorly) than broad ; outer ridge raised into an olitusely triangular cusp, much lower than p"'' ; inner ridge forming a low cusp opposite the outer cusp; the inner bases of the cusps so closely approximated as to form a narrow, more or less complete bridge across the median groove, m' similar in structure to p^, but outer and inner ridge lower, more flatly rounded ; no indication of a transverse bridge, m^ small, subcircular in outline, equal to about one sixth of m\ Lower incisors similar in form and size to upper ones, in contact with each other and with canines, or separated by minute spaces ; row slightly convex ; cutting-edges blunt, without any trace of a median emarsination. All postcanine teeth more or less separated. Pj small, from twice to four times the bulk of a lower incisor, but not reaching above level of cutting-edges of incisors ; as a rule situated nearer to the canine than to p^. Principal cusp of Pg (formed by the anteriorly completely fused cusps 2 and 4) about two thirds the height of the canine, acutely triangular, shar])ly pointed : posterior face of crown marked by a vertical groove (the remnant of the original longitudinal groove), p^ rather longer than broad; outer ridge raised into an obtusely triangular cusp, lower than p^ ; the opposite cusp on the inner ridge much lower, bluntly rounded ; anteriorly the outer and inner ridges are closely approximated, in some individuals completely fused (as in p ). m^ long, with rare individual exceptions equalling or exceeding the combined length of m., and m^ ; ridges lower than in y> , more flatl" convex ; sometimes a slight indication of a transverse liridge from the inner base of the outer cusp, m^ similar in structui-e to m,, but less than half the size of this tooth, m^ very small, subcircular in outline, equal to (or smaller than) Pj. PaJateridfjes. — In E.helvvm: 4-f3-H3, i.e. four anterior, undivided, three middle, separated in the median line, three posterior.

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First ridge teiminatiug at oiclosely behind tlie canine ; second at or closely behind p' ; third at ]/ ; fourth at front of p* ; fifth at front of m' ; sixth closely behind ra" ; seventh far behind m^ ; eighth to tenth situated near jjalation border. Palate-ridges in E. sahceum and E. dupreanum not examined. Head. — Muzzle long, from front of eye to tip of nostrils almost etjual to distance from front of eye to base of ears. Nostrils separated by a deep groove ; inner margins moderately projecting. Inner margins of lips fringed with short pajnlliie. Median portioa of tongue with oblique rows of rounded papilla?, each papilla with three small, straight, 8harj)ly pointed, backwardly directed horny spines. Ears. — Tij) of eai'-conch reaching hinder corner of eye ; outer margin miich more convex than inner, tip broadly rounded off. Autitragal lobe practically wanting. Wings. — Chief characters : second digit clawed ; wings from sides of dorsum, rather closer to spine than to Hanks, and from back of first toe. First phalanx of first digit twice the length of metacarpal. Second digit subequal to third metacarpal. Third metacarpal between six and seven tenths the length of the forearm, a little longer tlian the fourth metacarpal, which is a little longer than the fifth. First phalanx of third digit equal to two thirds of metacarpal ; second phalanx subequal in length to metacarpal. First phalanx of fourth digit rather more than half of metacarpal ; second phalanx always somewhat longer than first. First phalanx of fifth digit less than half of metacarpal ; second phalanx subequal in length to first. See wing-indices, infra. First digit included in the membrane by the metacarpal and base of first phalanx. Notopatagium acutely triangular. 16-20 long fasciae in the plagiopatagium, viz. 4-G postanconeal, 12-16 preanconeal.

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Kanije. — Madagascar ; AtVicau coiitiuojit, from Stiiiiaar and Seiiegatnbia in the nort)i, to Xyasaland and Namaqiialand in the sonth ; S. Arabia. Halrhs.—On the Middle and r})per Nile, v. Heuglin found " Fleropus jjrt?»jan«H" (i.e. Eidolon helvuni) mostly in flocks, frequenting Bornssiis (ethioijicus. the fruits of which, together with those of Ficns and Curdia, seem to be their favourite food ; they sometimes literally eat themselves into the Borassus fruit, a shot bringing the fruit together with the bat to the ground. They fly by night as well as by day, their sight being apparently unimpaired even in the brightest sunshine, though their flight by day is somewhat flickering and unsteady ; by night it is owl-like, straight, and they occasionally pursue each other, making rapid turnings with audible flapping of the wings. They are noisy and quarrelsome, alighting with great uproar on their roosting-places. — In Fernando Fo F. Newton found them feeding on the fruits of C'lrrica ixtpaya and Fersira gratlssima. — The British Museum specimens obtained by A. Whyte at Mt. Malosa, Nyasaland, were found suspended fiom the upper branches of tall trees, but " they also frequent caves in the rocks, v\ here they probably breed " ; those collected by E. Seimund in Fernando Po were mostly shot in palm or pLiutain trees ; ripe embryos or new-born young were obtained in Fernando Po by the same collector between the middle of February and the middle of March. E. sabceum was seen in considerable numbers near Lahej, Aden, early in March (Col. Yerbnry) and in the middle of August (A. B. Percival) ; they frequent the tops of the tallest trees, where they collect in large bail-like clusters of 10 to 50, but are by no means easy to detect ; " were it not for the characteristic chattering that they keep up incessantly, they would probably be overlooked altogether" (Yerbury). Percival found them very noisy in the roosting-places, "squeaking and swearing, making a great fuss eafly in the morning " ; they were apparently feeding on dates is hich (in August) were just ripe ; when the dates are ripening every bunch is put into a bag made of palm-leaves, for protection against these bats and the crows. Affinities. — Eidolon is related, though not very closely, to JloKsetius, with which it was united by Dobson in the genus •' Cynonycteria." It differs from Roiisetlns chiefly in the development of a short bony auditory meatus, in the distinctly Repavated premaxillaries, the proportionally longer rostrum, the less reduced size of p', the unusually long m^, the smaller m^, the larger number of posterior palate-ridges, the increased number of fascife in the lateral wing-membrane, the much larger size, and in being restricted to the Malagasy and Ethiopian regions (including S. Arabia). In the larger size of p' and proportionally longer rostrum, it would seem to be more primitive than Rousettns, but it is on a higher level of development in the peculiar tubular lengthening of Ihe tympanic, the sejjaraliou of the preraaxillaries,

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KIDOLON DOPREANUM. / the lengthening' of m^, the reduction of m^, the considerably larger dimensions, and the more pronounced sexual differentiation m colour. The three species are closely interrelated. E. dupreanum, from Madagascar, witli its relatively longer rostrum and less modified fur structure, is apparently the least specialized form. E. sabcpum, from S. \V. Arabia, is a small-skulled and broad-toothed representative of the African E. /lefvum. Nomenclature.— In 1810 (Ann. Mus. d'Hist. Nat. xv. pp. 90-98), E. Geoffroy divided the genus Ptcrojms into three sections, viz., " Roussettes sans queue " (Pt. e(htlis, edwardsi, vulgaris, ruhricoUis, griseus), " Iloussettes a queue " (Pt. stramineas, cfgyptiacus, amplexicaudalus, marginatus, minimus!), and " Roussettes a ailes sur le dos " (Pt. paJiatus). Five years later (1815, l.s.c), Kafinesque raised these sections to the rank oP distinct genera, restricting the genus Pteropns to the species of Geoffrey's first section, i)roposing the name Eidolon for the second, and Pteronotus for the third section. Pteropus stramineus, Geoff, (i. e. VespertiJio vampyras helvus, Kerr), as being the earliest known of the tive species included by Geoffrojin the section " Roussettes a queue," may be fixed as the type species of Eidolon, Raf. (Of. K. Andersen, Ann. & Mag. N. R. (8) i. p. 432 ; 1 May, 1908.) Sgnopsin of the Speciesa. Rostrum relatively longer : front of orbit to tip of nasals equalling or exceeding maxillary tooth-row; fur longer, moii woolly, not closely adpressed ; colour darker ; forearm 1'27-131 mm. (Madagascar) \. E. dupreanum, p. 7. b. Rostrum relatively shorter : front of orbit to tip of nasals less than maxillary toothrow ; fur very short, closely adpressed ; colour lighter. a'. Skull larger: total length .54:-o-62-2 mm. ; tooth-rows longer : c-m''' 21-23'8 ; molars narrower; forearm 117"5-132. (African continent) 2. is. /teiuiim, p. 8. b'. Skull smaller : total length 51'6-5o mm. ; tooth-rows shorter: c-m* 19 2-208; molars broader; forearm 114-127. (S.W. Arabia) S. E. sabf^um, p. 1 o. 1. Eidolon dupreanum, Schl. tjFull. CynomjcUris dapreana, Dobson, Cat. Chir. B. 31. p. '/8. Cvnonycteris stramineus [nee Geoff.), Peters, MB. Akad. Berlin, '186H, p. 88o (Nossi B6). Pteropus dupreanus, Schlegel Sf Pollen, P. Z. S. 1866, p. 410 (N.W. Madagascar) ; iid., in Pollen ^v. Dam,Rech. I'aune Madagascar, ii. pp. 17, 172 (1868: X.W. Madagascar).

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b FADULOS HBLVCJSr. L'ynouyctfris dupreauii *, Peters, MB. Akad. _flf/7f«, 18(37, p, 866 ; Jentmk, Cat. Oateol. Mnmm. p. 264 (1887) ; id., Cut. Syst. Mavitn. p. 152 (1888) (types : X.W. ^ladao-ascar) ; Trouessm-t, Cat. Mamm. 1. p. 85 (1897)'; Seabra, Jorn. Sci. Math. Lisboa, (2) \. no. 19, p. 159 1 (1898). Cvnoptenis (Xanthirpvia) dupieana, TrouesMirt, Rev. SiMag. Zool. '(3) vi. p. 207 (1878). ( 'ynonycteris straminea var. dupreana, Barthit, P. Z. S. 1879, p. 769 (S.E. Mada.L'ascar). Xantharpyia dupreaua, Matschle, Megachiropicra, p. 63 (1899). llousettus dupreanus, Trouessart, Cut. Mamm., Suppl. p. 59 (1904). I'terocyon diipivauus, K. Andersen, Ami. ^ Mug. X. H. (7) xix. p. 504 (1907); Miller, Fam. ^Gen. Bats, p. 56 (1907). Differs from E. helvum in the following particulars : — Cranial rostrum relatively longer and slenderer: measured from i'ront of orbit to tip of nasals equalling or slightly exceeding the length of the upper tooth-row from front of canine to back of m"; posterior premolars and molars slightly broader ; m' comparatively longer, being conspicuously longer than p^ ; p^ and m„ comparatively larger; fur on body longer, more woolly, and not closely adpressed ; colour of fur browner (details see below) ; length of forearm as ia large sjKicimens of E. helvum, but metacarpals and phalanges rather longer, as shown by the wing-indices, p. 5. Distribution of fur on ears, limbs, and membranes as in E. helvum. Colour. — General aspect : brownish above, tawny olive below ; a tawny half-collar in adult males. — Upperside of head, back, and ribiae brownish, approaching Prout's brown ; base of hairs tawny olive. Glandular hairs on foreneck tawny, paler on the sides of the neck ; absent in the only female examined. Breast and belly tawny olive on flanks, washed with dark hair-brown in the middle. Measurements. See table, p. 16. Range. Madagascar. Cotgpcs (two specimens) in the Leyden Museum. «. 5 ad. rk. ; sliiill. Madagascar. E. Bartlett [C.]. b. cJ ad. sk. ; skull. Viuauitelo, Betsileo, Hoyal Society Madaffasear; 27 May, [P.]. l^'^^iBr.C.I.Foriyth Mcjor). c. c? aJ. st. ; bknll. Itanibelo, Betsileo, Royal Society 1* Mndagascai. H June, [l^-]' \m&(Ih:C.I.Fo)i;iith Major). '2. Eidolon helvum, Kerr. Cynongcteris straminea, Dobson, Cat. Chir. B. M. p. 77. Cliaiive-Soui'is, Des Marchais, I'uyage e>i Gidnee, i. p. 81 (1730: off Guinea Coast). " I/est-er Tomate Bat, Pennant, Sgn. Quadr. p. 362, no. 274/3, pL xx.xi. * \ ariuujly written C. drprcaiiuf and (fiiprcana. 1 .MissiM-lt C. 'Iiii>rri^!-(i.

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i:iuoi,o>' HKi.vuM. y fig. 1 (1771;: /(/., Hist. Quadr. ii. p. 55:.^ pi. lii. iij,'. 1 (.17eil) ; ((/., op. cif. 3id ed. ii. p. 308, pi. civ. tig. 1 (1793). lloiissette jaime, G. Cuvier, Tabl. Hem. cT Ilkt. Nat. p. 104 (1798). Wspertilio vaiupyrus (nee L.), var. C, Schreber, Siiug. i. p. lo4 (1774). Var. y, Gmelin, Limi. St/st. Nat. ed. 13, i. p. 45 (1788). Var. 4, Turton, Linn. Syst. Nat. i. p. 24 (180i>). Pteropiis vainpvrus, var. y, Er.vlehen, 8i/st. Heyn. An., Mamm. p. 133 (1777). Var. y, Donndorff, Zo (1792). Var. A, Bechstein, Pennant'.^ Uehers. vicrf. Th. ii. pp. G19, 733, pi. liv. fif,^ 3 (1800). llli(je>; Abh. Akad. Berlin, 1804-11, p. 78, cf. p. 84 "\'espertilio vaiupynis belviis, Kcir, Animal Ktnqdotn, i. pt. i. pp. .wii, 91, no.' 108 (1792). PtL>roovon lielvus, K. Ayiderscn, Ann. ^May. N. II. (7) xix. p. 504 (11X)7). Pieviipu.s strainintHis, Ji. Genffvuy, Cat. Mamm. Mas. Nation. d'Hist. Nat. p. 48 (1803: habitat uuknown) ; id., Ann. Mus. d'Hist. Nat. XV. p. 95 (1810 : Tini'ir, errore) ; G. Fischer, Zooyn(ma,\\\. p. 557 (1814) ; Oken, Le/irb. Naturyesch. iii. Abtli. ii. p. 9.34 (1810); G. Olivier, Beqne An. i. p. 124, footuote (1817); Desmarest, Nouv. Did. d'Hist. Nat. xxix. p. 512 (1819); id., Enrycl. Mcth., Mamm. i. p. 110, no. 143 (1S20); Temminck, Mon. Mamm. i. p. 195, pi. xv. %s. 12-13 (skull) (1825) ; Lesson, Man. Mamm. p. 112, no. 291 (1827); Gray, in Griffith's Aniin. Kinyd. v. ]). 57, no. 100(1827); Desmarest, Diet. Sci. Nut. xlvi. p. 366 (1827) ; Is. Geofroy, Diet. Class. d'Hist. Nat. ^iv. p. 702 (1828); J. B. Fischer, Si/n. Mamm. p. 86, no. 15 (1829); Wayler, Sy.«t. d. Arnphibien, p. 9 (1830); Lesson, Hist. Nat. Mamm. (Coinpl. Biiffon) v. p. 55 (1836) ; Temminck, Mon. Mamm. ii. p. 84 (1837: Sennaar, Senegal); Gray, May. Zool. ^Bot. ii. p. 503 (1838); Blainville, Osteoyr. Mamm. i. livr. 5, p. 100, At his, Cheiropt. pi. vi. fi<^. 2 (skull) (1840: .Sennaar); Wai/ner. Schreber' s Siiiiy., Suppl. i. p. 357 (1840); Lesson, N. Tabl. lieyne A??., p. 14, no. 188 (1842); Siindevall, K. Sr. Vet.-Akad. Handl. 1842, pp. 198, 206 (1843: Sennaar); Sehinz, Verz. Siiuy. i. p. 129 (1844); Temminck, Esq. Zoul. p. 54 (1853: Gold Coast); Wayrier, Schreber' s Siiuy., Snppl. v. p. 603 (1853-55); Gervais, Hist. Nat. Mamm. i. ]). 189 (1854) ; Giebel, Sduy. p. 999 (1855) ; Heiiylin, Beise N.O.-Afrika, ii. p. 14 (1877). Xanthai'pyia straniinea, Grai/, List Mamm. B. M. p. 38 (1843); Gerrard, Cat. Bones Mamm. B. M. p. 58 (1862); Fitzingcr, SB. Akad. Wien, liv. Abth. i. no. 10, p. 544 (1866: Sennaar, 13abr el Abiad, Bahr el Asrak, Kordofan) ; id., op. cit. Ix. Abth. i. Heft 9, p. 458 (1870) ; Gray, Cat. Moi.k. ^c. p. 116 (1870); liochebnine, Faiine Scueyambie. pt. 2, p. 40* (1883); Mafschie, Sdvy. D. Ost-Afrikas, p. 17 (1895) ; Thomas, P. Z. S. 1896, p. 790 (Zoniba, Nya.i mi II ill.

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10 EIDOLON HELVUM. (tVoiu Sfiuiaar aud Abyssinia to Guinea) ; id., v. d. Decken's Reisen, iii. 1, Siiitg. p. 5 (1869) ; Greef, SB. Ges. Naturw. Marburg, 1884, p. 44 (San Thoni^) ; Martinez y Saez, An. Soc. JEspan. Hist. Nat. xv. p. 339 ( 1 88fi : Fernanda Po) ; Jentink, Notes Ley den JiJiis. x. p 5'2 (1887 : Liberia) ; Monticelli, Ann. Mus. Civ. Gifnom, (2) v. p. 524 (pt.) (1887 : Somaliland ) ; Jentink, Cat. Osieol. Mamm. p. 264 (1887) ; id., Cat. Synt. Mamm.T^. 152 (1888); H. Allen, Proc. Ac. Nat. Sci. Philad. 1889, p. 336 (wing-membranes) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) no. 1, p. 15 (1889 : Eio Cuillo, Caconda) ; Bidtikofer, Beisebilder aus Liberia, ii. pp. 362, 471 (1890) ; Bre/im, Tierleben, 3 ed. i. p. 349 (1890: habits) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) no. 7, p. 173, fig. (palate-ridges) (1892) ; Matschie, Mitth. deutsch. Schutzyeb. vi. Heft 3, p. 7 (1893: Liberia, Gaboon); Bocaye, Jam. Sci. Math. Lisboa, (2) iv. no. 13, p. 4 (1895: Fernando Po, note on food) ; Pousaryues, Ann. Sci. Nat., Zool. iii. p. 256 (1896 : French Congo)'; Sjo^-tedt, Bih. K. Sv. Vet.-Akad. Handl. xxiii. Afd. iv. no. 1, p. 46 (1897 : Camerocn) ; Trouessart, Cat. Mamm. i. p. 85 (1897) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) v. no. 19, p. 137, fig. (palate-ridges) (1898: Angola, S. Thom^); Seabra, ibid. pp. 169, 169, pi, i. fig. 13 (palate-ridges) (1898) ; Socage, op. cit. (2) vii. no. 25, pp. 27, 46, 55 (1903: Fernando Po, Principe I., Annobon) ; id., op. cit. (2) vii. no. 26, p. 66 (1904: S. Thomd); Scubra, ibid. p. 103 (1904: Liicinda, Angola). Pterocyon stramineus, Peters, Jorn. Sci. Math. Li.'fboa, iii. no. 10, p. 123 (1871 : Ajuda) ; id., MB. Akad. Berlin, 1876, p. 474 (Cameroon, Cap Lopez) ; Miller, Fam. ^Gen. Bats, p. 55, fig. 7 (auditory buUie) (1907 : Liberia). Cynonycteris (Pteropus) stramineus, Marchi, Atti Soc. Ital. Sci. Nat". XV. p. 517 (1872-73) (structure of hairs). Cvnopterus (Xantbarpyia) straminea, Trouessart, Rev. S{ May. Zool. "(3) vi. p. 206 (1878). Rousettus* stramineus, W. L. Sclater, Mamm. S. Africa, ii. p. 109 (1901: Namaqualand, Mashonaland) ; Thomas, in H. H. Johnston's The Uganda Protectorate, i. p. 422 (1902) ; Thoma.s, P. Z. S. 1903, 1. p. 295 (Khartoum); id., P. Z. S. 1904, ii. p. 187 (Fernando Po ; sexual colour-difference) ; Trouessart, Cat. Ma7)im., Siip2)l. p. 59 (1904) ; H. H. Johnston, Liberia, ii. p. 690 (1906). Rousettus (Pteropus) stramineus, Anderson ^de Winton, Zool. Egypt, Mamm. p. 91 (1902 : Upper Nile). Yespertilio caninus {nee Blmnenb.), var. b, Goldfuss, Veryl. Naturbeschr. Sauy. p. 98 (1809). Pterocyon paleaceus, Peters, MB. Akad. Berlin, 1861, p. 423 (Africa). Pteropus moUipilosus, IZ". ^//ew, Proc. Ac. Nat. Sci. Philad. 1861, p. 159 (1862 : Gaboon). Pteropus palmarum, Heuglin, Leopoldina, Heft v. nos. 3-4, p. 34 (June 1865: Middle and Upper Nile); id., Reise N.O.-Afrika, ii. p. 15 (1877). Xantljarpyia palniarum. Fitzinger, SB. Akad. Wien, Ix. Abth. i. H. 9, p. 456 (1870). Xantharpvia leucomelas, Wagner in Hit. {fide Fitzinger) ; Fitzinger, SB. Akad. Wien, liv. Abth. i. H. 10, p. 544 (1866: Sennaar, * Variously spelt 7iV?(^'f/')'«s or Eotissettits. , ,, .,•,..

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EIDOLON HELVOI. 1 1 Jiiilir el Abi;id, I'.alir el Asrak, Kordol'aii) ; ul., op. cit. Ix. Abtli. i. H. 9, p. 401 (1870). Leiponyx biittikoferi, Jentink, Notes Leyden Mus. iii. p. 59 (Jnn. 1881": Liberia); id., op. cit. x. p. 53 (1887: Liberia); id., Cat. Syst. Mamvi. p. 152 (1888: Liberia); BiHtikofer, Hei.sebilder am Liberia, ii. pp. 362, 471 (1890) ; Trouessart, Cat. Mamm. i. p. 89 (1897) ; Matschie, Meyachiroptcra, p. 85 (1899) ; Troziessart, Cat. Mamm., Stippl. p. 04 (1904). Pterocyon biittikoferi, Miller, Fum. Sj Gen. Bats, p. 56 (1907). The characters of this species, as compared with E. dupreanwa and sabceum, are pointed out on pp. 8 & 15. Fur. — Fur on head, back, breast, and belly short, closely adpressed, on neck longer, more woolly. Face in front of and below eyes almost naked. Ears naked posteriorly except at base. Above, humerus, proximal two thii-ds or three fourths of forearm, femur, and tibia clothed with short, adpressed hair ; interfemoral hairy in the middle and for a broad space along the tibiae, extreme lateral margin naked ; u})perside of feet short-haired. Below, humerus, and proximal half of forearm, membrane along inner and outer side of forearm, lateral membrane next to body, femur, proximal half of tibia, interfemoral in the middle and along sides of tibia3, short-haired. Colour. — General aspect : grizzled straw-yellow and hair-brown above, with a more or less pronounced tawny half-collar in adults. Sexual difference : females lighter than males, half-collar paler. Adult male — Base of hairs of tipperside almost buff, broad ti])s hair-brown ; the colour of the hair-tips predominant, giving the Avliole of the upperside a darker aspect than in females, darkest on the head, pO'^terior part of back, and upperside of tibia?. Haltcollar tawny, brighter than in females. Breast and belly yellowishbuff on sides, washed with hair-brown in the middle. Adult female — Base of hairs of upperside buff, tips hair-brown ; the colour of the hair-bases predominant, giving the whole of the upperside a lighter, more buffy aspect, grizzled with hair-brown on the crown, middle and posterior part of hack, and ujtperside of tibia\ Half-collar less pronounced, rarely brighter than ochraceous buff. Breast and belly buff on sides, grizzled with hair-brown in the middle. ^'oung and immature specimens are rather similar in colour to adult males, but generally of a somewhat darker shade, and with no tawny colour on the neck. Se.vual difftrence in size. — Inconsiderable ; in 15 fully adult males the length of the forearm ranges from 117"5 to 129 mm. (average 122-2), in 14 fully adult females from 119 to 131 (average 124-7). Measurements. On p. 16. Ramie. Africa : from Somaliland, Sennaar, and Penegarabia in the north, to Nyasaland, Mashonaland, and Namaqualand in the sou til. Tiipc probalily not in existence.

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12 KIDOLOX HKLVrM. VespertUlo vainjjtjrLits Jielvus, Kerr ; 1792. — The species of Eidolon described above was well known to the earh' post-Linneari systematists, who put it dowu as a variety of VespertUio (or Pterojnis) vamptjrus, L. The earliest recognizable figure aud description appear to he those given by Pennant, in 1771 (I. s. c), under the name " Lesser Ternate Bat," so called because Pennant considered it a lesser variet)' of Seba's " Canis volans Ternatanus orientalis." Kerr's V. vampyrus Jielvus is based on Pennant's description and figure of this bat. Type originallj' in Museum Leverianum ; no habitat given by Pennant, nor by Kerr. Senegal may be fixed as the type loculity of E. helvmn. Pteropus stramineus, Geoff. ; 1803. — Eased on three examples in the collection of the Paris Museum, viz., two specimens (nos. 92 and 93) from unknown locality, presented by Professor Fourcroy, and one (no.94)"de la collection du Stathouder," this latter wrongly supposed by Geoffrey to be the original of Seba's description and figure of Canis volans Ternatanus orientalis (Thes. i. pp. 91-92, pi. Ivii. figs. 1, 2). None of the specimens now in the collection of the Paris Museum can bo pointed out, with certainty, as the true cotypes of Ft. stramineus. Four characters in Geoffrey's description of this bat are, however, decisive evidence that his specimens were E. helvum, viz., upper incisors ' tres-ecartees " ; a short tail (10 mm.) ; total length 150 mm., expanse 640 mm.; colour '• jaunatre."— The " Cat. Mamm. Mus. Xation. d'Hist. Nat." (1803) was suppressed by Geoffroy himself, and the name Pteropus stramineus is therefore usually dated from his well-known paper in Ann. Mus. d'Hist. Nat. XV. (1810). In this latter Geoffroy based Pt. stramineus on two specimens, the one stated to be from Timor (Peron and Lesueur), the other without details (and possibly one of the cotypes from 1803). The error as to the locality of the former of these specimens was pointed out by Temminck (1837, I. s.c). Pterocyon paleace^i,s. Pet.; 1861. — Type locality, Africa ; no type. Owing to Geofl^roy's statement that Pteropms stramineus had been obtained in Timor by ]\^ron and Lesueur, and evidently unaware of the fact that this error had been corrected by Temminck, Peters proposed for the African species the name Ptcrocifon paleaceus. Eut Pt. stramineus was in reality not from Timor, but from Africa; and the very species named by Peters ijaleaceus. Pteropus mollipiilosus, H. Allen ; 1862. — Type locality : " W. Africa"; as belonging to DuChaillu's collections, the type is no doubt from Gaboon ; type presumably in the Philadelphia Museum. The essence of the original description is this : a very small tail present ; first upper premolar larger than incisors ; third digit 8". Eidolon hdviihi is the only known Afiican bat for which the combination of these characters holds good. Pteropus pahnarum, Heugl. ; 1865. — Type locality: "Am mittlern und oberu weissen Nil und zwischen Senar und Fazogl Ijings des blauen Flusses"; type not in existence (?) (see Heuglin, Eeise in N.O.-.\frika. ii. p. 15). DiflTers, according to Heuglin, from Pt. f/ranuif'is " durch Mangel eines Sohwanzrurjiment? luid mit

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EIDOLOX HELVUM. 13 Ausnahmo dcr Basis gauz frcieii Daumen."' The pollex is in E. helvum always included in the membrane only by the metacarpal aud the extreme base of the first phalanx. That the specimen had no tail is no doubt a mistake ; if, when skinning an Eidolon, the tail -vert ebrnc are extracted together with the body, the empty tailskin shrinks so raiTch that the specimen seems to be tailless. Xanlharpyia leucomeJas, Fitz. ; 1866. — In Schreber's ' Saugthiere,' Suppl. i. p. 358, footnote (1840), Wagner describes a female of '^ FUropus stramineus" (presumably in the Munich Museum) " aus den oberen Xilgegenden." Even if Wagner had not recorded the sex of this specimen, it would be easy to see, from the description of the colour of the fur, that it is a female of E. helinim. Fitzinger, being unaware of the sexual colour difference in this species, ])roposed for the specimen referred to by Wagner the name X. leucomelas. A skin in the British Museum (specimen c in the list below) belongs to the same series as the type of X. leucomelas. Leiponyx huttUcoferi, Jent. ; 1881. — Type locality: St. Paul's River, Miilsburg, Liberia; type in the Leydeu Museum. Chief characters, according to Jentink : postcanine teeth ^; second digit without claw. — I have examined the t3'pe in the Leydcn Museum, and find it in every respect indistinguishable from E. helvum, nor can I see any difference between British Museum specimens obtained at places near the type locality of Z. hiittlkqferi (Nigeria, Ashantee, Dahomey)and specimens from other places of Africa; the considerablo amount of individual variation in this species, in external dimensions and in the size of the skull ai;d teeth, is well shown in a British Museum series of fifteen adult individuals fiom Fernando Po. — The teeth in the type of L. hiittikoferi are excessively worn, some of the posterior molars entirely lost and their alveoli filled out. This explains Jentink's statement that the number of cheek-teeth is -., a result evidently arrived at as follows : — Upper jaw, left side : p\ p* ((hese premolars entire), two roots of p^ (m' lost and alveolus closed ; of m* a rudiment of posterior root present, but no doubt undetectable when the skull was in situ), giving an apparent total of four teeth ; upper jaw, right side : p', p^ two roots of p'' (ra' and m" entirelj' disappeared and alveoli closed), giving similarly a total of four : mandible, left side : p,, p.^, p^ (these premolars entire), a broad interspace representing raj (disappeai'ed, alveolus closed), two roots of m., (m, entirely wanting), giving seemingly a total of six; mandible, right side : p^, p.,. p^, two roots of m,. anterior root of m^ (])osterior root of m„ and m^ lost), giving similarly a total of six. — As to the absence of the claws of the second digits, it must be said that not the claws only but the w'hole ungual (third) phalanx is wanting ; the distal articular surface of the second jihalanx is. however, in both wings laid bare, nakedly projecting, so that the missing phalanges have undoubtedly been violently torn off. The I'rcsh condition of the wounds shows that this must have been done shortly before, or peihaps after, the death of the individual.

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14 EIDOLON HELVUil. e-f. ^ ad. sk. ; skull, c? ad. skull. 2 ad. sk. ; skull. (J imm. al. ; skull. •2 5 ad. sks. ; skull of e. Khartoum, Sudan ; Surg. -Major H. N. 15 Aug. 1902. Dunn [P.]. Khartoum, Sudan. Surg.-Miijor H. N. Dunn [P.]. Sennaar. Purchased(Parreys). 47.5.27-28. (Eepresents Xantharpyia leiicomelai, Fitz.) Ituri forest, betw. Ruwenzori Explo7.1.2.8. ration Comm. [P.]. g. Ad. skull. h. i-l. (J ad. al. 3 S ad., 1 2 ad. sks. ; skulls. S junal. ; skull. lrfad.,l.juv. sks.: skulls. (} ad. sk. skull. Mawambi and Avakubi, 2500' {R. E. Dent). Ugaya I., Victoria jS'yanza, 3000' ; 30 Dec. 1901. Ugaya I., Victoria Nyanza, 3000'. Zanzibar. Mt. Malosa, Nvasa, 5500'; Nov. 1898 (.4. Whyte). Zomba,Nyasa ; Jan. 1896 {A. Wliyte). Zomba, Nyasa ; 1 1 Aug.l898(^Cloimie). Mukimvika, mouth of Congo E. ; Jan. 1892. Gaboon. R. J. Cuniiighame, Esq. [P.]. R. J. Cuninghame, 2.7-5.3. Esq, [P.]. Sir J. Kirk [0.]. 68..3.1(>.13. Sir H. n. John97.10.1.14-17. ston [P.]. Sir H. H. John96.10.28.5. ston [P.]. General Manning [P-]. J jun. al. ; skull. 4d'ad.,52ad-, 1 2 imm. sks. ; skulls. 2Jad.,32ad., 1 d pull, sks. ; skulls. 12ad.,lpulL al. Juv. sk. ; skull. cJ ad., 2 adsks. ; skulls. (5 ad. sk. ; skull. Rev. J. M. [P.]. Purchased Higgins). H. Ansell, [P.]. Fernando Po Committee [P.]. Lewis Esq. Elloby District, Gaboon. Bilelipi, Fei-nando Po, 10 m.; 20 Feb. 1904 (£".&!mund). Bantabiri,Fernando Fei-nando Po Com4.7.1.22-27. Po, 10 m.; Feb.mittee [P.]. Mar. 1904 (£•.&/mund). Bantabiri,Feruando Fernando Po Com4.7.1.137. Po, 10 m. {E. mittee [P.]. Seiiuund). rellaVista,Principe Hon.W.Rothscbild I. ; June 1901 {A. Mocqiicrys). d'j"n.,2,jun. al. ; skull of ^'. 6 ad., S jun. sks. ; skulls. 2 ad. al. i^. cJ ad. sk. San Thome ; Nov. 1899. Roca Laura, San Thome ; April 1901 {A. Mocr/uerys). Old Calabar. Abuchi, S. Nigeria, 140 m. ; 19 Nov. Asaba, S. Nigeria. NearGombi.N.Nigeria,12°N.10°E. [P.]. A. Mo(;querys[C.]. Hon.W.Rothscbild [P.]. A. Murray, Esq. A. J. Braham, Esq. [C.]. 70.3.2iM4-15. 2. Dr. W. H. Crosse [C. & P.]. Expedition [P.],

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EIDOLON SAB.1:L'M. a^-b^. Jim. sk. ;

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16 ROTJSET'rrS. Measurement!^. E.dupreamrm.' JS. hdvum. I E. snh(P7im. ^fiN. Max. MiN. Max. Mix. Max. IlllU. Foreanu 1-7 Poll ex, c. u 49 2nd digit, metacarpal S8 „ 1st phalanx ! 17 2iid-3rd pbalaiix, c. u i l-^'S 3i'd digit, metacarpal ' 88 „ 1st phalanx | «^>9 , , 2nd phalanx 91 4th digit, metacarpal ; 84 1st phalanx ' 47 2nd phalanx j 54 5th digit, metacarpal I 79"8 ,, 1st phalanx j 37 2nd phalanx j 335 Ear, length from notch ,, greatest width, flattened 1 Front of eje to tip of muzzle Tail ... Lower leg . . . Foot, c. u ; 39 Skull, total length to front of premax. ...} 68 ,, width of brain-case at zygomata...' 22 ., zygomatic width 328 ,, width acro>s m-, externally j Ifi „ width across c, externally 11'7 ,, palation to incisive foramina ' 28-2 „ front of orbit to tijD of nasals I 21'7 Mandible, length 45-2 Upper teeth, c— ni^ 21'2 Lower teeth, c — m, 24 131 49-8 63-2 19-3 16 89 59-8 93 5 87 47-8 54-8 84 40 40-7 '59 40 (iO 23 34-7 17-5 12-2 28-8 22-8 47-2 22 25-2 mm. 117-5 42 61-2 15-3 10-5 76-5 50-2 74-5 74 39-8 47-2 70-5 30 32-3 27-2 18-5 23-2 10 47 31-5 54-5 208 32 16 9-8 26 19 43 21 23 mm. 132 50-5 m 18-5 16-8 88-8 56-2 91 -.T 86 46 8 55 84-5 39-7 40 28 20 25 15-5 52 38-5 62-2 23-5 36 18 11 30 22-8 49-5 23-8 26-2 mm. 114 42 51 15 12-8 72 49 76 72 39 45-5 70-7 30 30-8 26 8 19-5 21 15 47 30-8 51-5 20-5 30 15-8 9 25-4 188 41 19-2 mm. 127 47 68-8 105 15-5 84-5 57 87 80-5 44 52-8 78 35 36 27 20 24 15-5 52 36 55 22 322 17" 10-5 27 21 43-8 20-8 23-8 L\ ROUSETTUS, Grcuf. Cyuoiiycicris (pf.), Dob.soii. Cat. Chir. B. M. p. 70. Type. 1821. Rou.«ettui?, Graxi, London Medical liepository, XV. p. 299 (Apr. 1, 1821) R. fe?yptiacn!=>. 1829. Cercopteiopus, Bnrneft, Qvati. Jmirn. Sci. Lit. Art, 1829, pt, 1, p. 269 (Apr.-.Tune, 1829) I^PRg-yptiacus. 1843. Eleuthenna, C?;Y/y,i?s!^ Mamm. B. .If.p. xix. Nonien luiduni. 1843. Xanthaq)Yia, Grau, LiiY/?/, Vvyafie ' Sulp/mr,' i.T^.29. R. leachi. 1852. Gvnonycteris, Peters. Beise Mossa7nb., Zool. i. 'Satiff. p. 25 Rlecachi. 1870. Senonvcteri?, Gray, Ca>. .Monh. S^x: p. IM. . K. seminiulus.

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llOl'SKlTfS. Diagnosis. — Geueral shape of skull as in Eidolon, but no bony auditory meatus, premaxillaries in contact or co-ossitied in i'ront. Incisors f — H ; cheek-teeth j or ^ ; lower incisors (when unworn ) 2 J ' b b ' ^ ' bifid ; p' subequal in bulk to an upper incisor ; ra^ shorter than m, and m^ combined. Second digit clawed ; membranes from sides of back ; a short tail. Forearm 69"5-99 mm. Skull (figs. 2-4 *). — Brain-case in most species deflected very nearly to the same degree as in Eidolon, the alveolar line projected backward passing through upper part of occipital condyle or upper Fig.-. — RousiUus wifirpiiaius, rj. . C;uro, Egypt. r!Cu. 3. 12.S. 1. j. margin of foramen magnum ; in one species ( 7i'. lanosus: subgonu-s Stenonycteris') the deflection is so strong that the prolongation of the alveolar line goes through the middle of the supraoccipital, in another {R. anfjolensis : subgenus Lissomjcleiis) so slight that the liiie passes below, or througli the lower margin of, the occipital condyle ; in all species the deflection is distinctly greater in immatuv© and young than in fully adult individuals. Tympanic never elongated e.xternally into a bony auditory meatus. Ilostrum comparatively Fig. y oil p. 49, fij.'. 4 tiu p. b'l.

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18 EOrSETTUS. a little shorter than iu Eidolon; from front of orbit to tip of nasals less than length of maxillary tooth-row (except in E. lanosus, owing to reduction of cheek-teeth in this species), but much greater than lachrymal width of skull ; front of orbit vertically above middle or posterior half of m^ (cf. Myonycteris). Nasals not produced quite as far forward as front of premaxiUaries, Premaxillaries in perfect contact in front, in one species {R. angolensis) co-08sified without any trace of the original suture. Postorbital processes not reaching halfway between frontal and zygoma. Sagittal crest low or undeveloped, the temporal ridges often remaining separate throughout the life of the individual. Other cranial characters as in EidoJon. ,, i' i' c p' p' p^ m' m'' ^ o . Dentition (figs. 2-4 *),— Normally v-. 5— LJ X 2:= 34. ^ iii.cp, P3P, m^m^m, p^ and m^, p, and m^ reduced ; p^ permanent in most species, deciduous in li. seniinudns and brachyotis. A minute m^ occasionally present on one side or both sides t. In one skull in the collection (R. anr/olcnsis, S ad. ; G.12.4.5) a small supernumerary molar (very broad, but excessively compressed antero-posteriorly) is present on the right side, closely wedged in between m' and m'^. Upper incisors equidistant, or i'-i' slightly more separated than i^-i^. Upper canine and p' generally widely separated, sometimes rather closely approximated (individually in R. ampler icaudatus and, particularly, R. brachyotis), very rarely in actual contact (occasionally in R. brachyotis when p' is wanting), p' much more reduced than in Eidolon, being in cross section only equal to (or smaller than) an upper incisor, in some species deciduous (R. s'^mlnudus, brachyotis). m^ much smaller than ra', but not reduced to the same degree as in Eidolon. Lower incisors subequal in height and bulk, or the outer ones slightly larger in cross section ; cutting-edge in youngish individuals distinctly bifid ; the median notch of the cutting-edge continued for a short distance down the front of the crown as a faint vertical groove ; this groove is usually detectable even -when the cuttingedge has been worn straight, p^ small, in most species from twice to four times the bulk of a lower incisor, in one species {R. angolensis) not, or scarcely, exceeding a lower incisor ; as a rule situated nearer to the canine than to p^. m^ nearly always exceeding (sometimes only equalling) p^ in length, but never quite as long as m and m^ combined, m, smaller than, sometimes only half the size of, m^. m^ much smaller than m.^, elliptical or subcircular in outline. The molariform teeth, above and below, are uuusually narrow in R. lanosus ; an approximation to this is found in R. celebensis. * Fig. 3 on p. 49, fig. 4 on p. 52. t Details from 88 skulls examined, representing all species known : — m' present on one side : R. leacAi, cue adult, teeth well worn ( ; /?. aewimtdiis, one adult, teeth unwora (nut registered) ; m-* present on both sidf s : R. (pgijptiacuii, one adult, teeth practically unworn (

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BOTTSETTUS. 1 9 ruhtti'-riihjes. — Slightly varying according to species and individuals: — (1) In most species {11. Imchi, arahicis (soinctimes), leschenavlti, semini(diis, anijjJexicaii.ilatns, celebensix) 4 + 3-)1 ; first, ridge terminating at p\ or between c and p' ; second at p^ ; third at front of p^ ; fourth at front of m', or betAveen p^ and m' ; fifth afc m", or between m^ and nv ; sixth more or less closely behind m' ; seventh far behind m" ; eighth situated at palation border. Sometimes the fourth ridge is interrupted in the median line, the formula being 3-}-4-f-l ; this I have seen in B. lanosus, IL hraclnjotis, and some individuals of li. leachi, and according to Bocage it is (normally ?) the case in R. angolensis. A ninth, more or less indistinct, ridge is occasionally detectable (formula 4 + 3-1-2, or 3 + 4 + 2 ; see R. angolensis). (2) In some species (R. agj/ptiacus, often in arahicvs, rarel}in amplfxicau(fatic) an additional divided ridge is developed behind the sixth; formula 4 + 4 + 1 ; but the extremities of this ridge are either connected with, or situated closely behind, those of tlie sixth ridge. Head. — Muzzle proportiotiall}' somewhat shorter than in Eidolon ; measured from front of eye to tip of nostrils shorter than (only in R. lanosus almost equal to) distance from front of eye to base of ear. Nostrils separated by a deep groove ; inner margins moderately projecting. Inner margins of li])S Iringed with short pa])il]8e. Median portion of tongue with oblique lows of rounded papilla?, each papilla with three small, straight, sharply pointed, backwardly directed horny spines. Ears. — Tip of ear-conch reaching hinder corner of eye : outer much more convex than inner margin, tip rounded ; in some species (7^. arabicu-t, angolensis. htnosus) a shallow, but distinct, emargination of outer margin below tip of ear. Antitragal lobe small, flatly rounded (most sjiecies) or triangular (R. angolensis), in one species (7^. hinosus) practically wanting. Wings. — Chief characters : second digit clawed ; membranes from sides of dorsum and back of first or second toe. I'irst phalanx of first digit one and a half the length of metacarpal. Second digit subequal to (a little longer or shorter than) third metacarpal. Third metacarpal from six to seven tenths the length of the forearm, a little longer than fourth metacarpal, which is a little longer than fifth. First ])halanx of third digit two third? to tlircc fourths the length of the metacarpal ; second jihalanx four fifths to six sevenths of metacarpal. First jihalanx of fourth digit rather more than htdf of metacarpal : secotid phslunx a little longer than first. First phalanx of fifth digit equal to, or a Tutle less than, half of metacarpal; second phalanx a little longer or shorter than first. See wing-indices, on p. 2
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date-plaiitatious around villajrcs, iu sycamore, mulberry, and other trees; occasionally it is found singly, but generally a few are associated togeilier ; feeds on fruits of the date-palm, wild figs, &c. Visiting Cyprus in 1901-2, iliss D. M. A. Bate found li. (pr^i/ptiacas excessively common ; it does considerable damage to the fruit-crops, particularly to the oranges and dates, though while the latter are ripening they arc often envelo])ed in sacks or matting for protection ; during the summer it roosts in thick trees, in winter in closely packed bunches in the roofs of old buildings and caves ; it is very restless, and extremely noisy, even in daytime and when undisturbed (P. Z. 8. 1903, ii. p. 342). Ji. leacM has repeatedly bred in captivit)(Zoological Gardens, London and Cologne). During the act of copulation the male is suspended (as usual by thn hind feet) behind the female, its underparts being iti contact with the lower back of the female; an embracing with the anterior extremities does not take place ; period of gestation (in one case, Cologne gardens) precisely 1.5 weeks after last copulation ; during the first months after birth the young bat was rarely seen, suspended as it was from the mammfe of the female and completely covered by her wing-membranes ; by tlie end of the third month the young commenced to take part in the meals of the parents (the juice, not the flesh, of moist fruit), but remained in its protected po-itiou at the breast of the mother till the end of the fourth month ; at the age of eight months it was still not quite full-grown (Wundcrlich, Zool. Garten, xxxii. pp. 7881, 1891). — According to Dr. J. Anderson, j-oung of R. cer/)/j)tiacHS are born generally in February and Jlarch ; a female of M. arabicns killed on March 29 at Lahej, Aden, had a single young one at her breast, and other gravid females had single foetuses (Yerbury & Thomas, P. Z. S. 1895, p. 545); a female of ii'. amphxicmdatus obtained by A. Everett in Alor Inland, N. of Timor, had one young on March 24 (collector's label, British Museum). Scarcely anything is known about the habits of the Oriental and Austro-Malayan species. Hodgson's account of the habits and extensive niglitly migrations of " Fferojms pyrivorus" i. e. B. lexcJienanlfi, in Nepal (J. A. S. B. iv. p. 700. 1835) — copied by Hutton, P. Z. S. 1872, ])p. 693-94, under the head of Cynopterus marginatiis, and by Dobson, in his Catalogue, under Cynonycteris amplexicaudatvs, and again, by an oversight, under Cynopterus marginaf^'s — is based wholly on a misconception, as pointed out by Scully (J. A. S. B. Ivi. pp. 237-38, 1887). Dobson was informed that a colony of 7^. lesrhi'navlti living near the sea at ifoulniein were seen to feed on Mollusca left exposed by the tide (J. A. K. B. x)ii. p. 200, 1873) ; this would seem to explain the occurrence of an allied species (7?. arahicns) in so dreary a desert as the island of Kishm, Persian Gulf, where an exclusively fruit-eating mammal could hardly exist. Affinities. — Rousettus is allied (though not very closely) to EiloJ.on ; the two genera represent no doubt diverging branches from one common stem. In having no bony auditory meatus, the

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-KUUSETXUS. jjreinaxilhiries in coiir.>icL (not separated), lu, not lengthened, and lU" less reduced in size, Rousettus is less specialized tlian Eidolon ; but it is on a higher level in the rather shorter rostrum and more reduced p\ The range of the genus over the whole of the Ethiopian and Oriental regions, the close affinity of ]l. arahicus to the S. African 2i. Uachi, the absence of any representative of the genus from the whole of the Srediterraneau subregion * (except Egypt), are evidence that its origin dates back to a time when, owing to different physiographic conditions, Africa andS. x\sia were much more intimately connected than now. Eidolon is a peculiarly modified Ethiopian offshoot of the common prototype. The eleven known species fall into three natural, rather sharply separated groups (subgenera) : — (!) Subgenus Eousettus : — Brain-case moderately deflected ; premaxillaries in contact, not co-ossified (except sometimes in li. mjiiptiacm) ; cheek-teeth unmodified in size and shape ; p^ much larger in bulk than a lower incisor; wings from first toe; antitragal lobe small, but distinct. — This is the least specialized of the three subgenera of Housettus, in all essential characters perhaps the most primitive group of living Vlegacbiroptera. Range : the Ethiopian Itegion, through Egypt, Cyprus, Palestine, Arabia, 8. Asia, the Indo-Malayan and Austro-Malayan Archipelagos, eastward to the Solomon Islands. The nine species are referable to three types : — (a) K. lecichi, (egi/j^tiaai^, and arahicus : rather heavily built species, with strong rostrum and teeth, the second phalanx of third digit lengthened, the poUex comparatively long : distributed over Africa generally, Cyi)ru'5. Palestine, Syria, and Arabia, as far as Sind (Karachi), li. ivi/i/pticicns is a larger-skulled modification of the li. leachi type ; li. arahicua is more closely allied to the S. African R. leachi than to R. agyptiacm. — (b) R. lescheiiaidti, seminadus, nnrple.vicaudatus, wiinor, and hracliyotis : very closely related to the species of the former section, but rather more delicately built, wirh slenderer rostrum, feebler teeth, the second phalanx of tiie third digit not lengthened, the pollex comparatively shorter. The members of this section are probably on the whole slightly less specialized than those of the former. R. lexdiemmlti (continental S. Asia) and semiuudxs (Ceylon) come near to the S. African Ji. leachi in the width of the interspace between c and p^, the size and shape of m^, the width of the ears, and the shortness ot the tail ; in the Indo-lfalayan R. ampleA-icandatus there is a tendency to a reduction of the diastema c-p\ m^ is smaller and more circular in outline, the ears narrower, the tail averaging longer, the general dimeiisions smaller ; most of these characters * Botisettus fft/i'ltiri/i. 'froupssart ("Cat. Miuiiui., Suppl. p. fiO, 1904; baped on Cf/iioni/cteris (?) sp., Claude Gaillard, C. R. Aonrl. Sci. cxxv. p. (i20, 1897, and Arch. Mus. d'Hist. Nat. I,yon. vii., 2 meiu. p. 6, fig-. 1, 1899), known from a complete right humerus and the dista! extieinities of a risfht and lelt humerus, from the Middle Miooene of La Grive Saint-Allian, Lsere. is not a liuuseftus, nor even a FruitBr,t, but some large species of Microchiroptera, as proved by Gaillard's figure of the humerus (1. s. c. : high, flange-like deltoid crest).

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EousErius. 23 find a climax in the Austio-Malayau li. brachyotis : diastema c-p'^ still more reduced, p' deciduous, ears still smaller, size smaller. — (c) li. celebensis : palate and cheek-teeth narrower than usual, fur louf^er and richer, size small, wings proportionately long; probably a modification of the R. amplcxicaudatus type. (2) SxENOXiCTEUis* (subg. nov.): — Brain-case strongly deflected ; preraaxillaries never co-ossified ; check-teeth excessively narrow : width of p' about one fifth (in llousettus s. str. about one third) that of palate between fronts of p' ; p^ much larger in bulk than a lower incisor; wings from second toe; antitragal lobe obsolete; fur long and rather coarse. Etliiopian. One species: R. lanosns, Thos. — So far as the reduction of the cheek-teeth is concerned, this peculiar species stands in the same relation to the rest of the genus Rouset/iis as the narrow-toothed species of Pleropus (Ft. sahniger, personatus, woodfordi, sc(i2ndatus) to the normaltoothed Fteropi. If its habits were known, R. lanosus would probably prove to subsist cliiefly on food (juice of fruits) tliat requires little or no mastication. (3) LissoNTCTKRis t (subg. nov.) : — Brain-case only slightly deflected ; prcmaxillaries co-ossified in front, even in }'oung adults ; cheek-teeth peculiarly short and broad, subquadrate ; p^ reduced in size, being only subequal to a lower incisor ; wings from second toe ; antitragal lobe distinct ; fur limg and silky. Ethiopian. One species : R. aw/olensis, Boc. — This is the most aberrant species of the genus ; although in all important respects a " Rousettus," it shows, in the small deflection of the bi'ain-case, the shape of the occipital region of the skull, and the outline of the cheek-teeth, leanings towards the genus Epomophorns. Rousetius, Gray ; 1821. — Xo description. Type (only species): Pta-opus orpiptiacus, Geoff. The name Ronsctfii.s must have been suppressed or forgotten by Gray ; it does not occur in any of his later jtapers. Revived by Palmer, Proc. Biol. Soc. Wash. xii. p. 112 (30 April, 1898). Cercopteropus, Burnett ; 1829. — No description. Two species mentioned : C. (e.fjifpliacus and C. amplexicandatus. The former, as being the first-named, may bo fixed as the type. XantJunpi/ia, Gray; 1813. — Xo description. Three species, enumerated in the following order: The Xantharpye, Xantharptfia ample.vimndata ; Egyptian Xantharpye, X. wgyptiaca ; The Pale Xantharpye, A", stniminea. Both on the " first species principle " and on the " tautology principle " A'. ampJe.vicaudata must be regarded as the type of the genus. It should be mentioned that the two examples referred by Gray in this book (List Mamra. B. M. p. 37, 1843) to "X. ample-v'icavdata" are in reality not this species but Dohxonia pnllala ; however, by his reference to Geoffrey's original description and figure of Pteropus atnplexicaiidatus Gray * Irevos (narrow), ri'icrfpis (bat) ; in allusion to the I'xcessively narrow molnrs. t AiiT(TOf (soft), rvKTfpis ; from the silkv character of the fur.

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2i KousKrrus. clearly iudic-ittes wliicli species is meant bjthe name ample.vicau(lata. — A brief diagDOsis of the genus Xanthnrpiiia was given by Gray in Voyage of the ' Sulphur,' i. p. 29 {1844): distinguished from Eleutherurn by having the " tail with the base inclosed in the underside of the interfcmoral meml^rane." Eleutheriira, Grray; 1844. — First occurrence of the name in Gray's List Marara. B. M. p. xix (1843): no description, no species. Second occurrence in Voyage of the ' Sulphur,' i. p. 29 (1844) : type (only species mentioned), E. hnUentotta, Temm. ; a brief diagnosis showing that Gray based the genus on Temminck's erroneous description of Ptero'pus hottentottus : tail free (not connected with interfemoral), projecting from " a nick on the middle of the narrow interfemoral"; but Temminck's Pi. hoUeniotius is A. Smith's /'t. Jearln. Ci/noiii/cteris, Peters ; 1852. — Type (only species mentioned) : Pieropus coUnris, Licht. (nee ill.), i. e. Pteropiis Icachi, A. Smith. — The genus Cjfnoru/cterh, as understood by Peters in 1867 (Kevision of Megachiroptera, MB. Akad. Berlin, 18G7, ])p. 86.5-866), included all the then known species of liousettus and Eidolon. Dobson's Ci/nonj/ctens (Cat. Chir. 1878) corresponds to the genera Eidolon, liousettus, and Mi/oni/cteris of this Catalogue. Senoni/cteris, Gray ; 1870. — A subgenus of Xantlmrpiiia. Typo (only species) : Xantharpyia scminuda, Gray. Separated from Xantharpijia s. str. on account of its "very sparse and s'hort " fur, "especially on the back," and " nearly bald" forearm. tSi/nopsis of the Species.^ J. Brain-case conspicuously detJectfd ; preujaxilliiries in contact ; p, much larger in bulk tlian a lower incisor. A. Brain-case moderately deflected ; width of p^ about one tliird tliat of palate between fronts of p^ : wings from 1st toe ; antitragal lobe distinct Subg. lioisETTUs. a. Pollex .'iO-37-.5 : 2nd phalanx of third digit 50-5-61-5 mm. «'. l<;ars not attenuated at tip ; lower leg 40-4 So mm. «-. Length of skull 40'-'j-1.'>-8 mm. ; rostrum slenderer : palate-ridges normally 4-f 34-1 : forearm 8999 mm.' (S. Africa) }. H. k-achi. p. 2.5. b'. Jjeni;th of skull 4o-6-4()-7 nun. : rostrum heavier ; palate-ridges 4 4-44-1; foreanii 88-99 mm. (Angola to Palestiue) i'. R. (/i/i//)f;,irf/s, y. -J'.}. I'. Ears attenuated at tip ; lower leg .'17*•" •'j9-5 mm. ; forearm 87-96. (Arabia to Sind) y. Ji. unihieu.-: p. 3-1. h. P.dlex 24-30; 2nd phalanx of 3rd digit ••;(;17-2 mm. " On n. ,»/,/.>;, sve p, 4;l a.ud Acldeiidt at rnd of Voi.ii.i

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I'.uusrrms lkacui. c . Miihirt^ not iiiuisually narrow ; fur sliort: Botopatag'iuin naked. cK m., elliptical in outline, once and a half or Iwice as long' as Lioad ; width of ears 14'0-15'8 mm. ff\ p' not deciduous; fur on nape and shoulders not unusually scarce ; for'earm 80'5-87'5 mm. (India, Himalaya, to S. China) b^. pMeciduous; nape and shoulder.^ seuiinaked ; forearm 79-85/5 mm. (Ceylon) d'. m.. iu outline, only slightly longer than broad ; width of ears 10-1. 3 mm. <". p' not deciduous ; forearm 7787'2 mm. (Indo-Malayan) .. cP. p' deciduous; fosearm 69-8-76 mm. (Austro-Malayan) . . . . d'. ^Molars unusually narrow ; bony palate narrow ; fur longer ; notopatagium and tibife well haired ; forearm 72-5-75 mm. (Celebes) . . B. Brain-case strongly deflected ; cheek-teeth excessivelv narrow ; width of p' about one Kfth tLat of jialate between frimts of ])* : wings from :^ud toe ; antitragal lobe 1 'bsolete c. Fur long and coarse ; lower leg 39-40 ; forearm 88-5-90 mm. (Ethiopian) . . 11. Brain-case slightly deflected : prema.xillaries co-os?ifiod ; pi subequal in bulk to a lower incisor ; molars short and broad : wings from 2nd toe ; autitragal lobe distinct . . d. Fur long and silky; lower leg 29-31 ; forearm 79-83-5 mm. (Ethiopian) . . [p. 35. 4. li. Ifschenaulh, 5. R. seminuJus, p. 3S. LP-lO. 6. 1{. amplcxkaudatus, 8. R. iruchyotis, p. 44. 9. R. celcbensis, p. 4G. Subg. STENONYCTEniS. 10. R. hmosus, p. 49. Subg. LissoxYCTEnis. 11. R. angoknsis, p. 51. 1. Rousettus leachi, A. Sm. Cffuonycteris cuUaris, Dobson, Cat. Chir. B. M. p. 75. I'teropus collaris {nee III.), Licit rtisfei'n, J'erz. Douhl. Mas. Berlin, p. 3, no. 47, p. 5 (1828 : Terra Caffrorum) ; Giehel, Sdug. p. 1000 (1855). Cynonycteris collaris, Pctcvf:, Rrise MoRsambiqne, Sdiif/. p. 25 (1852: liiliaiiibane); id.. MB. Akad. Bfirlin, 1867, p. 865 (S. Africa); r. L. t'idnfp); P. Z. S. 1868, p. 404 (at sea oil" Natal); ojx cit. 1869, p. 602 (Natal) ; op. cit. 1870, p. 127 (fig. of $ ad. with young) ; op. cit. 1871, p. 478 (breeding in captivity); op. cit. 1873, p. 193 (specimens in captivity); Dohson, Cat. Chir. Ind. Mv.-<. p. 2 (1874 : S. Africa) ; Gulliver, R. Z. S. 1875, p. 493, pi. iv. tig. 7 (red blood-corpuscles) ; Dobson, Mo>i. A.i. 3 ed. i. p. 350 (1890) ; Wunderlich, '/.oil. (Uirtcn. .\.\xii. i)o. .'>. p. 78, toxt-fig. (I'^Ol : breeding in ciplivily t : 'I'miiv^mrl . Cat. Mmtmi. i. p. 8J ipt.) (1^07).

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26 EOUSETTCS LEAOHI. Cvnopterus colLiris, Kolenati, Moii. europ, Chir. p. 11 (18(30: Cape, Mozambique). Eleutherura coUaris, Gray, Cat. Monk. ^-e. p. 118 (1S70: Cape, Natal). Cynopterus (Cynonycteris) ccllaris, T^-ovessart, Jiev. ^Mag. Zool. _ (3) vi. p. 206 (pt.) (1878: Cape of G. H., Natal). Xantbarpyia collaris, Matschie, Megachiroj^tera, p. 66 (pt.) (1899). Koiisettus collaiis. W. L. Sc/afer, 3famm. S. Afr. ii. p. 106, text-fig. 118 (1901 : Cape, Natal) ; Anderson ^ de Winton, Zool. Egypt, Mamm. pp. 86, 88 (1902) ; Troitessart, Cat. Mamm., Sup2)l. p. 60 (pt.) (1904) ; Thomas ^Schcann, P. Z. S. 1906, p. 161 (Knysiia) ; Mtller, Fum. Sf Gen. Bats, p. 54 (1907). Pteropus am])lexicaudatus {nee Geoff.), Temminck, Man. Mamm. i. p. 260 (1827 : Cape) ; J. B. Fischer, Syn. Mamin. p. 86 (pt.) (1829: Cape). Cynouycteris amploxicaudata, Dohson, Cat. Chir, B. M. p. 73, .specimens e-j (1878). Pteropus leacbi, A. Smith, Zool. Journ. iv. p. 433 (1829: Cape); id., Bidl. Sci. Nat. xviii. p. 272 (1829 : Cape) ; Temminck, in Smuts s Enum. Mamm. Capens. p. 5 (1832: Cape); A. Smith, S. Afr. Quart. Joiiryi. ii. p. 53 (1833 : Cape) ; Lesson, Hist. Nat. 31a77im. [Compl. Buffon) \. p. 49 (1836: Cape); Temmijick, Mon. Mamm. ii. p. 88 (1837: Cane); Gray, Mag. Zool. ^ Bot. ii. p. 603 (1838 : Cape) ; tVagner, Schreber's Sdug., Suppl. i. p. 361 (1840 : Cape) ; Biippell, Mas. Senck. iii. Heft 2, p. 154 (1842: S. Africa); Lesson, N. Tabl. Eigne An. p. 14, uo. 192 (1842 : Cape) ; Schinz, Syst. Verz. Sdug. i. p. 131 (1844 : Cape) ; A. Smith, 111. Zool. S. Afr. pi. 48 (wbole fig., bead, skidl) (1847: Cape); Wagner, Schreber's Sdug., Suj)pl. v. p. 604 (1853-55: Cape, Mozambique); Gervais, Hist. Nat. Mamm. i. p. 191 * (1854) ; Schlegel, Eierkunde, i. p. 63 (1857 : S. Africa) ; Luyard, Cat. S. Afr. Mus., Mamm. p. 19 (1861). Xantbarpyia leacbi, Gerrard, Cat. Bones Mamm. B. M. p. 57 (1862 : Natal) ; Fitzi^iger, SB. Akad. Wien, Ix. Abtb. i. Heft 8, p. 466 (1869: Cape, Mozambique). RoLisettus leacbi, K. Andersen, Ami. ^ Mag. N. H. (7) xix. p. 506 (19071 ; Lonnberg, Kiliimmdjaro-Meru E.vpedilion, pt. 2, p. 6 ^908: Tanga). Pteropus hottentottusf, Tejuminck, in Smuts's Enum. MammXkqjens. p. 3 (1832 : Cape) ; id., Mon. Mamm. ii. p. 87, pl. xxxvi. figs. 16, 17 (skull) (1837: Cape); Wagner, Schrebei'^s Sdug., Suppl. i. p. 360 (1840: Cape) ; Eilppell, Miis. Senck. iii. Heft 2, p. 154 (1842: S. Africa) ; Lesson, N. Tabl. Rh/ne An. p. 14, no. 191 (1842 : Cape) ; Schinz, Syst. Verz. Sdug.'i. p. 131 (1844 : Cape) ; Wagner, Schreber's Sdug., Suppl. v. p. 604 (1853-55 : Cape) ; Gervais, Hist. Nat. Mamm. i. p. 191 (1854 : S. Africa) ; Giebel, Sduf/.-p. 1000 (1855); Grid, IC Sv. Vet.-Akad. Handl. n. s. ii. no. 10, pp. 9, 13 (1860: Knysua). Xantbarpyia bottentotta, Blyth, Cat. Mamm. Mus. As. Soc. p. 21, no. 56 71863: S. Africa); Fitzinger, SB. Akad. Wien, Ix. Abtb. i. Heft 8, p. 468 (1869 : Cape). Cynopterus brevicaudatua [nee Is. Geojf.), Gray, List Mamm. B. M. p. 39 (1843). Kousettus sjo.stedti, Lonnberg, Kilimandjaro-Meru Expedition, pt. 2, p. 7 (1908: Tanga) t. * 'Misspelt Pteropus ErachU. 1 Earliest spelling : Jwt/cntof/uf ; niifspeU hoflenfoUis by a fenof the juUiurs quoted. \ 8ee Addeui.x

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ROrsETTUS LKACHT. 27 />;(r/(K»s«'s.— Distiu^niished from all other species of the genus by the combination of the following characters:— Frontal region of skull between postorbital processes flattened; premaxillaries in co)itact, but rarely co-ossiiied ; total length of skull 40-5-43-8 mm. ; palate-ridges normally 4 + 3 + 1. Wings from back of first toe, or interspace between first and second toe; pollex (with claw) 31 -35-5 mm.; second phalanx of third digit 50-5-60 mm. ; second phalanx of fifth digit nearly always shorter than first plialanx ; cars not attenuated at tip ; fur short. Forearm 89-99 mm. Skull. — Brain-case moderately deflected (compare E. lanosus and Ji. cmgnlensis). rremaxillaries in contact in front, hut rarely, even iii very aged individuals, co-ossified (compare li. (pgijptiacus and R. nnr/oleiisis). Frontal region between postorbital processes flattened (compare R. anr/olensis) ; region between orbits cora))aratirely narrow, the width of the interorbital constriction in fully adult specimens being almost always distinctly less than the width of the postorbital constriction (compare R. cef/i/ptiacus). Temporal crests uniting into a sagittal crest at level with root of zygomata, but very often remaining separate throughout the life of the individual. Teeth. — Upper incisors almost eiiuidistant, or the interspace between i' and i' much less than between i* and c. Distance c-p' equal to or larger than between i" and c. p^ small, equal in size to i-. m' equalling or exceeding p' in length, m' rather less than half the size of m\ — Lower incisors crowded. A broad interspace between c and p,. p, in cross section from twice to four times the size of a lower incisor, situated nearer to ihe canine than to p^. m . m,, and ra , elliptical oval or almost elliptical in outline ; m.^ somewhat smaller than m, ; m^ from two thirds to less than one half the si/e of m.,. J'alate-ridgex.—'SoTmaih 4 + 3-|-l. Variations from the rule: the fourth ridge in some individuals interrujited in the median line (formula: 3 + 4 + 1); a 9lh, more or less indistinct, ridge occasionally detectable (formula : 4 + 3+2). Ears. — Outer much more convex than inner margin; tip not attenuated, broadly rounded off above (compare R. arabicus). Antitragal lobe flatly rounded. Winqs. — Pollex with claw equal in length to four fifths of second metacarpal (or equal to first phalanx of fourth digit). Second digit equal to metacarpal of third digit. First phalanx of third digit eijual to two tliirds of metacarpal; second plialanx only a little shorter than metacarpal. Second phalanx of fifth digit, with rare individual exceptions, shorter than first phalanx (compare R. avr/olensis). Wing-indices, see anted, p. 20.— Notopatagium semicircular in outline, naked ; lateral membrane inserted on end of first metal arsal, or between first and second toe. 10-14 long f.isciic in the lateral membrane, viz.. 2-3 postanconeal, 8-10 preanconeal. 7'(/(7. — From one half to three fourths the length of the hind foot. /.',„.._l.;;irs naked posteriorly, except at base. Face in front of and below ev.s short-haired. " Ftir on back short, soft ; longer.

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28 KOUSEXni.S LEAl'HI. more wooll)', and less closely adpressed on foreneck. Fur of body extending upon upperside of humerus and proximal half or two thirds of forearm ; front margin of antebrachial membrane fringed with short hairs ; notopatagium naked ; femur well haired ; tibia and hind foot to claws clothed with very short and thinly spread hairs ; central portion of interfemoral well haired, lateral portion along tibiae almost naked. Belowj proximal two thirds of forearm, plagiopatagium next to body, femur, and central portion of interfemoral covered with short, thinly spread, woolly hair. Colour.— General aspect : above, uniform dark brown with a tinge of slate, or more or less suffused with Prout's brown ; below, uniform smoke-grey, or more or less suffused with wood-brown. Eack and rump varying from dark hair-brown tinged with slate to bistre ; sides of back and rump next to membranes, in adult specimens of both sexes, often approaching Prout's brown, this colour sometimes extendiug over the whole of the upperside of the body ; crown and occiput markedly darker than back ; nape of neck lighter hair-brown. Underside uniform grizzled smoke-grey; in adult specimens of both sexes the foreneck, flanks, and under surface of humerus and forearm are often more or less suffused with wood-brown. / Measurements. On p. 34. Range. Cape Colony, Natal, Lower Zambesi (Inhambane), north to German East Africa (Tanga). Gotijpe in collection. Pteropus collaris, 111. ; 1815. — Type locality : " die ostlichen [afrikanischen] Inseln." — llliger's Pteropus collaris (Abh. Akad. Eerlin, 1804-11, pp. 71, 84; published 1815) is Erisson's " Roussette a col rouge" (1756"), Euffon's "Eougette" (1763), Kerr's Vespertilio vampi/rus subnir/er (1792), E. GeofFroy's Pteropus ruhricoJlis (\S10). In 1823 Lichtenstein (l.s.c.) wrongly identified llliger's Pt. collaris with the S. African F'ruit-Bat here under consideration, but the error, hidden as it was in the littleknown" Verz. Donbl. Mus. Berlin," passed for manj' years unnoticed, the species being constantly referred to as Pteropus leachi or Pt. liottentottns. In 1852 Peters (' Peise nach Mossambique ') confirmed Lichtcnstein's wrong identification of Pt. collaris, and from about that year the names Imchi and hottentotttts gradually went out of fashion, being replaced by collaris : from about 1870 leachi and hottentottns only appear in the lists of synonyms of collaris. Pteropus leachi, A. Sm. ; 1829. — Type locality : " Gardens about Cape Town"; cotype in collection. — Summary of description: " supra fusco-cinereus, infra sordido-cinereus, cauda libera." The earliest available technical name of the species. Pteropus hottaitottus, Temra. ; 1832. — Type locality : " circa urbem Capensem "; type in the Leydeu Museum (authentic specimens in the British Museum, see list below, specimens a-e). — Original description contributed by Temminck to Smuts's ' Enuraeratio Mammalium Capensium.' Temminck points out the differences between Pt. hottenfofius and Pt. amj-ih.ricyjtrlafvs, but was admittedly in doubt whether the former was separable from Pt, leachi

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KOl>;KTTVS .TXJVI'TIACIS. 29 (' de Fterofio leacJn tenendum vidptiir, eiim forfasse a praecedente [i.e. Ft. hottnitotto] nou distinctum esse specie"). Owing to a passage in the original description ("ad basin caudae perbrevis eernitur incisura, figurara inversae V pi'ae se ferens ; juxta banc Cauda, quae plane est libera, exoritur "), Ft. hottentottns As-as by subsequent writers ((iray, Waguer, and others) supposed to differ from Ft. leacJii by a V-sliaped eraargination of the interfemoral at the base of the tail, and by having the tail perfectly free from the interfemoral, diB'ercnces which undoubtedly were due to shrinkage of the median part of tlie interfemoral in the mounted, type of Jioltcutoitns. a-e. 4 ad., 1 iimn. ]^ear Cape Town Purchased (Leyden, sk.s.; skulls. (J.Sinuls). Museum). 38,40,67. { Autbentie specimens of I'teropus hottentoiius, Temm.) f-h. 2 S "d-. 1 6 Cfipe Town. Trustees of the S. iuv. al.; skull Africaia Museum of./; [P.]. /. Ad. skull. Cape Town (5i> .J. H. Ford, Esq. [P.]. Smilh). (Figured in 111. Zool. S. Afr.) j. $;ul.;il. Cape Town (.Sir .-1. Surg.-Gen. G-. E. ISmitli). Dobson [E.]. ( Cotijpe of species.) k. S j"T-^k; Cape of Good Hope. Purchased (Ver43.12.7.21. skull. reaux). l-f. 4cJ'ad..4 $ ad. Knvsna. 3 April, C. 1). Rudd, Esq. sks.; skulls. iyuo (C. //. .S. [P.]. Grant), f-v. (S ad., 2 id., Kny.-5nu {C. H. B. C. D. Eudd, Esq. $ juv. al. Grant). [P.]. w. Ad. St. ; skull. Natal. Purchased (Stockf holm Museum). .r. c^ad. al. Natal. Dr. Seem;ni [C.]. y.; skull. S.Africa. Purchased (Brandt). 2. Rousettus asgyptiacus, E. Geoff. Ctjnonycteris cegypiiaca, Dobson, Cat. Chir. B. M. p. 74. Pteropus ivgyptiacus •*, E. Geoffroy, Ayin. Mus. d'Hist. Nat. xv. p. 90 (ISiO: Lower Egypt); Oken, Lehrh. Natiirgesch. iii. Abth. il. p. 934 (1816) ; G. Cuvier, Begne Anim. i. p. 124 (1817); E. Geoffroy, Descr. de rEgypte, Hist. Xaf. n., Manim. p. 134, pi. iii. fit;-. 2 (1818) ; Desmclrcst, Nouv. Diet. d'Hist. Nat. xxix. p. 513 (1819 : Cairo) ; id., Encycl. Meth., Maimn. i. p. Ill, no. 144 (1820); Gray, Londmi Medical Eqxmtory, xr. p. 299 (1821); Lichfemtein, Verz. Douhl. Mus. Berlin, p. 3t (1823: Egypt); Lesstm, Man. Mamm. p. 112, no. 292 (1827: EgTpt) : Gray, in Griffith's Anim. Kingd. v. p. 57, no. 161 (1827 : Egypt) ; De-'^, Diet. 8ci. Xat. xlvi. p. 367 (1827 : Egypt, " Senegal ") ; J. B. Fischer, Syn. Mamm. pp. 85, 549 (1829 : Egypt, " W. coa.
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30 KOrSETltrs J.GTPTIACTS. c^Bot. ii. p. 503 (1838 : N. & E. Africa) ; BJainviUe, OsL Mamm. Atlas, Cheiropt. pi. xiii. tig. 2 (teeth) (1840) ; Gervais, Hist. Nat. Mamm. i. p. 191, fig. (teeth) (1854: Egypt, Nubia); Giehel, Stint/, p. 999 (1855); id., Odontof/r. p. 9; pi. iv. fig. 3 (teeth) (1855) ; Unger S)Kofscki/, Die Insel Cyiiern, p. 570 (1865) ; Gasco, Yiayqio in Erjitto, pt. ii. p. 95 ( 1 876) ; Klunzinqer, Upper Egypt, p. 148 "(1878). Xantharpyia fegy])tiaca. Gray, List Mamm. Ii. M. p. 37 (1843 : Egypt) ; id., List Osteol. Specim. Ii. 31. p. 10 (1847) ; Kolenati, SB. Akad. Wien, xxix. p. 344* fig. 30 (palate-ridges) (1858j ; Gerrard, Cat. Bones Mamm. B. M. p. 57 (1802: Egypt) ; Tristram, P. Z. S. 1866, p. 93 (Palestine) ; Fitzimjer, SB. Akad. Wien,\\\\ Abth. i. p. 544 (1866: Egypt) ; id., op. cii. Ix. Abth. i. p. 463 (1869 : Egypt, Syria) ; Maischie, Me
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Cynoptenis (Cynomcteris) collaris, Trouessart, Rw. ^Mnq. Zool. (3) vi. p. 206 (pt.) (1878: Gaboon). Xantharpvia collaris, Mntschie. Megachiroptera, p. 66 ^pt.) (1899). Kousettus collaris, Trouessurt, Cat. Mamm., Suppl. p. 60 (pt.) (1904). Cynonyctei-is sp., Bocage, Jom. Set. Math. Lisbon, (2) ii. no. 7, p. 176, te.\t-tig. 3 (palate-ridges) (1892; Pungo Audongo). Diagnosis. — Similar to 21. leavhi, but with larger skull, broader rostrum, broader frontal region, and heavier teeth ; palate-ridges normally 4-f-4-f-l. Forearm 88-99 mm. Skull (fig. 2, on p. 17). — Similar to that of li. leachi, but larger (see measurements, p. 34), and with markedly deeper and broader rostrum. Premaxillaries often co-ossified. Frontal region between orbits comparatively very broad, the width of the interorbital constriction in fully adult specimens distinctly larger than the width of the postorbital constriction. Owing to a stronger development of the temporal muscle (heavier cheek-teeth), the temporal crests unite into a sagittal crest at a short distance behind the postorbital processes, a sagittal crest being always developed in mature age and often present even in individuals with almost unworn teeth. For the same reason the coronoid process of the mandible is higher, the angular portion stronger and more projecting. Teeth (fig. 2, on p. 17). — As in IL leachi, but averaging markedly larger. Palate-n'Jr/es. — 4-(-4-t-l, i.e. essentially as in Ii. leachi, but with an additional middle (divided) ridge behind the molars; the formula given here is taken from a few alcoholic specimens only ; some slight individual variation may be found. External characters. — General size averaging a trifle larger than that of Ii. leachi, head proportionally much larger, muzzle deeper and broader, ears larger, but precisely of the same shape as in a. leachi; wing-structure as in H. leachi; upperside of tibia and foot, to the claws, more densely haired. Colour rather similar to that of It. leachi, but of a somewhat lighter shade. Back and rump hair-brown, with a slight tinge of brownish slate ; crown of head and occiput darker than back ; nape of neck light hair-brown, or drab. Underside grizzled smoky grey ; foreneck and flanks in adult specimens often more or less suSused with wood-brown. In dried specimens exposed to light the slaty tinge of the upperside very soon disappears, the colour fading into brown or yellowish brown. Measurements. On p. 34. lian(je. From Loanda and Gaboon to Egypt, Erythrea, Syria, Palestine, and Cyprus. Tjipe in the Paris Museum. Fterojnis agyptiacus, Geoff. ; 1810. — Type locality : " la basse Egypte " (" le plafond d'une dcs chambres de la grande Pyramide ''). Based by Geoffrey on " plusieurs individus '* ; of these one onljajipears now to be in the Paris Museum, an adult male, mounted, in bad condition, much faded, skull in situ ; labelled Egypt (lieg. no. A. 69). The species was figured eight years later in 'Description de I'Egypte ' {I. s. c).

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32 IIOIISF.TI'US .IJGYPTIACaS. Pleropus geoffroi/i, Temm. ; 1825. — Name proposed in lieu of cegypfiacas, " vu que I'espece se trouve au Senegal, et probablemeufc sur toute la cote septcntrionale d'Afrique," Eleutherura uiiicolor, Gray ; 1870.— Type locality : Gaboon ; type iu collection. — Gray did not give any tangible character by which to discriminate E. unicohr from E. rpr/i/ptiacus. The type is a faded specimen, indistinguishable in cranial and dental characters from examples from Egypt ; a specimen in the British Museum from Loanda (Pungo Andongo) is further evidence of the occurrence of R. cegypiiacus in W. Africa. ReniarJiS. — R. mjypt'mcas is a large-skulled northern representative of the R. leaclii type. It has not rarely been confused with the allied R. leachl and R. arahkus, more frequentlj' with the very different R. lanosus. Its distribution in Africa between Angola and Egypt, and the exact limit:^ of its area in Syria and Palestine (as compared with that of R. arahicjis), remain to be determined. a.

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ROUSETTUS AKABICUS. '.Hi 3. Rousettus arabicus, And. ^dc WInt. Cyiionycteris iimploxicaudata {tii'c Genff.), Dobson, Oct. Chir. Ttid. Mils. p. i' (pt.) (1874: Kisliin I.): Blanforrl, E. Persui, ii. p. Kl (1S70: Kislmi I.); Itoh.vm, Mon. Asiat. Chir. p. 190 (])t.) (187(3: ki-ihiii I.); J. Andfrsim, Cat. Mamm. Ind. Mus. i. p. 104 (pt.) (1881: Kl.sluiil.): Murmii, Vi-rt. Z. :i (1S84: Miikklre Hill.-, Siud); Trouessait, Cut. Mamm. i. p. 81 (pt,)(1897: Kishiu I.). Cyiio])terii9 (Cvnoiiycteris) amplexicaiid.itii.'',7Voz^e.s.s«;-/, lirv. Sf Maq. ^Zool. (3) vi/p. -iC) i (pt.) (1878: Per.-sian Gulf). Xanthuipyia auiplexicaiiilata, lilanford, Fnuna Brit. Lid., Mamm. pt. ii. p. 2U2 (pt.; (1891; Ki.shiii; Karachi); Thomas, P. Z. S. 1894, p. 44i) (-MiHcat) ; Matschic, Me/ackiroptera, p. 67 (pt.) (1899). RousHttii.s amiilexicaudatii.'^, Thomas, P. Z. .S'. 1900, p. 98 (Aden) ; Troue.smrt, Cut. Mamm., Viippl. p. 00 (pt.) (1904 : Aden, ^[ll8cat. Iv.i.slini). XaiUiiarpvia aig-\ptiaca [nee Geoff'.), Yerlurij SjThomas, P. Z. 8. ]S9.'5, p. .•')4o (Aden). Xanthai'])via collavis {nee III.), Matschie, Meqaehiroptera, p. 66 (pt.) (1899: Aden), lloiiseitiis arabicus, Anderson ^~ dc Winton, Zoo}.. Ei/ypt, Mamm. pp. 8(i, 88, 89-90 (1902: Aden); K. Andersen j Ann. i^Mai/. N. H. (7) xix. p. 507 (1907). Diagnosis. — Similar to It. IracM, but with shorter and lower ro^itriuu, narrower ear-tips, shorter tibia and foot. Forearm S796 mm. Details. — The skull of R. arabicus averaaes in ever}' respect smaller than in R. leaclii: total length in fully adult individuals 138 741 "8 mm., against 40'o-43'S in R. leachi; rostrum markedly shorter and slenderer : length from front of orbit to tip of nasals ];j-18'6 mm., against 13'8-15'2 in R. leachi. Teeth as in R. leachi, but smaller, and molars narrower. Palateridges sometimes 4-f-3-f 1, as in R. leachi, but very ofton there is, behind the 6th ridge, a more or less distinct additional ridiie, the extremities of which are either in direct connection with or terminating very closely behind thoseof the 6th ridge, giving the formula 4 -|4 -f 1, as in R. cpi/i/ptiaciis. Inner margin of ear-conch neaidy straight, tip of ear much narrower than in R. leachi • tibia and foot noticcabl}' shorter. Distribution and colour of fur as in R. ii'i/i/ptiacus. From R. o'ljijittiacas this species is readily distinguished bj" its much smaller skull, shorter and slenderer muzzle, much narrower ear-tips, shorter wdngs, and shorter tibia and foot. Measurements. On p. 3-1. Range. From Arabia (Aden, Muscat) to Sind (Karachi). Ti/pe in collection. Remarks. — R. arahicus is more closely related to the S. African R. leachi than to its geographical neighbour R. ar/if/diacus. The exact limits of its area (to the west, north, and north-east) arc still unknown ; it is not unlikely tliat it extends into Syria and ralestine.

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S4 ROUSE I IDS AKAHICrS. «-f. 2(5a(l.,2 2a(l., lahej, Aden. Col. J. W. Yerbiiry 95.6.1 o-C. 1 inill. al. [0. & P.]. /.; skull. Lahej, Aden ; Col. J. W. Yerbiirv 21 Mar. 1895. [C. & P.]. (T>/pe of ^pp^!ies.) ^-/i. c? ad., 5 imm. Lahej, Aden; Col. J." W. Yerbui-y 95.G.1.48-49. .^ks.; skulls. 21 Mar. l.'-9.=i. [C. & P.]. J-;. c?ad., c? imm., Lahej, Aden ; W. Dodson [C.]. 2 ad., 22 Aug. 1899. jimiu.sks. ; skulls. m-7?. c? ad., gimni. Lahej. Aden. Percival & Dodson al. [C.]. 0. 9 iirun. al. Muscat, Oman. Dr. A. S. G. Jayakar [C. & P.l. p-n. 2 ad., 9 inini. Muscat, Oman. Dr. A. S. G. Jayakar al. [C. & P ]. r. 9 ad. al. Karachi, Sind. Karachi Museum [E.]. Measurements of llonsettus leachi, 11. fpgyptiacus, and E,. arabiciis. 7?. hachi. E. (pgyptiacus. 11. arahkin MiN. Ma.\. Forearm Pollex, c. u 2nd digit, metacarpal ,, 1st phahiiix ,, 2rd-3rd phalanx, c. u ; 3rd digit, Dielacarpal ,, 1st phalanx I ,. 2nd phalanx '• I 4th digit, metacarpal „ 1st phalanx ,. 2nd phalanx 5th digit, metacarpal ,, 1st phalanx t „ 2nd phalanx Ear, length irom notch s ,, grpate:.t widtl), flattened Front of eje to t'p of muzzle Tail j Lower leg Foot, c. u Skull, total length to front of premax — ,, width of brain-rase at z3goniata...i „ zygomatic widtli ., posto]-bilal constriction „ interorbital constriction „ width across m-, e.xternally „ width across c, externally ,, palation to incisive foramina „ front of orbit to tip of nasals i Mandible, length , Upfer teeth, c— m^ ! Lower teeth, c — m^ mm. 89 31 36-2 8-8 9 54-5 38-2 50-6 54-2 29 32-2 54-5 27 25 20 15 17 15 40 24-5 40-5 167 24 7 7-7 7-S 12 5 8-2 19 7 13 8 31-8 15-8 17-2 nmi. Vi9 35-2 43-7 10-5 10 62 42-2 60 lil 3.S-8 38-7 CO 30'8 28-5 22-5 IC 19 2 18 8 41-8 27 43-8 18 26 9-2 87 14-2 92 21-2 15 2 34-8 17-7 19 Wi.\-. Max. mm. 88 32 40 8-7 9-5 57-2 39 53 54 30 33 54 28 25 22 17 17-2 13 41 25 43-6 17-7 26 7-2 S-2 12-8 8-7 208 15-2 33 16-7 18 mm. 99 37-5 44 11'7 123 63 44 61-5 61 34-5 39-8 59-5 30-8 318 24 19-7 19-2 15'5 45-5 27-8 46-7 18-8 29-2 8-7 9.8 14'5 9-8 23-7 16-7 37 2 18 8 20-8 MiN. M.\x. mm. 87 30 37 8-2 9 r3-2 37 50 5 50-8 28 31-8 50-2 26-2 24-2 20 15 15-2 9 37 22-7 o87 16-3 23-2 7(; 7-7 12 7-7 18-2 13 30-5 15-2 16-6 mm. 9(> 33 42-7 9-8 105 (>0 39-8 50 5 58-7 31 36-7 68-7 30 28-8 21-2 16-2 17-5 17 39-6 2.r5 41-8 17 25-8 8-2 7-8 13 8-7 20 13-6 33 16-7 178

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KOUSETTUS LISCIIKNAUMI. 35 4. Rousettiis leschenaulti, Desm. Ci/iionifcteris amfiltxicawlnta (pf'.)> Dob.son, Cat. Chir. B. M. p. 12. Pteropus lesclieiiiiLilti, T)esmarest, Eiiq/cl. Meth.. Mti/iiin. i. p. ]1(), 110. 142 (IHiJO: Pondichery) ; ifl, Did. Sci. Nat. xlvi. ]>. 3()o (1S27: Pondichery); Lesson, Man. M/inwi..\). 110, no. 28;'. (1827: Pondichery); (ri-(u/, in Grfffif/i'.s Aiiivi. Kini/d. v. p. 5/st. Verz. Savg. i. p. 1,80 (1844: Pondicliery) ; Wai/iu'i; Sc/ireOer's Sii'icj., Siifpl. v. p. 604 (pt.) (185.355: Pondichery, Calcutta); Giebel, Simp. p. 999 (pt.) (1855: Pondiclicry); Bbjtlt, Vat. Mamm. Mus. As. Soc. p. 21, no. 54 ())t.) (]8(!8: Coromandel ; Jcrdon, Mavnn. Itid. ^. 19 (1867: Madras, Trichinopoli) ; Sterndale, Mamm. India, p. 40 (pt.) (1884: Calcutta, Madras, Pondichery, TrichinopoJi). Xantharpvia leschenaulti, Fitzinyer, 8B. A/cad. ll'icn, l.\. Abth. i. Heft 8,"p. 472 (pt.) (18()9: India). Cynonycteris (Pteropns) lescheuiiulti, Mavchi, Atti Soc. Ital. Sci. Nat. XV. p. 517 (1872-7."} : structure of hairs). Cynouycteri.s leschenaulti, I'eters, MB. Akad. Berlin, 187.3, p.. 485 (type of species re examined). Itousettus lesclienaulti, K. Andersen, Ann. ^Maq. N. II. (7) xix. p 507 (1907). Pteropus ainplexicaudatus {neo Geoff.), Tennninck, Man. Mamm. i. p. 200 (pt.) (1825 : Siani) ; Desmarost, Diet. Sri. Nat. xlvi. p. .367 tpt.) (1827: Siam) ; J. B. Fischer, Si/n. Mamm. pp. 86,549 (pt.) (1829: Simn); Giebel, Siiuij. p.lO0O(pt.) (1855: India). Cynonycteris amplexicaudata, Peters, MB. Akad. Berlin, 1867, p. 865 (pt.) (Penj^'al, 8iani) ; Swinhoe, P. Z. S. 1870, p. (ilO (Anioy) ; Peters, P. Z. S. 1871, p. 51:'. (Burma) ; iJohson, Proc. A.S.B. 1872, p. 154 (Burma); Macalister, Phil. Trans. 1872, p. 125 & .«eq., pi. xiv. fifi-. 4 (myology) ; l)obso)i, J. A. S. B. xlii. ]it. ii. p. 200, footnote ( 187.3) ; id., op. cit. xlii. pt. ii. p. 202 (pt.), pi. xiv. iig. 8 (ear) (187.3) : id., Cat. Chir. Ind. Mas. p. 2 (pt.) (1874: Coromandel coast, Siugbhoom, Peo-u): id.. Mm. Asia': C'A(V. oPP-^•', 1'^^ (j't.) (1876: Indian Pen.) ; ? Leche, Lnnd.<: Univ Arsshr. xiv. p. 17, pi. ii. fig-. 9 (1878) (milk-teeth) ; ./. Ander-'ion, Cat. .Mamm. Ind. Mns. i. p. 103 (pt.) (1881 : Coromandel coast, Iiidia) ; ScxtUij. J. A. S. B. Ivi. pt. ii. no. ,3, y. 237 ( 1SS7 : Nepal) ; Jentink, Cat. Oste1 (pt.), text-iifr. 76 (ear) (1891 : India, Himnlava, JSurnui); Thomas, Ann. .Sins. Civ. Genora, (2) x. p. 921 (lS92: Moulmcin) ; Mn/svhie, Mei/arhirojitcra, p. 67 (pt.) (1809) ; S. .V Flower, P. Z. S. 1900, p. 340 (Lao.^ Mts.). p2

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86 EOUSETTUS LESCnKNAULTI. L\viii'])toi'us (Cyiii nycteris) ainplcxicandatiis, Troiwssarf, Iter. Sf ' Maij. Zcol. (?>) vi.'p. I'OG (pt.) (1878: Inciia, TsVptil, Jiiiniia). CynonTctt'ris (Pteropus) ani)>lexicaudatus, Thcohnld, in Masons JJiirmii, i. ]). 'l2o (1882 : Eenf>-al, S. India, Biiniia). lit.iisetUis 1 nipU'.\ic,uidiitu8, Trouessarf, Cat. Mamm., Sitppl. p. 00 (pt.) (1904: Himalaya). PteiM pus [,sp.], IMr/.soii, J. A. S. R. no. 7, p. 340 {\^?,± : Nepal). I'teropiis pvri\orii^, Ilodqsoii, J. A. S. B. iv. no. 48, p. 700 (1835: Nepal) ; ' id., V. Z. S. imn, p. 4G (Nepal) ; uh, Icon. ined. (B.M. copy) pi. xli. fifr. 1 (col. fit;'.), pi. xiii. fig. 2 (col. fig.). Pteropus pii-ivarus (sic), Hodyaun, J. A. S. B. x. p. 908 (1841 : Nepal). Oynopteiiis afiinis, Gray, List Mamm. B. M. p. 39 (1843 : Iliuialaya). Cynopterus ninrpnatus {nee Geoff.), Gray, List Mamm. B. M. p. 38, speciuieriR A, / (1843 : Nepal, the cotvpe.s ri Pt. pyriwrus, Hodgs.) ; Biyth, J. A. S. B. xiii no. l.OO, y. 479 (1844 : 'Ne} al) ; Gray, Cat. Hodgson Coll. B. j\J . ]). 3 (184(j : Nepal) ; Ilidton, P. Z. S. 1872, p.' (J93 (N.W. Hiiiialaya). Eleutlierura niar^inata, Gray, Cat. Blonh. S)-c. p. 118 (1870: " Nepal," really Nasirabad). ': Eleiitlierura fuli<:inosa, Gray, Cat. MovJc. i^c. p. 118 (1870: Laos Mts., Siani). .: Eleutlierura fusea. Gray, Cat. Monk. S)-c. p. 119 (1870 : " India P"). Cynonycteiis infuscata, Peters, MB. Akad. Berlin, 1873, p. 487 "(Calcutta). Diagnosis. — Allied to ii. arahicvs, but f-maller, with tlie imizzle shorter and slenderer, tlie tip of the ears not attenuated, the jiollex markedly shorter, wings shfirter, cs]jecial]y the first and second phalanx of the third digit, and the loot smaller. Porcaim 80-587"5 mm. ^ill•uU and teeth. — Similar to those of Ji. arahicvs, but skull averaging a little smaller, rostium slendeier; preniaxillaries ratlier more strongly projecting forward, palalion border more t'harjily angulate. Teeth on the whole ratlier smaller, raolais somewhat narrower ; p' not deciduous, unless perhaps in very aged individuals (cf. li. femini(tlns) ; m^ quite or almost equalling m, in length ; m, elliptical in outline (cf. 2i. umph x.cuudaiits). Fulate-ridges. — 4 + 3 -+1 ; number and arrangement as in U. leucJii. External cliaraeiers. — Muzzle averaging shorter and slenderer than in Ii. arahievs. IShaj e of ears as in Ii. Jeae^i ' outer miuh more convex than inner margin, tip broad, not attenuated as in li. arahinis; aiititragal lohe small, rounded. General size neaily always smaller Ihan in Ii. aralicvs : forearm 80-O-87-5 uim., against 87-90. Eirst digit absolutely shorter: length with claw '26-o-2i)-tj mm., against ;-50-;H;i in arubicus ; also jiroportioniilly the difference in this respect is well marked : index of iiist digit in Ii. lesvlunardti '6'65, in Ii. arahievs li'il. Second digit shorter, the tip of its claw in A', hsc/ienaulti as a rule falling a little short of, in Ii. arahiens as a rule being on a level with or reaching beyond, the tip of the third metacarpal. Metacaipals and

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EOUSCTTCS tESCHKNAULTI. 37 ]ih;iLuip;es of third, fourtli, and fifth dijjifs (except the second phiihiiix of the tilth digit) shorter thiui in R. aralncH>i, the dill'ererice bein;;; speciailjnotieoat)le in tlie absolute length of the first p'lahuiK of the third 'digit (33-8-;3(3'8 mm. in 11. hfchenanlH, 37-39-8 in li. (f rahicufi) and the second pliahinx of the third dvjit (41— Kr2 mm. in It. Iesch>'nn>dti., oO-[)-(V) in li. (imhicns: index of this phalanx in It. I'scheiiaidti 521, in It. arabicns 601). Length of tibia as in ii'. nrahicux, but foot averaging smaller : me isured witli claws 2U-23-5 mm. in li. leschenaulti. 22-7-25-5 in It. arahicus. Quality and distribution of fur essenrially as in R. arahinis, but tibia more thinly haired. Colour of a darker and browner shade, especially on the crown : — Back and rump dark and dull brown (darker than Front's brown) ; crown and occiput lirowiiish bistre; nape of neck varying from light drab to almost wood-brown; underside between drab and isabella colour. A considerably brightercoloured phase occurs : mars-brown on back and rump, Prout's brown on crown, wood-brown on the whole of the underside. Measurements. On p. 48. Itaiir/e. Himalayas (Nepal), extending southward over the Indian Peninsula, eastward through Bengal, Burma, Siam (Laos Mts.) to S. China (Araoy). Cutijpes in the Paris iluseum. rte'ropusleschennulti, Desm.; 1820.— Type locality : " les environs do Pondichery." Cotyi)cs, two adult males, mounted, much faded, hibolled "Pondichery", Leschenault "' ; Eeg. nos. A. 82 and A. 83; skull of no. 82 extracted, of no. 83 in situ. I'laeed by Desmarest in the section " Ronssettes sans queue," an error corrected by Is. (leoftVoy in 1828, /. s. c, on examination of Ihe type : (" sa queue, trcs-visible, u'est qu'a ])eine cngageu dans la membrane interfemorale, et a environ six lignes de long "). Pti'rojmx pi/rivorux, Hodgs. ; 1835.— Type locality: the central region of Nepal ; cotypes in collection, figured in Hodgson's unpulilislied drawings. Cifiiopierns nffini^, Gray ; 18-13. — No description. Two specimens were registered by Gray under this name, both from the "Himalaya," viz. no. 130rt [ =], which is the type, still in the collection of the British ifuseum, and 130 ft [ =, "younger," not found in the collection. Indistinguishable from R. leschrnaidti. FJcutherurci ftdiip'vosa, Gray; 1870. — Type locality : Laos Mts., Siam (Mouhot Coll.); tyjie incoUection.— Brief description of the colour of the fur. I am unable to discriminate this specimen from It. lesdienaxlti. Eh'uJherura fvsca, Gray; 1870. — Type locality uncertain ("India?"'; purchased from Par/udaki) : tyi>e in collection. — Separated by Gray on account of its "much brighter and redder" colour. Is the brighter-coloured phase of R. kxchemmlti. Cinwnycteris infmeutd. Pet.; Ib73.— Type locality: " angeblieh aus Calcutta" (a dealer's specimen); type in the Berlin Museum (no. 301 ). " Schr iihnlich der 0. lesrh^naKhii. in alien Verhtiltnissen

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38 "ROURETTUS SKMINITDFS. kleiiior, flnnkelbrnnn von Farbe, mit schwarzon Krallcn uud don erstoii falscluni liackzalin g-riisser " : detailed measurements given : forearm 68, tliiid metacarpal 42, tibia 29 mm. The type, I am informed by Trof. Matschie, is a young (not full-grown) individual ('die Epipbysen an den Fingergelenken sind noch nicht mit den Phalangen verwachsen," Matschie, in litt.) ; hence its small size. Remarks. — li. leschennaUi is at once discriminated from the foregoing species by the shortness of the second phalanx of the thi'd digit, a character which it shares with all the Eastern species of the genus. Piu-cbased (J. Tamer). .38 3.13.37. {Ti/pe i)t' Viniojjierus affiim. Gi-aj-.) B. if. H.idgsoii. Esq. [P.]. Not reg. (Coiypes of Pteriiqiuf, pyrivorus, Hodgs.) B.fl. Hodgson, Esq. [P.].<5. Cajit. \V. J. E. Boys 4.^.2.1.14. LO.]. Capt. W. J. E. Boys r |C.]. [49.8.16.y. (e &/ labelled Elcafhcrnra marqiiuifa. Gray.) Lieut. E. Y. VVatsun [P\J. Si;im. Moubot Coll. 62.8.18.r). {Type of F.lcnfhcrvrn fid'iqhui»a . (Trnv ) 'Pufcliasfd (Pai-ziidaki)! 4!>.8.23.1(). {Type of Eleufhrrura j'uFca, Gray.) Lidth de Jeiide Coll. (.7'.4. 12 32f5. 5. Roiisettus seminiidus, Gray. Oynoni/cferis am])le.vkandnta (pt.), Dobson, Gat. Chir. B. M. p. 72. Pteropus leschenaulti (nee Desni.), Blyili, J. A. S. B. xx. no. 211', p. l.')5 (1851 : Ceylon) ; id, op. cit. x.xi. no. 228, p. 34o (1852: Ceylon); Kelaart, Prudr. Faun. Zeylan. p. 27 (1852: Ce.ylou); Wayner, Schrebrr's Siiuy., Si/ppl. v. p. 604 (pt.) (1853-55 : Ceylon) ; Giehel, Siiuy. p. 999 (pt.) (1855: Ceylon); Blytli, Cat. Muvim. Mun. A.'t. 8oc. p. 21, no. 54 (}it.) (1863: Cevlouj ; Sterndale, Mamm. ImUa, p. 40 (pt.) (1884: Ceylon). Xauthaipyia leschenaulti, Fitzinyer, SB. Alcad. Wien, xlii. Heft 25, p. 389 "(1861: Ceylon) ; id ^' op. cit. Ix. Abtb. i. Heft 8, p. 472 (pt.) (1869: Ceylon). Cynonycteris (Pleiopus) leschenaulti, Zcklor, Beise ' Koiara,^ 'Siiiiy. p. 12 (1869 : Ceylon). C'ynoiiycteris amplexicaudata (pt., nee Geoff", j, Pefer.f, MB. Akad. Berlin, 1867, p. 865 (Cevlon); Ihilmjit, J. A. S. B. xlii. pt. ii. p. 202 (1873) ; id., Cat. i'hir.Ind. Mus. p. 2 (pt.) (1874 : Ceylon) ; id., Mvn. Asint. Chir. pp. 29, 190 (187f>: Cf-ylon) ; J. Anderson, Cut. Ahniim. Ind. Mus. i. p. 103 (1881: Cevlon); Truuessart. Cat. Mcnmn. i. p. 84 (1897). Cynopterus (Cyuonvcteris) amplexicaudatus, Trouessdrf, Bee. ^• May. Zoul. (3) vi.'p. 20(j (pt.) (1878: Ceylon). a.

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RaUSETTUS SEMTXTTDUS. U9 Xaiith;ivpyia aniplfxicandata (pt.), Blanford, Fami. Brit. Lid., Mamm. pt. ii. p. 201 (18;)1 : Cevlouj ; Matschie, Megachiroplrra, p. (37 (pt.) (1899). Xantliarpyia tiemiiuida, Gray, Cat. Monk. Sfc. p. 115 (1870: Ceylon). Rousettus seniinudus, K. Andersen, A7in. ^May. N. H. (7) xix. p. 538 (1907,1. Diagnosis. — Similar to R. leschenaulfi, but p' deciduous, iiajie and slioulders scmi-uakt'd, general colour of fur lij^hter. Forearm 79-8o-5 mm. Detaih. — Size and shape of skull as in 7^. JescJienaulii. p' deciduous* ; m, quite or almost equalling m, iu length ; m^ elliptical in outline. Palate-ridges as in Ji. hschenaulti. Pur markedly shorter and more closely adpressed than in 7^. hucheiumltl ; nape and shoulders covered with such sjiarse and short hairs as to appear semi-naked ; hairs on membranes and upperside of forearm and tibia shorter and sparser : in some individuals these parts appear to be almost naked. Colour lighter than ill 7i. Icsclienaultl : upperside intermediate between marsbrown and wood-brown, or very nearly wood-brown ; head darker; underside wood-brown. M.-asiiremenis. On p. 48. Range. Ceylon. Type in collection. Xaiitharpijia .'leminuda. Gray ; 1S70. — Typo locality : Ceylon.— The name Pteropus semimidits is commonly assigned to Kelaart, but Kelaart seems never to have published any description of the species ; in the paper usually referred to by authors, viz. Blyth's account in J. A. S. li. xxi. p. 3-15 (1852) on a collection of mammals sent by Kelaart to the Asiatic Society of Bengal, the name ajipears only as a synonym, without comment, of Pt. lescheiianlti, and the same is the case in Kelaart's ' Prodromus Fauna Zeylanicae ' (1852). It remained a nomen midiirn, until iu 1870 (Z. s. c.) Gray published a brief description (quality of fur, colour, length of forearm) of " XaatJuirpyia seminada" and the British iluseum specimen on which he based this descriiition is, therefore, the type of the species. o. (^ ; skull. Ceylou (l/nvaites). Mr. Cuming's Coll. Not reg. (Ti/pe of fpocies.) 6. cJ ;skull. Ceylon. Dr. Kelaart [C. & P.J. .'i2.5.9.10. (Authentic specimen of"' Pteropus semiinuliis. Ke\"^ c. J ad. sli.; skull. Punduloya. Cevlou, E. E. Green, Esq. [P.]. 9,j. 7.27.1. 4:joo' ; 30 Apr. is;i.^. * Details from three skulls : — one im'nature, p^ present on both sides; one ad., tei'th very slightly worn, p' present on one, nbsent on the other side: one ad., tfetli unworn, p' ili>i»nt on both i-'Hoa

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40 RorSETTDS AMPLEXICAUDATUS. 0. Rousettus amplexicaudatus, E. Geoff. Cijnonijcteris atiij_)lv.iicaadata (pt.)i Dobsou, Cat. Chir. \^. M. J). 12. rteropus amplexicaudatus, i'. Geoffroi/, Ann. Mus. d'Hist. Nat. XV. p. yC), pi. iv. (whole tif,r) (1810: Tiinoi-j ; Okeii, I.elirb. XatunjeKch iii. Abth. ii. p. 934 (1816 : Timor) ; G. Vucier, Ithjue Anim. i. p. 1:24 (1617) ; Desinarest, Now. Did. ci' Hi^t. Nut. xxix. p. r)13 (1<"?19 : Timor) ; id., JEmycl. Meth., iVJatinit. i. p. Ill, no. 145 (1820: Timor); Temmiiwk, Moii. Maiiiin. i. p. 200 (pt.), pi. .\iii. (whole iig.), pi. xv. lig. 16 (bead) (182".: i'imor, WumaU-a) ; Lei^son, Muii. J^iamm. p. 112, do. 203 (1827 : Timor) ; Gir/t/, in Griffith's Anim. Kiti(;d. v. p. 57, uo. 162 ( 1827 : Timor) ; Desmaresf, Diet. Sci. Nut. xlvi. p. 367 (pt.) (1827: Timor, Sumatra); Is. Geo/f-oy, Diet. Class, dllist. Nut. xiv. p. 703 (pt.) (1828: Timor, iSumatra) ; J. B. Ftsclier, Syn. Mnmni. pp. 86, 549 (1829: Timoi', Sumatra); li. Geofroy, Mist. ^ at. Mamvi. lei;. 13, p. 25 (1829); Wayler, Syst. d. Aniphihiiii,-^. 9 (1830) ; Lesson, Hist. N(d. Manwi. (Co7npl. Biiffhn) v. p. 55 (i:t.) (1836 : Timor, Sumatra) ; Temvdnck, Man. Mamnt. ii. p. 90, pi. XXX vi. H>^s. 18, 19 (sivuU) (1837) ; S. Mi'dler, in 7'evmiinrk's Nat. Gesch. Nederl. overz. Lez., Zoof/d. ]ip. 20, 58, 59 (pt.) (1839-44 : Timor, Sumatra) ; Waijncr, Sdirebers Siiui/., SuppL i. p. 359 (pt.) (1840; Timor, Sumatra); Lesson, N. Tail. Ithjue An. p. 14, no. li)3 (pt.) (1842: Timor, Sumatra) ; 8ulanz, iSyst. J'erz. iSduy. i. p. 131 (pt.) (1844 : Timor, Sumatra ) ; U'ayni-r, Schrehers Siiuy., fSuppl. V. p. 604 (pt.) (1853-55: Timor, Sumatra]; Gielxd, Siiuy. p. 1000 (pt.) (1855: Timor, Sumatra) ; tSihleyd, Dierhttnde, i. p. b'-i (1857: Sumatra); Fatsih, J^eu-Guinca, p. 150 (pt.j (1865: Timor). PacliYSoma amplexicaudatum, Wntcrhouse, P. Z. S. 1843, p. 67 (i'iiilippines). Pteropus ((!\niinycteris) amplexicaudatus, Peters, MB. Akad. Berlin, 18()1, p. 707 (Luzon, Samar). Xantbarpvia aniplexicaudata, Gerrard, Ccd Bones Mamm. B. M. p. 58 (1862); Fitzuu/er, SB. Akad. Wien, 1.x. Abih. i. Heft 8, p. 470 (pt.) (1869: Timor, Sumatra); Thovms, Ann. Mus. Civ. (Jenova, (2) .xiv. p. 108 (1894: Eiigano) ; id., Nov. Zool. iv. p. 263 (1897: Savu I.) ; id., Trans. Zool. Soc. xiv. pt. vi. p. 383 ( 1898 : Luzon) ; Hurtert, Nov. Zuul. v. p. 456 (1898 : Alor I.) ; Matschie, Meyaehiroptera, p. 67 (pt.) (18i.'9). Cynt)uycteris ample.xiraudata, Peters, MB. Akad. Berlin, 1867, "p. 865 (pt.) (Timor, Philippines) ; Dobson, J. A. S. B. xlii. pt. ii. p. 202 (pt.) (1873) ; Peters, MB. Akad. Berlin, 1873, p. 485 (type of species re-examined) ; Dobson, Mon. As. Chir. pp. 29, 190 (pt.) (1876 : rhiiippine;;) ; id., P. Z. ,S. 1878, p. 877 (Cambodja) ; id., Pep. Brit. Assoc. 1880, p. 1 73 (Cambodja) ; Rubin, C. R. Acad. 8ci. xc. p. 1369 (1880: anatomy); id., Ann. Hci. Nat., Zool. (6) xii. art. 2, pp. 4lS: seq., pi. v. lig. 30, vii. bg. 47 (organs ot reproduction) (ls81); J.Anderson, Cat. Mamm. Ltd. Mus. i. p. 103 (pt.) (1881: Pbilippiaes) ; Jeidink, Cat. Osleol. Mamm. p. 263 (pt.) (1887 : Timor, Sumatra) ; id.. Cat. Syst. Mamm. p. 150 (pt.) (1886: Timor) ; D. G. Ediot, Field Coliunb. Mus. Piibl., Zool. i. no. 3, p. 79 (1896: Negios L) ; 'J'ronessart, Cat. Maiinn. i. p. 84 (pt.j (1897); tieabra, J. Sci. Math. Lisbon, (2) v. no. 19, pp. 161, 168, pi. i. lig. 10 (palaty-ridgc.-) (1S'.»8: Timor).

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RorsF.rrns ,\Mri,r,xiCArpATrs. 41 ('viinii\ct('i'is (I'ternpiis) nniplc.xicauilatiis, Mnvchi, Atti Sic. Ital. 'Sii. Ndt. XV. p. 517 (187i'-7.'< : structure nt' Imirs). (.'viiopierus (Cynonvcteris) iuii])l(;xicaudatus, Trouessart, licv. iSMm/. ' Zuol. (;J) vi.'p. :266 (pt.) (1878). Kousettus fiinplexifaudatu.-;, Troiwsxiivt, Cat. Mamm., Siip/il. p. (iO (pt.) (llKU); Miller, Pruc. U.S. Nat. yiiis. xxx. p. 824 (1906: Euyaiio) ; K. Anih-rsvn, Ajvi. ^Ma;/. X. H. (7) xix. p. '"-08 (1907) ; Miller, Fan,, if Gen. Bats. p. o4 (pt.) (1907). Elontlierura int'uniata, Gray, Cat. Monk. ^-c. p. 118 (1870: Flores). Eleutherura philippiuen.-is, Graij, Cat. Monk. SfC. p. 119 (1870: Manila). PuTopus ])lulippineasi.s, Elern, Oit. Sid. Faiiua Filij)inas, i. ]). 6 (1895: Luzon; Samar ; Min^'anao ; Palawan). Itousettus philippinensis, Miller, Fam. ^Gm. liat^, p. 54 * (1907). Cvnoiivcteri.s bocaiiei. Seabra, J. Sci. Math. Lisboa, {'2) v. no. 19, 'pp. it"0, 1G9, pi. i. tifr. 11 (paktu-ridge*) (1>^98: Timor). liouseltus bocagei, Truutssart, Cut. Mumm., Siippl. p. UO (1904). IHa;f)Wsis. — Similar to li. leschenauM, bttt m^ suljcireular in outline, ears narrower. Forearm 77-S7"2 mm. Skull and teeth. — Skull essentially as in Jt. hf'clunaulti, but avera^^ing smaller; rostrum proportionally .slenderer. — Teeth smaller: maxillary row 12-8-14:2 mm., against 14-15-7 in 7i. /esrJwnaulti. m^ subcircnlar in outline, not elliptical as in It. leschenaitlti. In li. amj^) there is a distinct tendency to a reduction of the interspace between the upper canine and p\ i^ being in some individuals rather closely wedged in between those two teeth ; but the character is not iixed, many examples (independent of geographical habitat) having the interspace c-p' as broad, and p' as distinctly separated from either of these teeth, as li. lexelienaulti; li. amplexicaudntns forms in this respect a transition between R. ItsehenavUi and by-ac/njoiis, in which latter p^ is deciduous, c and p'' occasimially in contact. r(date-rid
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42 EOrSETTUS AMPLKXICAUDATUS Range. Carabodja, Piiiliiipiues, Borneo, Sumatra, Engano, Flores, Savu, Alor. Timor. Tjipe in the Paris Museum. Fterojms am/>Je.cicaiulatn.s, Geoff. ; 1810. — Based bjGeoffroy on "plusieursindividus" obtained in Timor during Peron and Lesueur's voyage (Capt. Baudin) ; only one of these appears now tc be in the Paris Museum, a young individual, mounted, much faded, skull extracted, labelled " Timor, Exp. Baudin" ; lieg. no. A. 79. ELeutlierura infumata, Graj' ; 1870. — Type locality : Flores (A. 11. Wallace) ; type in collection. — The name would seem to indicate that Gray separated the Flores specimen on account of its " blackish browu "' colour ; it ditfei's, however, neither in this nor in other respects from a majority of examples oi R. amphxicaudatus. Eleulherura pJiilippinensis, Gray; 1870. — Type locality: Manila (Hugh Cuming); type in collection. — Gray gives a brief description of tbe colour of the specimen, without pointing out his reasons for separating it from Pleropus amplexicaudatus, Geoff. The type, as well as more recently acquired specimens from the Philippines, differ m no respect from R. amplexicaudatus. Ci/7ioni/cteris hocar/ei, Seabra ; ISUS. — Type locality : Dyli, Timor (Fr. Newton); type in the Lisbon Museum. ^ — Separated by Seabra from R. ahip>lexicaudatus on account of the supposed greater zygomatic width of the skull (and a trivial difference in the form and position of the sixth and eighth palate-ridges). Zygomatic width of the type skull {S ) not given ; in the figure {I. s. c. pi. i. fig. 11) it measures scarcely 23 mm. ; in a British Museum specimen ( c? ) from Alor I., north of Timor, 24 ram., in another ( $ ) from the same island 21*2 : similar variations are found in R. amplexkaudatus and allied species from an)' locality. Remarlcs. — R. amplexlcaudaius is readily distinguished from R. lescJienaulti by its much narrower ears and the difi'erent shape of nig. ill. hschenaidti is continental in range, R. amplexicnudatus chiefiy Indo-Malayan (insular) ; their areas probably touch each other somewhere in S.E. Asia ; R. leschcnaidti is represented in the collection from Burma and Siam, R. ample.clcaudatas from Cambodja. — Some authors (Peters, Anderson and de Winton) have laid stress on the greater length of the tail in Indo-Malayan specimens {amplexicaudaius) as compared with examples from tlie Himalayas and the Indian Peninsula (lescJienaulti) ; the tail averages, in fact, decidedly louger in wmplexicaudcitus, but the character, as being subject to a good deal of individual variation, is practically not of much use for a discrimination of the two species. a. Ad. sk. ; skull. Cambodja (iVouTomes Coll. hot Coll.). b. 2 imiii. al.; Philippines. Zool. Soc. Coll. Not reg. skull. c. S ad. sk. ; Manila. Luzon. Hugh Cuming, Esq. .skull. [C.]. (Type oi Elci'.lherura philip2>ineHsu, Gray.)

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ROtrSKTTTJS MTNOK. 4.J d-e. S 'tI" " '«'• E<'iiKuet. N. LuJ. Whiieliead, Esq. OT.o.^.o-C. sks. ; skulls. zm, i^lHJO' ; [C.j. 14 Febr. 18i)4. ./-.9. c? iw^.. (5 P"llBanuii, ISarauak. Dr. Chas. Koso [P,]. 0.7.29.'2-3. al. ; skull of/: //. ^ acl. sk. ; Suinalra. Purchased (J. Tiir3S.3.13..'ib. skull. iier). i. 2a(; skull. Bua-Kua, EnMarquis C4. Doria gano {Dr. K. [P.J. ModiyUaui). j. skull. Flores. Dr. A. R. Wallace (i3.12.26.12. [C.]. {T(ipc of Eleafhrrurti hifumata, Gray.) >!•. ;skul!. Saru I.. W. of A. Everett [C.]. Tiincir ; Aug. 18«G. l-m. d' ad., 9 ad. AJor I., ^\ of A. Everett [C.]. sks. ; skulls. Timor; Mar. 1897. n. luiui. St. Tiuior. Purchased (Frank;. 44.4.4.(5. 7. Rousettus minor, Dohs. Ci/iionycteris minor, Dobson, Cut. Cliir. B. M. p. 73. Pt-n-opn.s aniplexicaudatus (pt., nee Geoff.), Besmaypst, Diet. Sci. Nat. xlvi. p. 367 (1827 : Java) ; Lesson, Hist. Nut. Mumm. {Conipl. Hufon) V. p. o5 (18-JG : Java) ; 8. Miiller, in 'i'eiiiniitick's Nut. Gesch. Nedeii overz. hcz., Zuoyd. pp. 20, 58 (1839-44: Java); Les>
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44 EOUSETTUS nnAcnTOTis. (only one (|nite fiill-sizcfl skull of brachi/otis available for comparison). oiil}' tangible difference in the teeth would seem to he the slightly smaller width of the premolars and molars in li. minor, but the character must be taken with caution, inasmuch as there is some individual variation in this respect in 11. hrachi/vtis. p^ a little smaller in bulk than an inner upper incisor, separated by miuute spaces from the canine and p'^. ni^ subcircular in outline (a little longer than broad). Rvternal characters. — According to Dobson the ears in R. minor are much longer than in li. bracJu/oiis (see his description of li. hrac7i(/otis, Cat. Chir. p. 74). Tliis statement is erroneous ; the oars in the type of i?. yninor (the left ear-conch is well preserved, the riglit much damaged) are of the same length as in dried skins of li. hrachi/otis, or if there is any difl'erence, they are perhaps a trifle shorter ; the width of the ears cannot be estimated with certainty, but would seem to be slightly smaller than in hrachifotis. In the relative length of the muzzle (wrongly stated by Dobson to be shorter than in brachi/oiis), the distribution of the fur, and the measurements of the forearm, metacarpals, phalanges, and tibia, there is no difference from li. braclnjotis. Measurements. On i). 4S. liange. Java. Tjiiie in the Calcutta iluseura. liemarks. — Judging from the single specimen examined, li. minor appears to be more closely allied to It. brachi/oiis than to li. amplcvicaiulattis. From the latter species it is readily distinguislied by its smaller size and relatively smaller ears. Its differences from M. brachyotis remain to be determined *. 8. Rousettus brachyotis, Bobs. Cjinonycteris brcicJiyotis, Dobson, Cat, Chir. B. M. p. 74. Pt'.^ropus amplexicaudatus (pt., nee Geoff.), Desmaresf, Diet. Set. Xat. xlvi. p. yi)7 (18:^7: Ambnina) ; Is. Gfofroif, Diet. Class. d'Hisf. Nat. xiv. p. lO'.'j {\S'26 : Amboina) ; J. B. Fiseher, Sj/n. Manim. p. SO (1829: Amboina); Lesson, Hist. Nat. Mamm. (Compl. liiiffiit)) v. p. oo (183(i: Amboina);
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BOfSETTlS BKACnyOTIS. 45 Cviion7ctfvi.« ani])loxicaii(la(ii {i\t.), I'cfcrs, Mil. Al-ad. />,r!,'n.]fi(u, ]i. si).") fAiuboina); Jentink, Cut. Si/st. Maiiun. p. 1")1 (]8r!8: Aiuboina). Xaiilliiiri)\ia amplexicaiidata (])t.), T'ltziiK/er, SB. Ahod. ]]'ien, Ix. Abih. i". lleCr <, p. 470 (1860 : Amb.iiiia). CvnMpii,'ius (1 viioiivcieri.s) airplexicaudatus, Troiiessdrt, Iin\ S; ' M) vi. p. l'OG(pt.) (1878: Aiuboina). Cvnonvctm-is brachvcti^, I)ofj.^f^t<, (ip. 47o, 483,484 (.Soh.iiiun Is.) ; Jcntiiil-, Cat. S'/ft Mcotim. p. h")! (1888: Duke of York I.); 'J ruuessart, tat. Mainiu. i. p. 84 (pt.) (1^97). C}noptorii.'3 (Cviiouvcteri.s) braclivotis, Truues-sart, liev. 4'3/"'/. Zool. (3) vi. p. 201') (1878). * Xanthai'pyia brachvotis, Mat.'uhie, Mcf/achiropterti, p.fiS (pt.) (1899 : New Guinea, New Ireland, .Solomon Is.). Ilousettus braclivotis, Trouessart, Cat. Mamii)., StippL p. 60 (]it.) (1904) ; K. Andersen, Ann. if Mar/. X. H. (7) xix. p. .3UD (1907) ; Milk-r, Fam. ^Gen. Bid>:, p. o4 (1C07). Diiicjiio^is. — ISiiuilar to JL ampli'dicaxdatus, but smaller, with shorter and narrower ears ; p' deciduous ; toutli-rows shorter. Forearm 70-75 uim. kt in collection. Remarks. — This, the most eastern representative of the genus, is * Details from six skulls: — one jiiT. (not quite full-grown), two youug adults: p' pre.-ent on both sides; one young adult : p' pre.seut ou one side CiAy ; two adults (teeth slightly worn) : p' absent on botli sides.

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46 B0XI3ETXrS CELEBENSIS. very closely allicil to lu ampJe.vlccmdaUis, dilleriiig only in its smaller size, i)roportionally smaller ears, slenderer rostrum, deciduous p\ and (as a rule) narrower interspace between c and p'. T!ie western limits of its range (islands VV. and S.W. of Nevv Guinea) remain to be detei-mined ; a specimen in the Genoa Museum from Amboina (collected by 0. Beccari, and examined by the present w liter) is E. hrac/ii/otis, but its alleged occurrence in Celebes (Jentink, Notes Leyden Mus. v. ]>. ITo, l^So) probably rests on confusion with a distinct species {JL cfle/>e)i.sis, infra). a. 2 ^^sl^-' Unlie of York I. ]{,ev. J. Brown skuU. !<.'.]. ( 7//;?P of species.) b.; Duke of York I. Ee.v. J. Brown 77.7. 1«.4. skull. [C.]. c. cJad. sk-; Duke of York I. Rev. J. Brown 7.S.15.5. skull. [C.]. d-e 2jun. sks. Fauro, Solomons ; 0. M. Woodfortl, S7. 1.18.4-5. 'skulls. Mmv 18S(;. Esq. [C.]. /. Jun. sk. Gn;ul'i,,!can:u-, J. Meek, Esq. [0.]. 1.11..5.4. Solomons ; 12 May, I'JUl. 9. B,ousettiis celebensis, A'. An J. CiinoiiUctcris ample.ricnuihitd, (pt. ), Dobsou, Gat. Chir. B. M. p. 73. Cyiionycteris amplexicaudata (pt., nee Geof.}, Dohson, J. A. S. B. xlii. pt. ii. p. 20:^ (187-5 : Celebes); Jentink, Cat. Osteul. Mamm. p. 2C>S (1887: Celebes); id., Vat. >Si/d. Mamm. p. 150(1888:, Celebes); Trim/'snart, Cat . Mumvu i. \). S-i (1697 : Celebes). Roiisettus amplexieaudatus (pt.), Trouesmrt, Cat. Mamm.. .Snpjil. p. GO (1904: Celebes). Cvuoptenis (Cvnonvcteris) amplexieaudatus (pt.). TruuefSMm/. y Leyden 3f)i.'<. v. p. 17;j (1883 : N. Celebes) ; Trouessart, Cat. Manaa. i. p. 84 (pt.) (1897: Celebes). Xantbarpyia bracbyoti.s (pt.), Mat.schie, Mer/ackiroptera, p. 68 (1899: Celebes). Kousettns bracbvotis (pt.), Troncsfiaii, Cat. Mamm , Snppl. p. 60' (1904: Celebe.^). Cvnonvcteris minor {nac Dobs.), Hickson, A Naturalist in A'. Cclchcs, 'p. 84 (1889: I.). Xantbarpyin minor, A. B. Meyer, Abli. 3Lis. Dresden, vii. no. 7, p. ^(iNgO: X. Celebes, iSangbir). R-ousettus celebensis, K. Andersen, Ann. Sf Mag. iV. IL. (7) xix. p. 509 (June 1, 1907: Mt. jMasarang, Celebis). Diagnosis. — Bony palate narrower than in any of the foregoing si)ecies; last jjremolar and molars, above and below, unusually narrow. Eur longer and richer ; notopatagium partly (or wholly) hairy ; general size small. Forearm 72-5-7 >'> nim. ,'^l-ull. — General size as in Ii. ani2ih.ricau
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UOUSEXTUS CEI.IOUKNSIS. 47 ])alate unusually narrow jjosteriorly : width extornally across m--m'"' !)•? mm. (two adults), against 10-2-1 1-8 in aniple.vicKvdntus (ten adults) ; frontal region between postorbital processes flat ; temporal fossa narrow ; temjjoral ridges forming a lov/ Sigittal crest in fully mature individuals. Teelh. — Upper canine and p'' widely separated ; p' in the centre of the inters|)ace between these two teeth, not deciduous ; last premolar and molars above and Vielow very nanow : m^ at least twice as long as broad ; m^ small, less than half the size of m' ; p^ three or four times the size of a lower incisor; m.^ about half the length (or less) of m^ ; m, subcircular in outline. Pdlatc-riihjes. — 4 + '^+ 1 ; arrangement as in li. cunple.vlcaudafus. Ears, u'bu/s, tail. — Ears essentially as in li. (implea-icdudaUis: narrow, not attenuated below the tip, "the tip itself broadly rounded oft' ; antitragal lobe small, rounded. General size of the animal as ill li. bracJiifotis (smaller than E. ampIe.ricaudtUus), but digits proportionally longer than in any other eastern species of the genus (index of poUex 392, of third digit 1(546, against 33-5-41 and 1529-41 respectively in all other eastern species); for details see wing-indices, above p. 20. Tail long, probably about 20 mm. (only dried skins examined). /'MJ-.— Longer, richer, and more velvet than in 11. ctiuidexicaudatus and allied eastern species ; notopatagium clothed with dense fur : hairing on forearni.s, tibia^, interfemural, and underside of lateral membrane longer and richer ; face more densely haired. Colour. — Brighter than in B. ampJexicandatus. Back light Trout's brown, rump more inclining to mars-brown tinged with russet ; sides of back and tibial next to membranes almost vandyckhrown ; crown and occiput dark brown, approaching bistre ; nape of neck broccoli-brown ; a glandular tuft of mummy-brown hairs on each side of the neck in both sexes ; entire underside of body dark greyish drab. — Immature individuals are similar in colour to adults, but without the mummy-l)rowu neck-tuft. Measurements. See table, p. 48. llatif/e. Celebes; Sanghir Islands (Dresden Mus.). Tt/2^e in collection. liemarlcs. — Without close inspection this species, owing to its small size, may be easily (and has in fact repeatedly been) confused with It. brac]ii/otis. The larger skull, very narrow palate, narrow molars, non-deciduous p\ much longer poUex (28-30 mm., against 24-26 in hracJi;/otu-), longer wings (chiefly owing to the longer metacarpals), much louger fur, hairy notopatagium, and much more densely haired tibia readily distinguish it from B. brachi/ctls, rt. c? ad. sk. ; skull. Rurukan, N. Celebes ; Dr. Clias. Hose [C.]. 97.1.2.t>. Oct. 185)5. A. linni. sk. : skull. Jit. Masaraiig, N.Celebes, Dr. Chas. Hose [C.]. 4<)W)'; Oct. IWt.i. <;. 5 ad. sk. ; skull. Ml. Ma^arang, N. Celebes, Dr. Clias. Hose [C.]. 97. 1 .2.8. 3.300' ; Oct. 1895. ( Ti/pe of species.;

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PAGE 155

ROrSETTUS LANOSUS. 49 10. Ronsettiis (Stenonycteris) lanosus, Thos. Xantharpyia ffigyptiaca (wee Geof.), Horxfield, Cat. Mamm. Mus. E. hid. Co. p. 29 (iSol : Abyssinia). Eleutherura ae-ryptiuca (pt.), Gruy, Cat. Monk. Sfc. p. 117 (1870: Abyssinia). Cynonvcteris a?gvptiaca, Dohsim, Cat. Chir. B. M. p. 7o, specimen e ^ (1878: Abyssinia). Cynopterus (Cynonyrteris) ajsryptiaca (pt.), TroufSiart, Rev. i)May. Znol. (.3) vi. p. 206 (1878: Abyssinia). lioiisettus peo-yptiacus (pt.), Trouessart, Cat. Mainin., Sappl. p. GO (190i: Abyssinia). Eousettus lanosus, Thomas, Ann. ^3Iag. N. H. (7) xviii. p. 137 (Aug-. 1, 1906: Ruweuzori East): K. Anderson, n/j. cit. (7) xix. p. oil (1907). Diagnosis. — Basicrauial axis strongly deflected ; molars exeessively narrow. Antitragal lobe obsolete: wings from back of second toe; second phalanx of litth digit longer than (irst phalanx ; fur long and coarse; uotopatagium hairy. Forearm 88-5-90 mm. •-^ Fig. 3.— Rouset/ Its {Steiwni/cicrii;) lanosus, (^. Bmvenzori East. No G 7 1 •> Type. 1. SkitU (fig. 3). — General size as in Jl. hachi and anc/o7ensis, but more delicately built, the bones thinner and lighter. Brain-case 80 strongly deflected that the alveolar margin projected backward passes through middle or upper half of supraoccipital. Eostrum

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."(I RorsETTrs LAXOsrS. sloTulor ; ]vrenia\illaiics in simple contact (uot co-ossiiiecl ), and produced distinctly more forward than in li. Itcichi; incisive foramina, therefore, longer and broader than nsual in the genus. Owing to the strong reduction in the size of the cheek-teeth, the temporal muscle is weaker, the teraporal fossa markedly shorter and narrower than in B. Jeachi and aii
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RoUSinTl'S ANGOLliXSlS. 51 ^/lasll)^elllenh. Uii p. 54. J{ant/e. Shoa ; lluwcuzoii East (5UU0-13,000'). 'J'l/pe in collection. Eemar/,-s.—-Thia peculiar species, the type of the subgenus Steiionj/cteris, is chietly characterized by the thin, almost papery condition of the bones of the skull, the unusually strong deflection of the brain-case, the extremely narrow cheek-teeth, the practically complete obliteration of the aniitragal lobe, the insertion of the wings on the second instead of the first toe, and the long and coarse t'ur. Though much reduced in size the molars do not difi'er in structure from those of otlier species of the genus. In the quality of the fur It. laiwsus is unlike any species oi liousettus, but closely similar to the coarse-haired species of Pteropus. c7. Iiiiiu. sk. ; skull. Slioa. Sir W. Cornwallis 61.2 30 6. Harris [C.]. h. Jatl. al.; -kuU. Ruwenzori East. Eiivvenzori Ex6.7.1.2. 13,000' (H. B. ploration Coinm. Wooinum). [P.]. c-U. !;^ail., 5;? jnv. ill. Ruwenzori p:ast, Ruwenzori Ex6.7.1.3-4 12..5()013,000' ploration Comm. {P. B. Woomiam). fp.]. e. $ inini. sk. ; Ruwenzori East, Ruwenzori Ex6.124 11 skull5000"; 13 Man -h, ploration Coniuj. I'.IOO (R. E. Bent). [P.]. 11. Rousettus (Lissonycteris) angolensis, Boauje. Cynonycteris jegvptiaca, Bocaye, Joni. Sri. Math. Lishoa, (2) i no 1 p. 15 (1889 : Pmigo Andongo). " ' ? Cynonycteris unicolor (nee Gray), Matschie, Arch. Xatitra i 3 p. 351 (1801: Cameroon). ' J• • Cynonycteris sp., Bocar/e, Jorn. Sci. Math. Lhhoa, (2) ii. no. 7, p. 174, lig. 2 (palate-ridges) (1892 : Pungo Andongo, Cahata,' Qiiibula). Xaiitharpyia collaris (nee III.), Matschie, Sam. I). Ost-Afrika,v. 17 (1895: Tanga, Bukoba). ^ Cynonycteris collaris, SJiistedt, Bih. K. Sv. Vet.-Akad. Hamll. x.viii. Afd. iv. no. 1, pp. 13, 15 16. 46 (1897 : Cameroon) ; id., Mitth, Deutsch. Schutzgeb. x. Heft 1, p. 7(1897: Cameroon). Rousettus coUnri.s, Thomas, in U. H. Johnstons ' The Uqanda Proleclorutc; l.\^.422 (\m-2). Cynonycteris angolensi.x, Bocaye, Jorn. Sci. Math. Lisbua, (2) \-. no. 19, p. 133, 138, text-lig. (palate-iidges) (1898: i'lmoo Andongo, Cahato, Quibula) ; Seahra, ibid. pp. 159, 109, pi.' i. tig. 9 (palate-ridges) (1898). Xautliarpyia (Myouvcteri.s) angolensi.'*, Mats,hi/>, Mcnachiroplera p. G4 (1899: Angola, Togo, Jjukoba). Rousettus angolensis, Troue.
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0:i ROUSETTXJS ANGOLEXSIS. ]>, subequal in bulk to a lower incisor. Antitragal lobe distinct; wiugs from back of second toe ; lower leg ver_v short (2^-41 mm.) ; lur long and silky ; notopataginm hair}-. Smaller than R. Jeaclii : forearm 78-83'5 mm. Skull (fig. 4). — Brain-case only very slightly deflected, the alveolar line if continued backward passing below, or through, the lower margin of the occipital condyle. Occipital and hinder parietal region considerably more flattened than in any other species of the genus, supraoccipital much lower, outline of lambdoid crest, in back view, therefore more flatly convex. Owing to the combined eff"ect of the slight deflection of the basicranial .axis and the flattening of the Fig. 4. — Bo-iiKetfiis (Lisscnyctcris) angolends, J. Riiwenzori Eas-t. No. 6.1"2.4.ri. |. (A supernumerary upper molar present on one side in the skull figured has been omitted in the drawing ; cf. antea, p. 18.) occiput, this latter portion of the skull is directed more backward, less obliquely downward, than in other species. Premaxillaries firmly co-ossified in front, forming an unbroken bar without any trace of a median suture, even in slightly immature individuals ; ascending rami of premaxillaries slenderer than in other species, particularly in their up])er half. .Supraorbital margin and bases of postorbital processes more raised above the le\cl of the upper surface of the skull, making the frontal plateau between the processes very distinctly concave from side to side. Palation border and mesoptei'vgoid fossa rather narrower than usual.

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RousnTTUs AXGni.KNsis. 0:3 Ticlh (tig. 4). — c-p'' aiifl i>''-])', as well us c p, and \>-\\ more broadly separated than iii li. Icnchl. p' and pj more reduced in size, p, in cross-section only equalling or slightlj' exceeding a lower incisor. Molariform teeth, above and below, peculiarly short and broad, in the upper jaw almost squarish. Pulate-ridi/es. — H-\--i-\-2 (three of four alcoholic specimens) or, the fourth ridge being not interrupted in the median line, 4-|-3-|-2 (one specimen). Ears. — Outer margin of ear-conch very slightly attenuated below the tip, the tip itself broadly rounded off; anr.itragal lobe well developed, triangular, subacute. Wiiir/s. — Inserted on back of second toe. 12-15 long vertical fascise in the lateral mcmbratie, viz. 3—5 postanconeal, 8-1 1 preanconeal (five alcoholic specimens examined). Forearm shorter than in li. leachi : 7>S-83-5 mm., against 89-99. All digits proportionally considerably longer than in R. leachi : itidex of pollex 440 (361 in leac^n) ; second digit (index 786) decidedly longer than third metacarpal (index 705 ; in leachi respectively 652 and 644) ; second phalanx of fifth digit as a rule longer than first ])halanx. For further details see wing-indices, supra, p. 20. Tail and hind limb. — Tail shorter than in R. leachi : S'5-13 mm., against 15-19. ],ower leg much shorter: 2S^-33 mm., against 40-42 in R. leachi. Far. — Ears naked posteriori)', except at base. Face in front of and below eyes much more strongly haired than in R. leachi. Fur of body much longer, softer, more silky. Distribution of fur on humerus and forearm, above and below, as in R. leachi. Notopatagiuni densely haired. Upperside of femur, tibia, and interfemoral (a narrow portion next to caloar excepted) long-haired. Colour. — Different from that of li. leachi : general aspect, chestnut above, dark wood-brown below. Adult male. — IJack and rump rich brown, approaching chestnut or burnt-umber. l)ase of hairs light drab or wood-brown, sometimes with a tinge of fawn ; crown dark grizzled brown ; fur on forearrr., sides of back next to membranes, interfemoral and tibia usually of a brighter tinge, approacliing mars-brown or russet. Breast and belly wood-brown ; throat and foreneck dark mars-brown tinged with fawn. — The bleached coat ( cj ad., liuwenzori East, ilareh 13, no. presents a very different aspect : back and rum]) rich tawny olive : crown and occij)ut dark raw umber ; breast and belly as above : throat and foreneck cinnamon. Adult female. — Similar to the male, but throat and foreneck scarcely differing in colour from the i-est of the underparts. Mea.'
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54 EOrSETTUS ANGOLENSi:^ the slight deileeliou of the brain-case and the shape of the occipital and hinder parietal region it shows distinct leanings towards the genus Epomophorus. a. 2 imm. al.; Quibula, Beiiguela. Lisbon Museum [P.]. skull. {Cotype of species.) A-e. 3 (^ ad., 2 Ruwenzori East, Ruwenzori Exploration ad. sks.; 5000-.5500^ March, Coium. [F.]. 1906 (/?. K. Dent). Euwenzori p]ast, Riiwenzori Exploration o500' ; March, 1906 Couim. [P.]. {D. Carruthen). Ruwenzori East Ruwenzori Exploration E. Dent). Comni. [P.]. 114. Kodja, Gaima Alexander-Gosling Ex7.7.8.24. Range, N.E.Congo, peditiou [P.]. 3° 30' N., 29= E. ; 26 Aug. 1906. Measurements of E,ousettus lanosus and angolensis. hh

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.5/J 3. BONEIA, Jcntlnl: Type. 1879. Boneia, Jentink, Notes LeAfdvn Mm. i. p. 117 (April, 1879) .' B. biden?. Diagnosis. — Closely allied to Housfttas, with wliich it accords in most of its cranial and dental and practicalljall external characters (second digit clawed, a short tail, membranes from sides of back), but differing in the following particulars : palate much broader anteriorl}', premaxilla; separated in front, upper and lower canines excessively heavy at base, lower canines directed strongl}outward, inner pair of upper incisors lost (at least in adults), outer pair of lower incisors larger than inner, crowns of molariforra teeth flatter. General size (of the single species known) as the larger species of llousettaa: forearm about 95 mm. Fig. 5. — Buncia bidcna, (^ . Meiiado. No. \)~.l.'2.i) (type of B. iiiciiadnisk). \, >Skull (tig. 5). — DifTcring from that of Jioastttu^ in the two cliaracters referred to in the diagnosis above ; in all other respects

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5f? EONEIA. perfectly Rousetlinc. In liouscltus tlic breadth of the pnlat.e ( palatal plates of niaxilhe) anteriorly between the cingnla of the canines is lews than, or at most equal to, half the breadth between m'^-ra-, the maxillary tooth-rows therefore very conspicuously converging in postero-anterior direction, and the rostrum, in ujiper view, distinctly tapering anteriorly, the breadth across the outer surfaces of the canines being much less than the lachrymal breadth. In Boneia the palatal breadth between the cingula of the canines is about two-thirds the breadth between m"-m", the maxillary tooth-rows not conspicuously converging postero-anteriorly, the rostrum in u])per view, owing both to the greater distance between the canines and to the unusually heavy sockets of these teeth, not distinctly tapering anteriorly, the breadth across the outer surfaces of the canines subequal to the lachrymal breadth. Premaxilla? separated anteriorly by a space of about 1 mm. Deflection of brain-case greater than in typical Rovsettus, nearly as in Steiionycteris {liousettvs lunosus, p. 49, fig. '•'>), alveolar lino if projected backward passing through supraoccipital near upjier margin of foramen magnum. Occiput not tubular (comparo Ptn-opus). Length of rostrum much greater than lachrymal breadth ; front of orbit above middle of m'. Palate, apart from greater breadth iu front, quite as in Mmtsetfus, postdental portion not long as in Pteropus. Premaxilhe perfectly similar in shape to those of Eidolon and Roiiseitus, narrow, u])per extremity curved forward and tapering to a point. Tympanic unknown. Sagittal crest low, scarcely raised above level of brain-case ; postorbital processes as in llousettus. Coronoid low, much sloping, rather similar to that of Stenonycteris, though still lower and somewhat broader antero-posteriorly ; condyle of mandible below level of alveolar line. Dentition (fig. S).-.-!--^!'-^^-'"'— x2 = 32. p' and m=, i^i,cp,p3p,m, m,m3 Pj and 51)3 reduced. Dilferential characters, as compared with dentition of liovsetfus, see diagnosis of genus, siiprci. i^ small, subterete, crown very slightly differentiated from shaft, cutting-edge rounded off; premasilla projecting on inner side of i'^ (see fig. 5), suggesting the possibility that a rudimentary i' may be present in young individuals, i^ similar in size and shape to i"; i„ scarcely differing in general shape, but higher and considerably larger, crown area twice or twice and a half that of i^. Upper canine long, very broad f antero-posteriorly) at base, compressed from side to side, its socket conspicuously projecting on side of rostrum ; anteroposterior basal diameter subequal to length of ji' (in 7^o«'4W^?/.s about two-thirds the length of p''); profile of front margin of crown straight ; anterior surface of crown marked with a broad and deep vertical groove : a similar deep but narrow vertical groove on hinder surface of crown ; cingulum narrow. Lower canine very heavy at base, crown slanted strongly outward.

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p' slightly larger ill crown area lliaii i^ p, about twice tlie bulk of i,. p^-in* and p^-m., similar to corresponding teetli of liouscttufi, but; crowns Hatter, longitudinal ridges lower; outer and inner ridge of ]>' and Pj less completely united anteriorly than in liousettjis ; m" smaller, scarcely one half the area of m' ; m, little more than half the bulk of m^ ; ra^ similar in size to p,. Falate-ridijes. — 4 + 3 + ? (formula derived from Jentink's description, I. s. c). Wings. — Second digit clawed. Lateral membranes arising from sides of back, separated by a space about 15-17 mm. in width, and inserted posteriorly on back of first (or junction between first and second) metatarsal. Relative lengths of metacarpals and phalanges essentially as in Stenoniidans (Bousetfy.s lauosns, see p. -0). The subjoined wing-indices are calculated from measurements of only two specimens : — 1 ,, Pollei

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53 BOXEIA It I HENS. lower canines and somewhat enlarged outer lo\\'er incisors make the sympliysis of the mandible distinctly broader and longer (anteroposteriorly) than in Bougeitu,^. The tip and trenchant hinder edge of the lower canines has carved a deep vertical groove in the front face of the crown of the npper canines. The heavy canines and anteriorly protruding lower jaw render this portion of the skull and dentition somewhat bulldog-like in appearance. The cheek-teeth are relativeh" weak (longitudinal ridges unusually low), and, in accordance with this, the coronoid process low and much sloping, the deflection of the brain-case greater than in any species of Eousettus, except the weak-toothed E. lanosus. — The separation of the premaxillae might suggest aflinities with Eidolon, the somewhat enlarged i., with Pleropus ; but in no other characters does Boneia show any leanings whatever to these genera. Hisloiv/ of c/emis in literature. — .Secondarj' references: Dobson, Hep. Brit. Assoc, for 1880, p. 174 (near Eousettus) ; Winge, E Mus. Lundii, ii. pt. 1, p. 59 (1892 : near Eousettus) ; Matschie, Megachiroptera, p. 69 (1899: near Eousettus); Miller, Fam. & Gen. Bats, p. 61 (1907 : near Pteropns), 1. Boneia bidens, Jtntlal-. Boneia bidens, Jcntinli, Xoteg Leyden Mus. i. p. 117 (April, 1879: Bone, N. Celebes) : Dobson, Eep. Brit. Afsoc. for 1880, p. 174 (1880) ; Jentinl; Cat. Ost. Mnmm. p. 264 (1887 : Gorontalo) ; id., Cat. Sijst. Mmnm. p. 152 (1888 : Gorontalo ; Bon^) ; Floiver cV Lydehker, Mnmm. p. 653 n891) ; Trouesscni, Cat. Mamm. i. p. 85 (1897 : Bone) ; Matschie, Mef/achir. p. 69 (1899 : Bone) ; Troiiessart, Cat. Mamm., Suppl. p. 60 (1904: Bone): Willink, Nat. Tijd. N,;derl. Ind. Ixv. p. 275 (1905 : Celebes) ; Miller, Fam. ^'Gen. Bats, p. 62 (1907). Boneia nienadensis, 2^homa.i, Ann. S,Mag. N. H. (6) xviii. p. 242 (1 Sept. 1896: Menado); Troi/e.-^sart, Cat. Mamm. i. p. 85 (1897: Menado); Matschie, Majacldr. p. 69 (1899: Menado); Trouessart, Cat. Mamm., Suppl. p. 60 (1904: Menado); Willink, Nat. Tijd. Nederl. Incl. Ixv. p. 275 (1905: Menado); Miller, Fam. ^ Gen. Bats, p. 62 (1907). Forearm about 95 mm. Ears. — Similar in size to those of Eousettus leacJii, but much more distinct!}' attenuated above, tip narrowly rounded ofl' ; antitragal lobe small. Fur. — Short ; someivhat adpressed on back and underside of body, spreading round neck. Upperside of forearm thinly clothed with short adpressed hairs for proximal half; femur and adjoining interfcmoral furred above ; tibia naked. Approximate length of hairs, back 8, nape cf neck and belly 9-10 mm. Least breadth of furred area of back 15-17 mm. Colour. — d ad. (skin), type of i>. mcuadensis : Back deep Prout'.4

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TCINEIA lUDKXS. 59 hrowu, tinged with rus.sot o!i rump. Entire underside, from throat to interfemoral, inchiding tlanks, drab with a slight tinge of brownish, rather darker on throat and foreneck than on breast and belly. Nape and sides of neck golden bufty. A tuft of rigid unctuous hairs on each side of neck, deep ochraceous at base, golden buft'y at tip ; colour of tufts, in arranged fur, not conspicuously differing from that of surrounding hair. Occiput, crown, sides of head, and face darker brown than back. Sexual differentiation. — Females of this species are unknown. They probably differ in having the neck tufts undeveloped or less developed than males. Measurements. On p. GO. Specimens examined. Three males (two skulls), in the collections of the Leyden and British Museums, including the tyjie of tlie species and of B. menadensis. So far, these appear to bo the only specimens known (December, 1908). Range. North Celebes : Menado, Eone, Gorontalo. (" Eotie," the type locality of the species, is not, as naturally presumed by Thomas and all later writers, the well-known Bone or Boni in S. Celebes, but a mountain range and river in N. Celebes, very near Gorontalo ; sec Bosenberg, Malay. Arch. p. 257, 1878, and P. & F. Sarasin, Beisen in Celebes, i. map iii, 1905.) Tijpe in the Leyden Museum. Boneia hidens, Jentink ; 1879. — Type locality. Bone, N. Celebes ( Bosenberg) ; type, an adult male, preserved in alcohol, skull in situ. The second Leyden specimen, also a male, is mounted, and ticketed Gorontalo {Rosenberg) ; skull extracted, very incomplete. Boneia menadensis, Thomas ; 1896. — Type locality, Menado, N. Celebes ; type in collection. In this specimen the golden buffy colour is restricted to the nape and sides of the neck, whereas in the type of B. bidens it extends to the occiput ; in all other characters the two specimens are identical. In view of the fact (which was not known to Thomas) that B. menadensis is very nearly a topotype of B. bidnis, there can bo no doubt that the difFerence referred to is only individual. a. S ad. sk. ; skull. Menado. N. Celebes, Dr. Chas. Hose [C.]. 3500' ; Oct. 1895. {Type of B. menadensis.)

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60 BONKIA BIDBXS. Measurements of Jioneia bidens i 1

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61 4. PTEROPUS, Brisson. Pternjnis (pt.), Dobsoii, Cat. Chir, B. M. p. 15. 17(}'2. Pceropus, Bn's-sv/i, llr
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02 witliiu sliort^. distance of correspoiidini;' small ijrocx'sses ou zygoma. Cororioid process moderately strong, somewhat sloping ; coronoid Leight of mandible less than length of lower tooth-row, c-ra^, subequal to c-m'; condjle of mandible considerably above level of alveolar line. Pig. &.^-Ticntpus JnnKmiclanvf. fomesi, ^ . '-' ' ::' ' ". No. 94.7.] 4.2. Dai-vel B.iv, X.E, Borneo. Tlifi typical Pteropine skull (without, or with only slight, modifications) is found in the Lirge njajority of species of the genus, viz. in all species of the Pi. Injpomehinvx (except /'/. S)ih,ii;/fr), mar'iannus.

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rTEKoi'is. 63 Ciiniirps, ritfi(s, melanotus, niel(inoj)ogon (I'ostriun somewhat shorteucd ill I't. Jcei/ensis), vampyrus^ alecto, i'onspicillatiix, and neohibernicus groups. Leaving all minor differences out of consideration, the prii]ci[)al aberrations from this general type may be classed under the foHowing headings [(Jj) and (4)] : — (3) Modifications of skull in species with weak dentition (figs. 70, 8 C). — Species with small or excessively narrow premolars and molars sho«invariably all, or most of, the following modifications of the skull : — Deflection of brain-case conspicuously greater than usual : alveolar margin if projected backward passing through middle or Tipper half, or even through upper margin, of sujiraoccipital ; coronoid process of mnndiblc weak and much sloping; condyle of mandible situated lower than usual, at level of or only slightly above alveolar line ; temporal ridges more or less closely approximated, but not fused to form a sagittal crest, or not fused throughout their whole length ; brain-case much less constricted in front than usual, making postorbital broader than interorbital constriction. Extremes of these modifications of the skull are seen in Ft. scapalatus, tuoodfordi, and jiersonatas (cheek-teeth excessively narrow), Pt. sidmlger, molosahms, insrdaris, plueocephalas, temmiiicJci, epidarins, and mncrotl'i (teeth smaller than usual). A similar adaptation of the skull is shown by the extremely narrow-toothed liouseitus lanosus (fig. 3, p. 49) compared with the normal-toothed species of llousettus (cf. also ilacroglossime). Some species with weak dentition show no appreciable, or at least no considerable, reduction of the length of the rostrum ; in these, the teeth are iiormally spaced or may even be more spaced than usual (Pt. scapidatu:^, tvoodfordi). Generally, however, weak dentition is combined with a conspicuous shortening of the rostrum and consequently rather more crowded arrangement of the teeth. In a few species the shortening of the rostrum seems to be due chiefly to eidargement of the orbits (Ft. temmincli, personcaus, e2ndarit's, nuicrotis). (4) Short rostrum combined with heavy coronoid process (figs. 7B, 813). — In the typical Pteropine skull (as shown in figs. 6, 7 A, and 8 A, 7'<. hypomelanus) the rostrum is rather long, the front of the orbital cavity approximately vertically above the front of m', the coronoid process moderately strong and somewhat sloping, tfie coronoid height of the mandible less than the length of the lower tooth-row, exclusive of incisors. In a small number of species, viz. the typical forms of the Pt. pseJaphua group (Ft. pseJaphon (fig. 8 B), 2n^06-i«s, tidiercidatus), all forms of the Pt. scunocnsiat type ( Pt. nauaiensis, moioens-is, anetinnus, fig. 7B), and the typical iorma of the Ft. lomhocens'is group (Pt. Jomhofcusis, 4-o/»7rov'»,s), these characters are modified as follows : rostrum considerably shortened, front of orbit above back, or posterior third, or even middle of \)\ coronoid process uiiusually liigh, broad (aiitero-postcriorly), and steeply ascending, coronoid height of mandible more than length of lower tooth-row, exclusive of incisors. In all of these sj)ecip.s the dcntitiim is heavier than usual, and in nearly all peculiarly modified

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64 PTEROPTTS. iu one or other direction (either by eiilnrgemeiit of the posterior basal ledges of tlie molariform teeth, or development of an inner basal ledge in the lower cheek-teeth, or tendcnc}to splitting of inner ridges of certain lower cheek-teeth, or reduction of m^ and m^, or excessive development of cingulum of canines and incisors, or by several of these modifications combined). Compare skull and numdible of I'leralojx'.v. I Fig. -A, typical Plei-oijine skull (Pt. hyjjomdn hilh tomci^i,; see text p. Gl. B, skull of a species with short rostrum and heavy dentition (Pt. cnieliam(f, 7. 1.1. 2;" 6) ; text p. 63. C, skull as modified in species with excessively weak cheek-teeth (Pt. icapulatus,&lX>A.2); text p. (iS. All figures \. The three principal types of Pteropine skulls described above as (2), (;i), and (4) must not be considered sharply separated. There is, on the contrary, a perfectly gradual transition from (2) to (3) (viz. by species showing various stages of reduction in the sizes of the teeth, and therefore also more or less pronounced leanings towards the cranial charactei's of weak-toothed species), and several transitional stages betAveen (2) and (4) (some of the most instructive examples are found in the I't. iiselajiliut group ; certain jieripheral

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VTKHol'US. g5 species of this group, viz. I>t. insularls aud jJi<>-mY/,hah(.^, C"uroli„e Islands, and Ft. laicopten'S, J'hilippines, while preserving most of the pselaphon characters in skull and dentition, show a remarkable decrease in the size of the teeth, and in these species the short and heavy rostrum of a Pi.jyseJnphon is combined with a normal or evea rather weak coronoid process: similarly in certain peripheral forms Fig. 8— A, tjpicnl Pteropine mandible (Pf. hi^pomefanus iome^i qj 7 U '), . see text p. 62. B. vnandible of a with short rostrum,' Leav; r Q">"o^'f^ ""'^ steeply ascending coronoid process (Pt. psell phon 92.2AI) ; test p. 63. C, n>anclible of a species with exces^i/. y weak clieek-teeth {Pt. mipiilatiis, ; text p. 63. All figures J. of the lomhocemls group, viz. Pt. roJricemis, Kodriguez, and molos»i,nt,, Caroline Islands). Hut any essential modifications of the skull, other than those briefly referred to abov(s *lo not occur iu tlijs genus. Dn^titioa (figs. G, f), 10).I' ^ ' P' ''' P' °^' °^"' x •^-•'4 * r 1, 'i'.iCp, p p m, m,, m, '"* • Prudimentary, deciduous ; p„ m^, and m^ much reduced in size. * Individual abnormalities in the tooth formula appear to be exceedinalv r«re „, all of P/..o;„„., except Pt.,capuk,tus (a secies cbara'tSSv d»-ek-..e.h obht.rated), and va.,p;,n,. (cio.,ely A\[J io yiy.utca.,. 0?^ F

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(it? rri:7;nns. (1) Differential characters, as compared with dentition of Eidolon osterior basal ledge (cingulum) generally obsolescent in i', narrow but distinct in i'. Diastema i"-c wide, subequal to transverse diameter of lower canine at middle, and to diastema cp'. Ujqier canines long, siibccjual to ascending branch of prcmaxilla^, slender, gently recurved, in certain species practically straight ; cingulum always distinct, as a rule prominently developed and forming a conspicuous lira oi' ledge at inner and jiosterior base of tooth : front face of crown nuirked by a broad ar'd deep vertical groove terminating a short distance above tip of tooth, inner face by a sharp median keel from cingulum to tij). p' a minute spicule, deciduous, p^ slightly shorter (antero posterior extent) than, but nearly as broad as, p^ ; ])' subrectaiigular in basal outline, distinctly longer than broad ; m' longer and slightly narrower than p' ; m"" much reduced, sulx'qual in cross-section to i\ outline of crown circular or elongate. Posterior basal ledge of p^ and p' distinct but short, marked off ])ostero-e.\ternnlly by shallow notch from base of outer main cusp of teeth. Dower incisors crowded or very narrowly spaced, in contact with or separated by minute space from canines ; crown distinctly ditferfiOO skulls pxaniinecl by llie writer, and represpnling all species and subspecies, only 10 show deviations from llie normal condition ; no less tJian live of these skulls are of Pt. scapulaft'S (total number of sliulls examined, 21). three ot Pt. gic/anleus (of a total number of 29 skulls), two of Pt. vawpi/ru!^ (15 skulls), none of other species. The aberrations found in I't. f/iff'ni/i>iis anti vamptfrus are as follows (on those of Pt. scajmlatug see tliis species, p. 40iJ) : — (1) a well developed " i.,' is present on both sides in one skull of Pt. vampi/rus (B. M. o5.12.26.y0, locality unknown); the right and left i^ are perfectly alike in size, larger than but essentially similar in form to i.,. and situated on inner side of tooth-row, at base of canines ; (2) uij and its alveolus missing on one side: one Pt. vampijrits t'(7«(^)?/r«s (jg. ad., Java, ; (3) ni^ and their alveoli missing on both sides: one Pt. f/ifiantcvs (ad., teeth slightly worn, uncertain locality.; (4) an " m^," similar i?i bulk to i., of the same skull, present on one side: one T't . t*'ing lliul " m' ie supernntnerary, and " m ' " homologous with the Moru;al m'1,

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entiatcd, :it least in i, ; usunlly a faint median notch in front edge of crown of i.^ ; i^ twice or twice and a half the hulk of, hut only slightly higher than, ij. Lower canines generally a little shorter and more recurved than upper ones, in front view more conspicuously diverging; cingulum distinct or even strong; usually a well-marked vertical groove on outer face of crown. Diastema c-p^ wide, subequal to antero-posterior basal diameter of upper canine. Pi much reduced, once and a half or twice the bulk of, but scarcely higher than, i,, permanent ; crown subcircular or somewhat elongate in outline, crushing surface concave ; nearly always situated closer to canine than to p^. p^, p^, and ni^ subequal in cross-section, longer tlian broad ; m., smaller than m^ ; m^ much reduced, subequal to i,^, outline circular or somewhat elongate. Posterior basal ledges of Pj and ]\ as in corresponding upper premolars. A typical Pteropine dentition, similar to that described above or differing only in minor details, is found in about half the number of known species, viz. all forms of the Ft. Jn/ponichnnis (except Ft. suhniyer), viaritninus. canleejjs, rufiis, melunotas. and melanopor/on groups. (3) The principal modifications of the typical Pteropine dentition may conveniently be epitomized under the following headings: — Upper incisors.— General size larger and cingulum conspicuously broader than usual, forming a noticeable shelf posteriorly, in all species of the Pt. pselaphon group, viz. Ft. pselaphon, pilosus, tuberculalus, leucopterus, 'mstdaris, and phaoirphahis. (Compare Fteralopex.) Lower incisors. — In most species i„ is about once and a half, or twice, or twice and a half the bulk of ip but in some species the disproportion in size is considerably greater. This is sometimes due chiefly to a reduction of i,, making this tooth only about \-^ of i„ : Ft. Jdiiihoceii^sis, solitarius, rodnccnsis^ and inoloxsit'iis. More frequently, however, to actual enlargement of i.^, sometimes combined with a slight reduction of i, : Ft. iiawaiettsis, mmoensb^, anetiatiHS^ psel((phon, p'dosus, tuJiercidati's, and letwopterus ; in some of these species i.. is about four or five times tlie bulk of i^, (Compare Pteralope^v.) — In Pt. srap^datus and woodfordl all incisors are reduced in size. Canines. — As a general rule, weak cheek-teeth (no posterior basal ledges, or general size of teeth conspicuously reduced) are in this genus combined with longer, slenderer, and straighter canines with narrow cr even somewhat ill-dctined (but never completely obsolete) cingulum, whereas heavy cheek-teeth (strong basal ledges) are associated with rather shorter, stouter, and more distinctly recurved canines with strong, sharply detiiicd, shelf-like cingulum. In species with very broad cingulum of the canines, the edge of the cingulum sometimes shows a pronounced tendency to subdivide into a row of small, round, more or less incomidetely separated tubercles: Pi. sumoeni^is, andianuK, psehtphon. pilosus, I, uropientjf, tubfrculatiis, iiig}dari.i, plKTocephalus (cominro with this Pfernlopex). Tn Pt. liiohts.iitiHx the crown of the canines is more strong!)' compressed fi'nm ^ide to -.idc ihnn ii^u;il. Tlic vertical gro
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68 front face of the upper canines is shallow or soraetiraes oltliteiafod ill Pi. natidis and livwgsicmei. Pi. hihercuhit^i''; is unique in the genus in having a small', but perfectly well-marked, cusp-like projection on posterior trenchant edge of upper canine above middle of tooth (compare Pterahpe.v). A' B' D' Jig. t). — p', p^, ni', profile of left row, pulate view of right row. All figures |. A, A', typical Pteropine dentition {Ft. hypcmelamu hiteus, immature, B, B', heaTj dentition, strong piisterior basal ledges {I't. ])selajilion,&d\i\\,^^1.2A.\). C, C, teeth short (anteroposteriorly) and broad, posterior basal ledges practically obliterated » {Ft. nenhibernicus, immature,, type of Ft. corenia/Ka). D, D', degenerate dentition (Ft. svaptdafus, adult, pi. — Eather larger than usual, sometimes equal to or slightly laro-er than i , and with crown distinctly differentiated in Pt. pilostui, leucopterus, ncnuciiensis, samoensis, anetianus, lomhocensis, and solitarius. p . Vnusually large in Pt. nawaiensis, scvtoeiisis, anetlamis, psdophoyi, jvlosug, tulermlatus, and leucopterus. This peculiarity is in all of these species associated with an enlargement of i.^. (Compare Pteralopex.) Posterior basal ledges of molariform teeth. — Species with typical Pteropine dentition possess short or moderately strong posterior basal ledges in p\ pS p^, and p, ; not infrequently the ledges are also more or less distinctly perceptible in m' and m,. From this general type the modihcations go in two opposite directions: —

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PTKROPITS. 69 Either towards complete or iiearljcomplete oliliteration of the ledj;;es, as in the s[)ecics of the l^t. ie of inner (right) side of tooth {Ft. anetianu.-'. adult,';r)). R, D', teeth short (ant«ro-posteriorly) and brotid ; piisterior bisnl ledges practically obliterated (PL reohiliernii-iis. immnture, 8().7.9.1. type of I'f. rvroiianis). E, E', degeuerat*.' ileutitioii (Pt. scapuUilus, adult, unusual strengthening of the ledges, as in Pt. rodrirensis, nawaimsis, sarnohms, anttliniu^, innuJdri.i, phceocephaJus, pselaphon, pih.tus, and t'lhercnlntus ; in most of these species the ledges are well-marked also in m' and nij. and not infrequently indicated in m.. Inner hasal lodge in lower molarit'orm teeth (tig. 10 (V). — I't. uiietianus is unique in the genus (but similar to Acerodon) in ha'^iug

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70 rrEKOPCTs. a bi'oarl, well-defined inner basal ledge iu p,. nip and ni^. this ledga being in reality (quite as in Acerodon) a direct continuation of the posterior ledge along the inner side of the teeth. (_)n close examination a very faint trace of an inner ledge in the lower cheek-teeth is sometimes perceptible in the related Ft. sanwensis and 2^iiosits. Traces of antero-internal basal tubercles in p', p', p,, and p^. — In a few species the antero-internal cingulnm of p^ and p'' is somewhat more differentiated than usual, i. e. more distinctly projecting as a narrow ledge : Ft. molossinus and rorh-icensis ; in the latter species the rim of this narrow ledge occasionally develops one or two minute tubercles. Similarly, the antero-internal cingulum of p^, or both p., and p^, shows in certain species a more or less noticeable tendency towards differentiation into a projecting ledge : Pf. nuniaiensis, samoensls, (t»ctinni(s, insidaris, and phaoccjihaJ us ; at least in J'f. aiieti.anns this ledge develops a minute tubercle. In Ft. pselajifion the antero-internal portion of j)^ is rather sharply marked off, by a deep groove, from the inner main cusj) of the tooth. — Compare Acej-odon. Indications of splitting of ridges of lower molnriform teeth. — In the large majority of species the inner and outer longitudinal ridges (cusps) of all molariform teeth arc perfectly simjilc. On close inspection of p, of Ft. sanioensis and in/osus (allied to Ft. j^selo'phon) a faint transverse dejiression in the inner ridge of that tooth is detectable, at least in some individuals, suggesting an initial stage towards a splitting of tlie ridge into a?) anterior and posterior portion. This leads to Ft. i^selophon, in which the inner ridge of m^ is always slightly but quite distinctly subdivided into an anterior and posterior portion. And finally, in Pt. leiicojitcrns (allied to jiseJaphon) not only the inner but, at least in some individuals, also the outer ridge of m, and m., are similarly subdivided. — Compare Fteralopex. m^ and m''.— m.^ is more reduced than usual in the Pt. rayneri group (/'('. cor/iKifuf:, rrn/nen, rHhia)nis, Ifu'eUanits, (jrandis, and chrijsoprocti(s). Eoth m.^ and m" are reduced in the Ft. loinhocensis (Pt. londiocensis, solitavius, rodricensis., and molossinns) and scaj^ulatus groups ( Pt. scapidatus and woodfordi) ; in the two latter species these teeth are quite rudimentary, and at least in I'r. scupuhtus m^ is fiometimes, but nr rarely, lost. Modifications of general shape (basal outline), and noticeable reduction of size, of cheek-teeth. — The tyjjical Ptrropine shape and size of the cheek-teeth lead, on the one side, through practically all intermediate stages to the extremely heavy dentition of a Ft. pnelaphon, and on die other through numerous transitions to the small niid excessivel}narrow cheek-teeth of a Ft. scapnlatus. Obliteration of the posterior basal ledges of the cheek-teeth (a character which of itself is perhaps indicative of a beginning degeneration of the cheek-teeth) is rather often associated with a distinct reduction of the size of the teeth : Ft. r/culdi, epalai-iuf, macrotia, poUocephalux. The cheek-teeth are small but not much narrower than usual in Ft. niolossiiii's; peculiarly sjiortened, witli rounded corners, in P/. papuKHUs and )icohdjcrnici'..^if\gs. 9C', 101) ); and so much shortened

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riKuoPUs. 71 as lo be ik^ui'Ij' si]u,iris!i in /'/. hucu/iU'nt.s ( [>;iil icuhirly in fJio U])pi'r jaw). All the check-teeth are reduced, especially in width, in PL suhnvjer. And the extreme of degeneiation is reached h}J't. pgrsonittus, sca/>nh(tHS (figs. 9 1), i)' , lU E, E'), and uwodfon/i, in which the cheek teeth are excessively narrow ; a perfect parallel to this peculiarity is seen in one species oIl Jiousetiux { li. lanosus) and in the Macroglossi ; in the general shu])e of the teeth Pt. personatus, aai/mhitus, and woodfordi bear in fact no small resemblance to Eonyder'ts sjiehp.a. It should l)e noticed that a reduction of the general size of the clieek-teeth (m length only, bieadth only, or both in length and breadth) is a character developed independently by species of entirely different groups of the genus {Pt. suhnhjer, a member of the hifpomelaaus group ; Pt. moJossinus, of the loniboceu-tis group; Pt. ]ierso)iatHS, of the teinniiucJii group; l^t. .sOTj)M?« and woodfordi, geminate species fbrming a small natural group), and therefore far from being indicative of natural relationship. The moditication is ])robably in all cases due to adaptation to a kind of food which rcijuircs less mastication than that taken by the majority of species. Fig. 11. — Palale-ritlges, typical Pteropiiie number and arrangeiueMl, funiiiila 5-|-r)-l-3 (see text, infra) {Pt. hypomelanus tomesi, 94.7. 14.^). 1. Falate-riclf/es (fig. 11). — (1) The apparently commonest formula is 5 + 5 + 3, i.e. five anterior, undivided, five middle, separated iu the median line, and three posterior, approximately wedge-shaped, situated near the hindeiborder of the i)alate ; tliis formula has been observed iu Pt. alecto, inii-tiittws, cJiri/.^ca'chen., tormosi(.s-, (jirjaiittus, i/riseus, hi/jwiiieianus, k<'t/eiislx, melaiwpoijon , iiielanotits, nttwaiciisig,, iisi laphon, rfii/neri, rudricensis, ruhianus, ton(janus, and v((>upi/ru,t. In some of these species there is a more or less incomplete, sometimes nearly fully developed, additional ridge between tiie normal ninth and tenth ridges (formula approaching 5 + 6 + 3: Pi. toni/cnnis. ;/i
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(2) o 44-)2 (or 3) pnlafe-ridges have been found in six species : A"/, molossinu.-i, ingul.nri!:. phcfocejjhtihis, epidarius,, woodfordi ; in at least two of these (Pt. insidaris anA pJuvocephahis) the formula appronches 6 + 3 + 2, owing to the sixth ridge being only very alightl}' inteiTupted in the middle. (3) In two species, Pf. papwinns and 7uohd)ernicvs, the number of middle divided ridges is noticeably larger than usual, formula 5 + 8 + 3. — The palate-ridges have been examined by the present writer only in tho tJnrty-one species referred to above, but inasmuch as these rei)resent practically all groups of the genus, it appears unlikely that any formula essentially difieront from those given above will be found. E:trx. — In most species moderate in length (not reaching eye, when laid forward), subtriangular, exposed ; inner margin convex from base to tip, outer margin convex in lower two thirds, straight or very flatly concave in upper third ; tip obtusely pointed or .somewhat narrowly rounded off. The deviations from this general type go chiefly in two opposite directions: — (1) Ears considerably reduced, half exposed, or so much reduced as to be nearly concealed in the fur ; this modiflcation is predominant in two Polj'^uesian groups, the Pt. samuensis and ps^elaphon groups, and occurs also in a few members of other groups, e. g. Pt. snhnir/er, w
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Wil.hiu thegenus Plcro/'U.^the wing-structure is rather uniform, the variations chietlj' confined to the greater or lesser relative length of the terminal phalanges, and even in this respect the variations are rather inconspicuous. .Subjoined the wing-indices of Ft. hiJj>omeJanus, calculated from measurements of 56 adult specimens representing the eleven known races of the species: —

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Colour. — The most churucteri.slic ieuf.ure is [he clcvelcjuueiil, in a majorit}of species, of a brilliimtly coloured mantle (nape of neck), contrasting with the colour of the back *. liut the character is far from being constant ; the mantle shows any colour tinge, from light yellowish buff or even cream-butf ur whitish, through any shade of buff, tawny, russet, or chestnut, to dark brown, seul-brown, or blackish. liy darkening of the normal biiglit colour the mantle may become similar to the back ; if at the same time the colour of the back is lightened, the mantle may even become much darker than the back {Pt. aruensi:;, some races uf Fl. Jii/pomehinus, and others). — The numerous variations in the total aspect of the colour of the pelage may be roughly classed as follows: — (1) PeLige approximately dark brownish above and beneath (often sjirinkled occuiring in the Liu-kiii Islands, viz. Pf. dasymalltui. has long and spreading fur extending upon the upperside of the tibiie, and that the lliree Masciirene species are all hairy-legged (one of them, Pt. suhuiyer, a s)>ccies hiding in caies, also long-furred), and it is probably on these facts that Dobsou based his statement. But tlie second species inhabiting the Liu-kiu Islands. Pt. loochoensis, has tiie fur rather short and the tibi.T naked above; one of the two species known from Aneiteum (New Hebrides) is long-furred and )iah"y-legged (7Y. anetianii>i), the other very short-furred and naked-legijed {Pt. gcddici); of the five Australia.n species, one is long-furred with the tibia; luiusually thickly clothed {Pt. iio'iocephalu)^), in a second the fin* of the body is moderate in length and tl)e tibi;e clotlied above {Pf. /iriiimei/.s). while the three otiiers are short-furred and naked-legged. If all the hairv-legged species are taken together, they cover the Philippines. Celebes, Moluc(';is, Lesser Sunda Islands. Bismarck Archipelago, Solomon Islands, Australia, Polynesia, Micronesia, Formosa, Liu-kiu Islands, and Masearenes. All the naked-legged and short-furred species taken together cover jtraotically the whole area inhabited by tlie genus, including nearly all the small wind-swept oceanic islands, with the onlj' noteworthy exception of tlie Mascarenes. * The dilfiirence in colour between mantle and back, so conspicuous iu many species of P/croputi, lias been interpreted as an instance of " jjrotecti ve mimicry " (in the sleeping attitude of these bats "the brightly-coloured neck and head are presented to the view and resemble the extremity of a ripe cocoanut, with which the larger species closely correspond iu size," Dobsou, Cat. Chir. B. M. p. 17). This explanation, though perfectly natural, perhaps almost inevitable, in the golden age of the theories of " natural selection," " protective mimicry," &c., would scarcely meet with universal acceptance now. It would seem, in this case, no other theory is i-equired than the very simple one : similar causes, similar effects. The real fruits and the pseudo-fruits (Pteropi) hang side by side, from the same branches, in precisely similar positions, in jjreciselj' similar surroundings, subject lo precisely the same ellects of light and shade, and so it has been for countless generations of both ; no wonder, therefore, if they become similar in colnur. It has been stated (/. .?. c), obviously to support the protective-mimicry-theory, that the contrast in colour between mantle and back is " much less developed in the smaller species of the genus" (which, "by their small size, are less easily seen "); but the statement is entirely wrong ; a strikingly bright-coloured mantle or a mantle practically similar in colour to the back are characters equally common in species of all sizes, from the very smallest, through the medium-sized, to the very largest. Also it would seem that, if Nature had given these bats a protective colour, then she would, so to say, have given with one hand and taken back the benelit of her gift with the other; the colour of some species may render them not too easily detectable for the untrained eye, but at the same time all species have so strong an odour that any of their enemies, be he man or animal, easily detects ihem with closed eyes ... , „ ,, „,,,» .„..,... .-• ,1., ,.^. .^, .-.n, ,„,,.

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with greyish hairs, pariicuhuly on uuclt'i'inirU), and viLh l)ri^htcr<;olo\ireii niaiitle (many spocies. particularly smaller, hairy-legged forms, predominant style e.g. in the Pt. samoensis group); (2) back blacki^ih or brownish (often thinly sprinkled with greyish hairs), mantle bright-coloured, underparts paler than back (many species, style purest in Pt. rufus, girinnteus, and their closest allies) ; the pale colour of the underparts may be restricted to the centre of the breast (Pt. melanotus and others), or wholly replaced by bbickish or brownish {Pt. vam/ii/rns and othors); the normally very inconspicuous pale sprinkling of the back may be greatly exaggerated, making the colour of tlie back nearly hair-brown (Pf. 7iiadicits) or even silvery greyish (some races of Pt. /u/por>ieJ(iina<, Pt. arnensii) ; (3) blackish above and beneath, generally more or less thinly sprinkled with greyish hairs, and with strongly contrasting yellow ish-bulF mantle (this stylo is found only in the eleven species of the Pt. mariannus and conspkillatn^ groups, and in Pt. macrotis and epulavius) ; (4) uniform blackish above and beneath, with or without some trace of a "tippet" (Pt. modigJiani, nafalis, some races of Pt. I'ainpi/rus, Pt. aJccfo, gotddi, lii'iny&tonei).The upperside is tricoloured (pale mantle, dark hack, pale rump) in most of the species of the Pt. rai/tieri group inhabiting the Solomon Islands ( Pf. rai/»eri, ruhidjius., Inrelhiims, gra»dis) and in a related ^oluccan species (Pt. chri/sojjroctus). The whole of the fur of the body is unusually pale, nearly silvery buff or silvery whitish, in Pt. temtniwlci, persoiuttus, and capistratvs. One species is nearly uniform buffy above and beneath (]'t. Iceijeiisis). Pt. persomitus and capistratus are unique in the genus in having the head marked with sharply pronounced dark brown stripes on pale ground (compare Stijlocteniam). Sexual differentiation. — Canines in males of nearly all species longer and heavier than in females. I'ur of mantle in most species which show no other external sexual differentiation more rigid and unctuous in males, sotter and more spreading in females (nearly all species of the Pt. hypoixehinv^, mariannus, rufus, melanotus, ntelanojtogon.alecto,conxpicillatus, a nd/ifo/aAeTMtoiis groups, but the character is not e(]ually conspicuoiis in all of these species) ; in some of these the concealed base of the fur of the mantle is dark brownish in females, contrasting with the light yellowish-lnifl: tips of the hairs, whereas in males the fur of the mantle is light-coloured to the extreme base (h. mariannus and conspicillatus groups). Males of certain species have a well-developed rigid, generally richer-coloured tuft of glandular hairs on each side of the neck, while these tufts are either entirely wanting or indistinctly differentiated in females (Ft. hruntiei'.s, cognaius, rai/neri, ruhianus, lavellanu-s, grandis, lomhocensis, soliiarius. giqanteus, ariel, li/lei, epulanus, macrotis, scapulntus, vinodfordi). — l)obson's statement (Cat. Cliir. pp. 17, 49, 54) that the mantle of the female is generally " darker or less brilliantly coloured than that of the male" must have been based on insuHicicnt material ; the individual variation in the colour of the mantle is in some species considerable, but in no species

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exftmined Ij}' the present writer is there any appreciuble sexual ditl'erence in the cvposed colour of the mantle. Vi"*/))*/^.— Malagasy region generally, but not extending to the continent of Africa : Oriental region ; Austro-Malaya ; North and East Australia (not known with certainty from Tasmania) ; western Polynesia, north-west to Volcano, Bonin, 8. Liu-kiu Islands (not Japanese main islands), and Formosa, east and south-east to Caroline, Sta Cruz, Fiji, Tonga, and Samoa Islands. See geographical review of species, infrci pp. 79-86. Affinities. — Pteropus has originated from a Itousettus-\i\e type. It comes nearest the living species of Itouseftus in the general shape of the skull, and the dental formula is the same in both genera. In at least one point Pteropus appears to be slightly more jirimitive than Jiousettus ; the t\mpanic is rather nearer in form (narrow, annular) to the generalized insectivorous type than to that of IloHsettns (broader, annular ; compare F/idolon : tympanic still broader, with short bony auditory meatus). Already in llousettus the occipital region shows a slight tendency to lengthening (compare HovskUus with Oynoptenis and allied genera) ; in Pteropus it is more distinctly lengthened into a short " tube"; the rostrum is relatively somewhat longer, the palate narrower, especially in its postdental portion, the skull on the whole (in all typical forms) more heavily built, with stronger crests, and broader and more steeply ascending coronoid process of the mandible. The dentition of Pteropus is conspicuously heavier, i.^ enlarged (a slight enlargement of ij, as compared with i,, is already seen in certain species of Nonsett^is ; compare also lioneia), p' still more reduced and generally deciduous (comj)are liovsettus hrachyotis), the cingulum of the canines and incisors stronger,' and the posterior portion of the molariform teeth generally differentiated into a more or less conspicuous basal "ledge" (in certain specialized forms of the genus this ledge is again obliterated, but even then a trace of the ledge is often detectalile in the young unworn teeth). Also externally, Pteropus is decidedly on a higher level than : the tail has disappeared, its wing-structure is slightly less primitive (see p. 72), the mnntle generally conspicuously different in colour from the back, and the size of the animals generally much larger. Both genera are distributed over continental S. Asia, Indo-Malaya, and Austro-Malaya ; but whereas Roiisettus extends to Palestine, Cyprus, and the Ethiopinn region, exclusive of Madagascar and its adjacencies, Pteropus finds its western limit in the Indian Peninsula, including Ceylon and the Maldives, is rather richly represented in the whole of the Malagasy region, but perfectly unknown in the African continent ; and eastward, liousettus does not extend beyond the Solomon Islands, and is unrepresented in Australia, whereas Ptero^ms has spread over the whole western portion of Polynesia, northward over Micronesia, and southward to Australia. Suhdivisions. — The 82 species (1*9 distinguisliable forms) are arranged in seventeen groups according to their probable natural affinities, viz. :— the Pt. hi/pojiielanvs {Hi species, 26 forms j, )nnri-

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annvx (8 species), caniccjis (3), rafus (5 species, 6 iorms),mehinotiis (5 species), melanopogon (4), ratjneri (6), lomOocemis (4), samoensis (3), padaphon (G), teumuncld (3), vampyras (5 species, 11 forms), «Zerfo (4 species), mnspicillatus ('i), ncohihernicm (2), 9?ia(To;i»(3), and scapulatus groups (2). — From a broader point of view these seventeen groui)s may be classed in three primary sections, as follows : — (1) The first six groups, the I't. hypoimlanns, marhmmis, caniceps, rufus, inelaiiotm, and inelano/togon groups (altogether 41 species, 52 forms, thus almost exactly half the number of known forms), are chiefly characterized by the typical Pteropine slcull and dentition (posterior basal ledges of premolars distinct) ; the large mujority of species are short-furred and naked-legged ; and they cover practically the whole Pteropine area with exception of the Himalayas, India, and Ceylon. (2) The second section, the Ft. rnyncri, lombocensis, sdmoeiisis, pselapJion, and tennnlncJri groups (22 species), accords with the first in having distinct, or even unusually strong, posterior basal lodges in the jn-enjolars, but the rostrum is always shortened and the dentition modified, either by reduction of m^, nr, and i,, or by enlargement of i^ and p,, or the whole of the dentition is noticeably (sometimes even excessively) weakened; nearly all species are hairy-legged; and they range over the whole of Austro-Malaya, Polynesia, and Micronesia, being represented also in the Mascarenes ; but are unknown from the whole of the Oriental region, except the Philippines (one species). (3) The third section, the Ft. vampyrus, alccio, consptcillaius, neoJiibernicus, macrotis, and icapulalns groups (19 species, 25 forms), is characterized by the practically complete obliteration of the posterior basal ledges of the cheek-teeth ; the skuU is tyiiical Pteropine (except in a few species with degenersite dentition) ; all species but one are naked-legged and short-furred ; they range over the whole of the Oriental region (being the only section represented in the Himalayas, Ijidia, and Ceylon) and Austro-Malaya, as far east as the Solomon Islands, extending also to Australia ; but is unrepresented in Polynesia, Micronesia, and the Malagasy region. Fteropus^ Brisson ; 1762. — Three species were "included by Erisson in his genus Fteropus ; the first headed " Fteropm,'' the second ' Pteropus culh riihro;' and the third " Fleropus auricuUs patulis;' The first species, as being evidently considered by Brisson the Pteropus, must technically be taken as the type of the genus, on the tautology princii)le. It is marked in Brisson's book with two asterisks (indicating it as a species examined by the author himself in the Koaumur Museum), diagnosed " Pteropus rufuK aut niger, aiiriculis brevibus aeutiusculis " ( tlie last throe woids absolutely excluding Ft. vcmpyrun), and stated to occur in the island of Bourbon ( Reunion) ; these facts are evidence that Brisson had before him a Ft. niger, Kerr (Ft. vulyco-is, E. Geoff. J, not a Jt. vampyrus. But, as might be expected, Brisson places in the synonymy of the lieunion spc(;ies also references to Seba's Can-is I'olaits Ter)iutatin.i orientalis (which is Ft. vampyms) and Linne's VespertUio cnuda nvlht (l)ased on Seba), and therelore gives as

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liabitat not only Ueunion lait al&o Ternate. — Owiuo to a uiislakcn identification of Brisson's first species of Fteropiis, tlie type of this genus has hitherto been given as Fi. vnmpyrus (Merriam, Science, (n. s.) i. p. 376, 1895 ; Miller, Fam. & Gen. Bats, p. b'o, 1907: probably by a slip Matschie gives on p. 6 of his ' Megachiroptera ' the type as Pt. conspiciUatus, on p. 12 && Ft. cdano = Ft. vampyrus). The genus Fteropus as understood by Dobson in his ' Catalogue of Chiroptera ' (187S) corresponds to the genera Ftcropvs, Acerodon, and Styloctenium of the present catalogue. The genus as hero defined is equivalent to Fteropus and Desmcdopex in Miller's ' Paniilies and Genera of Bats ' (pp. 56 and 60, 1907). Spectrum, Lacepede ; 1799. — Type (only species), '^ Spectrvm vampirus." Linne's Vespertilio vamj^yims (1758) is a composite species, including Ft. vampyrus, mger, and rufus. Of these, Ft. niger uas apparently the only species known to the early French mamnjologists from autopsy (see Brisson, /. s. c, and Bnffon, Hist. Nat. X. p. 55, who distinguishes only two species of Fruitbats, the Koiissette, which is i;nquestionably Ft. niyer, and the Rongette, which is Ft. suhmyer, l)oth from the Mascarenes), and maj' therefore be fixed as the type of Lacepede's Spectrum. — In a revised reprint of Lacepede's 'Tableaux methodiques,' published in 1802 in the Didot edition of Buff'on (Quadr. xiv. p. 18S), Daudin includes two species in the genus Spectrum, S. vampirus and S. ruhidum ; the former is Ft. niger, the latter Ft. subniger. Euiiycteris, Gray; 1866. — Type (only species), '-Fteropus phaioyjs," Tenim. Temminck's Ft. pJia-iojis, 1825 (Mon. Mamm. i. p. 178), is Pt. rufus, E. GeolT., J 803 {Ft. edwardsi, E. Geoff., 1810); Temminck's Ft. pltaiopis, 1837 (Mon. Mamm. ii. p. 65), is Ft. melanopogon, Peters. From Gray's diagnosis of Euvycterts (only important characters: uings attached to the vertebral line, the part over the back bald, or nearly so) it is clear that the " P/. pliaiops'' which he proposed to separate generically from Ft('rop)us is Ft. melanopogon.— Eunycteris has been considered a synonym of Fteropus by all authors except Jfatschie (ilegachir. pp. 6, 11 ; 1899), who revived the name for a subgenus of Fieropnis including four (in reality three) forms, Vt. melanopogoti, papuanus,