(E. GR.; M. O. B. C.)
CHIPPENDALE, THOMAS(d. 1779), the most famous of English cabinetmakers. The materials for the biography of Chippendale are exceedingly scanty, but he is known to have been the son of Thomas Chippendale I., and is believed to have been the father of Thomas Chippendale III. His father was a cabinet-maker and wood-carver of considerable repute in Worcester towards the beginning of the 18th century, and possibly he originated some of the forms which became characteristic of his son’s work. Thus a set of chairs and settees was made, apparently at Worcester, for the family of Bury of Knateshill, at a period when the great cabinetmaker could have been no more than a boy, which are practically identical with much of the work that was being turned out of the family factory as late as the ’sixties of the 18th century. Side by side with the Queen Anne or early Georgian feeling of the first quarter of the 18th century we find the interlaced splats and various other details which marked the Chippendale style. By 1727 the elder Chippendale and his son had removed to London, and at the end of 1749 the younger man—his father was probably then dead—established himself in Conduit Street, Long Acre, whence in 1753 he removed to No. 60 St Martin’s Lane, which with the addition of the adjoining three houses remained his factory for the rest of his life. In 1755 his workshops were burned down; in 1760 he was elected a member of the Society of Arts; in 1766 his partnership with James Ranni was dissolved by the latter’s death.
It has always been exceedingly difficult to distinguish the work executed in Chippendale’s factory and under his own eye from that of the many copyists and adapters who throughout the second half of the 18th century—the golden age of English furniture—plundered remorselessly. Apart from his published designs, many of which were probably never made up, we have to depend upon the very few instances in which his original accounts enable us to earmark work which was unquestionably his. For Claydon House, the seat of the Verneys in Buckinghamshire, he executed much decorative work, and the best judges are satisfied that the Chinese bedroom there was designed by him. At Harewood House, the seat of the earl of Harewood in Yorkshire, we are on firmer ground. The house was furnished between 1765 and 1771, and both Robert Adam and Chippendale were employed upon it. Indeed, there is unmistakable evidence to show that certain work, so closely characteristic of the Adams that it might have been assigned to them without hesitation, was actually produced by Chippendale. This may be another of the many indications that Chippendale was himself an imitator, or it may be that Adam, as architect, prescribed designs which Chippendale’s cabinetmakers and carvers executed. Chippendale’s bills for this Adam work are still preserved. Stourhead, the famous house of the Hoares in Wiltshire, contains much undoubted Chippendale furniture, which may, however, be the work of Thomas Chippendale III.; at Rowton Castle, Shropshire, Chippendale’s bills as well as his works still exist.
Our other main source of information isThe Gentleman and Cabinet Maker’s Director, which was published by Thomas Chippendale in 1754. This book, the most important collection of furniture designs issued up to that time in England, contains one hundred and sixty engraved plates, and the list of subscribers indicates that the author had acquired a large and distinguished body of customers. The book is of folio size; there was a second edition in 1759, and a third in 1762.
In the rather bombastic introduction Chippendale says that he has been encouraged to produce the book “by persons of distinction and taste, who have regretted that an art capable of so much perfection and refinement should be executed with so little propriety and elegance.” He has some severe remarks upon critics, from which we may assume that he had already suffered at their hands. Perhaps, indeed, Chippendale may have been hinted at in the caustic remarks of Isaac Ware, surveyor to the king, who bewailed that it was the misfortune of the world in his day “to see an unmeaning scrawl of C’s inverted and looped together, taking the place of Greek and Roman elegance even in our most expensive decorations. It is called French, and let them have the praise of it! The Gothic shaft and Chinese bell are not beyond nor below it in poorness of imitation.” It is the more likely that these barbs were intended for Chippendale, since he was guilty not only of many essays in Gothic, but of a vast amount of work in the Chinese fashion, as well as in the flamboyant style of Louis XV. TheDirectorcontains examples of each of the manners which aroused the scorn of the king’s surveyor. Chippendale has even shared with Sir William Chambers the obloquy of introducing the Chinese style, but he appears to have done nothing worse than “conquer,” as Alexandre Dumas used to call it, the ideas of other people. Nor would it be fair to the man who, whatever his occasional extravagances and absurdities, was yet a great designer and a great transmuter, to pretend that all his Chinese designs were contemptible. Many of them, with their geometrical lattice-work and carved tracery, are distinctly elegant and effective. Occasionally we find in one piece of furniture a combination of the three styles which Chippendale most affected at different periods—Louis XV., Chinese and Gothic—and it cannot honestly be said that the result is as incongruous as might have been expected. Some of his most elegant and attractive work is derived directly from the French, and we cannot doubt that the inspiration of his famous ribbon-backed chair came directly from some of the more artistic performances in rococo.
The primary characteristic of his work is solidity, but it is a solidity which rarely becomes heaviness. Even in his most lightsome efforts, such as the ribbon-backed chair, construction is always the first consideration. It is here perhaps that he differs most materially from his great successor Sheraton, whose ideas of construction were eccentric in the extreme. It is indeed in the chair that Chippendale is seen at his best and most characteristic. From his hand, or his pencil, we have a great variety of chairs, which, although differing extensively in detail, may be roughly arranged in three or four groups, which it would sometimes be rash to attempt to date. He introduced the cabriole leg, which, despite its antiquity, came immediately from Holland; the claw and ball foot of ancient Oriental use; the straight, square, uncompromising early Georgian leg; the carved lattice-work Chinese leg; the pseudo-Chinese leg; the fretwork leg, which was supposed to be in the best Gothic taste; the inelegant rococo leg with the curled or hoofed foot; and even occasionally the spade foot, which is supposed to be characteristic of the somewhat later style of Hepplewhite. His chair-backs were very various. His efforts in Gothic were sometimes highly successful; often they took the form of the tracery of a church window, or even of an ovalled rose window. His Chinese backs were distinctly geometrical, and from them he would seem to have derived some of the inspiration for the frets of the glazed book-cases and cabinets which were among his most agreeable work. The most attractive feature of Chippendale’s most artistic chairs—those which, originally derived from Louis Quinze models, were deprived of their rococo extravagances—is the back, which, speaking generally, is the most elegant and pleasing thing that has ever been done in furniture. He took the old solid or slightly pierced back, and cut it up into a light openwork designexquisitely carved—for Chippendale was a carver before everything—in a vast variety of designs ranging from the elaborate and extremely elegant, if much criticized, ribbon back, to a comparatively plain but highly effective splat. His armchairs, however, often had solid or stuffed backs. Next to his chairs Chippendale was most successful with settees, which almost invariably took the shape of two or three conjoined chairs, the arms, backs and legs identical with those which he used for single seats. He was likewise a prolific designer and maker of book-cases, cabinets and escritoires with doors glazed with fretwork divisions. Some of those which he executed in the style which in his day passed for Gothic are exceedingly handsome and effective. We have, too, from his hand many cases for long clocks, and a great number of tables, some of them with a remarkable degree of Gallic grace. He was especially successful in designing small tables with fretwork galleries for the display of china. His mirrors, which were often in the Chinese taste or extravagantly rococo, are remarkable and characteristic. In his day the cabinetmaker still had opportunities for designing and constructing the four-post bedstead, and some of Chippendale’s most graceful work was lavished upon the woodwork of the lighter, more refined and less monumental four-poster, which, thanks in some degree to his initiative, took the place of the massive Tudor and the funereally hung Jacobean bed. From an organ case to a washhand-stand, indeed, no piece of domestic furniture came amiss to this astonishing man, and if sometimes he was extravagant, grotesque or even puerile, his level of achievement is on the whole exceedingly high.
Since the revival of interest in his work he has often been criticized with considerable asperity, but not always justly. Chippendale’s work has stood the supreme test of posterity more completely than that of any of his rivals or successors; and, unlike many men of genius, we know him to have been warmly appreciated in his lifetime. He was at once an artist and a prosperous man of business. His claims to distinction are summed up in the fact that his name has by general consent been attached to the most splendid period of English furniture.
Chippendale was buried on the 13th of November 1779, apparently at the church of St Martin-in-the-Fields, and administration of his intestate estate was granted to his widow Elizabeth. He left four children, Thomas Chippendale III., John, Charles and Mary. He was one of the assignees in bankruptcy of the notorious Theresa Cornelys of Soho Square, of whom we read in Casanova and other scandalous chronicles of the time. Thomas Chippendale III. succeeded to the business of his father and grandfather, and for some years the firm traded under the style of Chippendale & Haig. The factory remained in St Martin’s Lane, but in 1814 an additional shop was opened at No. 57 Haymarket, whence it was in 1821 removed to 42 Jermyn Street. Like his father, Thomas Chippendale III. was a member of the Society of Arts; and he is known to have exhibited five pictures at the Royal Academy between 1784 and 1801. He died at the end of 1822 or the beginning of 1823.
(J. P. -B.)
CHIPPENHAM, a market town and municipal borough in the Chippenham parliamentary division of Wiltshire, England, 94 m. W. of London by the Great Western railway. Pop. (1901) 5074. Chippenham is governed by a mayor, 4 aldermen and 12 councillors. Area, 361 acres. It lies in a hollow on the south side of the Upper Avon, here crossed by a picturesque stone bridge of 21 arches. St Andrew’s church, originally Norman of the 12th century, has been enlarged in different styles. A paved causeway running for about 4 m. between Chippenham Cliff and Wick Hill is named after Maud Heath, said to have been a market-woman, who built it in the 15th century, and bequeathed an estate for its maintenance. After the decline of its woollen and silk trades, Chippenham became celebrated for grain and cheese markets. There are also manufactures of broadcloth, churns, condensed milk, railway-signals, guns and carriages; besides bacon-curing works, flour mills, tanneries and large stone quarries. Bowood, the seat of the marquess of Lansdowne, is 3½ m. S.E. of Chippenham. Lanhill barrow, or Hubba’s Low, 2½ m. N.W., is an ancient tomb containing akistvaenor sepulchral chamber of stone; it is probably British, though tradition makes it the grave of Hubba, a Danish leader.
Chippenham (Chepeham, Chippeham) was the site of a royal residence where in 853 Æthelwulf celebrated the marriage of his daughter Æthelswitha with Burhred, king of Mercia. The town also figured prominently in the Danish invasion of the 9th century, and in 933 was the meeting-place of the witan. In the Domesday Survey Chippenham appears as a crown manor and is not assessed in hides. The town was governed by a bailiff in the reign of Edward I., and returned two members to parliament from 1295, but it was not incorporated until 1553, when a charter from Mary established a bailiff and twelve burgesses and endowed the corporation with certain lands for the maintenance of two parliamentary burgesses and for the repair of the bridge over the Avon. In 1684 this charter was surrendered to Charles II., and in 1685 a new charter was received from James II., which was shortly abandoned in favour of the original grant. The Representation Act of 1868 reduced the number of parliamentary representatives to one, and the borough was disfranchised by the Redistribution Act of 1885. The derivation of Chippenham fromcyppan, to buy, implies that the town possessed a market in Saxon times. When Henry VII. introduced the clothing manufacture into Wiltshire, Chippenham became an important centre of the industry, which has lapsed. A prize, however, was awarded to the town for this commodity at the Great Exhibition of 1851.
CHIPPEWA1FALLS, a city and the county-seat of Chippewa county, Wisconsin, U.S.A., on the Chippewa river, about 100 m. E. of St Paul, Minnesota, and 12 m. N.E. of Eau Claire, Wisconsin. Pop. (1890) 8670; (1900) 8094; (1910, census) 8893. It is served by the Minneapolis, St Paul & Sault Ste Marie, the Chicago & North-Western, and the Chicago, Milwaukee & St Paul railways, and by the electric line to Eau Claire. The first settlement on the site was made in 1837; and the city was chartered in 1870.
1For the Chippewa Indians seeOjibway, of which the word is a popular adaptation.
1For the Chippewa Indians seeOjibway, of which the word is a popular adaptation.
CHIPPING CAMPDEN, a market town in the northern parliamentary division of Gloucestershire, England, on the Oxford and Worcester line of the Great Western railway. Pop. (1901) 1542. It is picturesquely situated towards the north of the Cotteswold hill-district. The many interesting ancient houses afford evidence of the former greater importance of the town. The church of St James is mainly Perpendicular, and contains a number of brasses of the 15th and 16th centuries and several notable monumental tombs. A ruined manor house of the 16th century and some almshouses complete, with the church, a picturesque group of buildings; and Campden House, also of the 16th century, deserves notice.
Apart from a medieval tradition preserved by Robert de Brunne that it was the meeting-place of a conference of Saxon kings, the earliest record of Campden (Campedene) is in Domesday Book, when Earl Hugh is said to hold it, and to have there fifty villeins. The number shows that a large village was attached to the manor, which in 1173 passed to Hugh de Gondeville, and about 1204 to Ralph, earl of Chester. The borough must have grown up during the 12th century, for both these lords granted the burgesses charters which are known from a confirmation of 1247, granting that they and all who should come to the market of Campedene should be quit of toll, and that if any free burgess of Campedene should come into the lord’s amerciament he should be quit for 12d. unless he should shed blood or do felony. Probably Earl Ralph also granted the town a portman-mote, for the account of a skirmish in 1273 between the men of the town and the county mentions a bailiff and implies the existence of some sort of municipal government. In 1605 Campedene was incorporated, but it never returned representatives to parliament. Camden speaks of the town as a market famous for stockings, a relic of that medieval importance as a mart for wool that had given the town the name of Chipping.
CHIPPING NORTON, a market town and municipal borough in the Banbury parliamentary division of Oxfordshire, England, 26 m. N.W. of Oxford by a branch of the Great Western railway.Pop. (1901) 3780. It lies on the steep flank of a hill, and consists mainly of one very wide street. The church of St Mary the Virgin, standing on the lower part of the slope, is a fine building of the Decorated and Perpendicular periods, the hexagonal porch and the clerestory being good examples of the later style. The town has woollen and glove factories, breweries and an agricultural trade. It is governed by a mayor, 4 aldermen and 12 councillors. Area, 2456 acres. Chipping Norton (Chepyngnorton) was probably of some importance in Saxon times. At the Domesday Survey it was held in chief by Ernulf de Hesding; it was assessed at fifteen hides, and comprised three mills. It returned two members to parliament as a borough in 1302 and 1304-1305, but was not represented after this date, and was not considered to be a borough in 1316. The first and only charter of incorporation was granted by James I., in 1608; it established a common council consisting of 2 bailiffs and 12 burgesses; a common clerk, 2 justices of the peace, and 2 serjeants-at-mace; and a court of record every Monday. In 1205 William Fitz-Alan was granted a four days’ fair at the feast of the Invention of the Cross; and in 1276 Roger, earl of March, was granted a four days’ fair at the feast of St Barnabas. In the reign of Henry VI. the market was held on Wednesday, and a fair was held at the Translation of St Thomas Becket. These continued to be held in the reign of James I., who annulled the former two fairs, and granted fairs at the feasts of St Mark, St Matthew, St Bartholomew, and SS. Simon and Jude.
CHIQUITOS(Span, “very small”), a group of tribes in the province of Santa Cruz de la Sierra, Bolivia, and between the head waters of the rivers Mamoré and Itenez. When their country was first invaded they fled into the forests, and the Spaniards, coming upon their huts, the doorways of which are built excessively low, supposed them to be dwarfs: hence the name. They are in fact well formed and powerful, of middle height and of an olive complexion. They are an agricultural people, but made a gallant resistance to the Spaniards for nearly two centuries. In 1691, however, they made the Jesuit missionaries welcome, and rapidly became civilized. The Chiquito language was adopted as the means of communication among the converts, who soon numbered 50,000, representing nearly fifty tribes. Upon the expulsion of the Jesuits in 1767 the Chiquitos became decadent, and now number short of 20,000. Their houses, regularly ranged in streets, are built of adobes thatched with coarse grass. They manufacture copper boilers for making sugar and understand several trades, weave ponchos and hammocks and make straw hats. They are fond of singing and dancing, and are a gentle-mannered and hospitable folk. The group is now divided into forty tribes.
CHIROMANCY(from Gr.χεἰρ, hand, andμαντεία, divination), the art of telling the character or fortune of persons by studying the lines of the palms of the hands (seePalmistry).
CHIRON, orCheiron, in Greek mythology, one of the Centaurs, the son of Cronus and Philyra, a sea nymph. He dwelt at the foot of Mount Pelion, and was famous for his wisdom and knowledge of the healing art. He offers a remarkable contrast to the other Centaurs in manners and character. Many of the most celebrated heroes of Greece were brought up and instructed by him (Apollodorus iii. 10. 13). Accidentally pierced by a poisoned arrow shot by Heracles, he renounced his immortality in favour of Prometheus, and was placed by Zeus among the stars as the constellationSagittarius(Apollodorus ii. 5; Ovid,Fasti, v. 414). In a Pompeian wall-painting he is shown teaching Achilles to play the lyre.
See articles in Pauly-Wissowa’sRealencyclopädieand W.H. Roscher’sLexikon der Mythologie; W. Mannhardt,Wald- und Feldkulte(1904).
See articles in Pauly-Wissowa’sRealencyclopädieand W.H. Roscher’sLexikon der Mythologie; W. Mannhardt,Wald- und Feldkulte(1904).
CHIROPODIST(an invented word from Gr.χεἰρ, hand, andποῦς, foot), properly one who treats the ailments of the hands and feet, or is consulted as to keeping them in good condition; the use of the word is now restricted, however, to the care of the toes, “manicurist” having been invented for the corresponding attentions to the fingers. The word was first introduced in 1785, by a “corncutter” in Davies Street, London.
CHIROPTERA(Greek for “hand-wings”), an order of mammals containing the bats, all of which are unique in the class in possessing the power of true flight, and have their fore-limbs specially modified for this purpose.
Fig. 1.—Skeleton and Wing-Membranes of the Noctule Bat(Pipistrellus noctula). × 1/3c, Clavicle.h, Humerus.r, Radius.u, Ulna.d1, First digit.d2, d3, d4, d5, Other digits of the fore-limb supporting wm, the wing-membrane.m, m, Metacarpal bones.ph1, First phalanx.ph2, Second phalanx.ph3, Third phalanx.am, Antebrachial membrane.f, Femur.t, Tibia.fb, Fibula.c, Calcar supporting im, the interfemoral membrane.pcb, Post-calcaneal lobe.
c, Clavicle.
h, Humerus.
r, Radius.
u, Ulna.
d1, First digit.
d2, d3, d4, d5, Other digits of the fore-limb supporting wm, the wing-membrane.
m, m, Metacarpal bones.
ph1, First phalanx.
ph2, Second phalanx.
ph3, Third phalanx.
am, Antebrachial membrane.
f, Femur.
t, Tibia.
fb, Fibula.
c, Calcar supporting im, the interfemoral membrane.
pcb, Post-calcaneal lobe.
The mammals comprised in this order are at once distinguished by the possession of true wings; this peculiarity being accompanied by other modifications of bodily structure having relation to aerial locomotion. Thus, in direct contrast to all other mammals, in which locomotion is chiefly effected by action from behind, and the hind-limbs consequently greatly preponderate in size over the fore, in the Chiroptera the fore-limbs, being the agents in propelling the body forward during flight, immensely exceed the short and weak hinder extremities. The thorax, giving origin to the great muscles which sustain flight, and containing the proportionately large lungs and heart, is remarkably capacious; and the ribs are flattened and close together; while the shoulder-girdle is greatly developed in comparison with the weak pelvis. The fore-arm (fig. 1) consists of a rudimentary ulna, a long curved radius, and a carpus of six bones supporting a thumb and four elongated fingers, between which, the sides of the body, and the hinder extremities a thin expansion of skin, the wing-membrane, is spread. The knee is directed backwards, owing to the rotation of the hind-limb, outwards by the wing-membrane; an elongated cartilaginous process (the calcar), rarely rudimentary or absent, arising from the inner side of the ankle-joint, is directed inwards, and supports part of the posterior margin of an accessory membrane of flight, extending from the tail or posterior extremity of the body to the hind-limbs, and known as the interfemoral membrane. The penis is pendent; the testes are abdominal or inguinal; the teats, usually two in number, thoracic; the uterus is simple or with more or less long cornua; the placenta discoidal and deciduate; and the smooth cerebral hemispheres do not extend backwards over the cerebellum. The teeth comprise incisors, canines, premolars and molars; and the dental formula neverexceeds i. 2/3, c. 1/1, p. 3/8, m. 3/3; total 38. Despite the forward position of the teats, which is merely an adaptive feature, bats are evidently mammals of low organization, and are most nearly related to the Insectivora.
In consequence of the backward direction of the knee, a bat, when placed on the ground, rests on all fours, having the knees directed upwards, while the foot is rotated forwards and inwards on the ankle. Walking is thus a kind of shuffle; but, notwithstanding a general belief, bats can take wing from the walking posture.
The bones of the skeleton are characterized by their slenderness and the great size of the medullary canals in those of the extremities. The vertebral column is short, and the vertebrae differ but slightly in number and form throughout the group. The general number of dorso-lumbar vertebrae is 17, whereof 12 are dorsal; the cervical vertebrae are broad, but short. Except in fruit-bats (Pteropodidae), the vertebrae, from the third cervical backwards, are devoid of spinous processes. From the first dorsal to the last lumbar the vertebral column forms a single curve, most pronounced in the lumbar region. The bodies of the vertebrae are but slightly movable on each other, and in old individuals become partially welded. The caudal vertebrae are cylindrical bones without processes; their number and length varying in allied species. The development of these vertebrae is correlated with habits, the long tail in the insectivorous species supporting and controlling the position of the interfemoral membrane which aids bats in their doubling motions when in pursuit of insects by acting as a rudder, and assists them in the capture of the larger insects. In the fruit-bats this is not required, and the tail is rudimentary or absent. In all bats the presternum has a prominent keel for the attachment of the great pectoral muscles.
The shape of the skull varies greatly; but post-orbital processes are developed only in somePteropodidaeand a fewNycteridaeandEmballonuridae; inPteropus leucopterusalone does a process from the zygomatic arch meet the post-orbital so as to complete the orbital ring. Zygomatic arches, though slender, are present in all except in some of the species ofPhyllostomatidae.
The milk-teeth differ from those of all other mammals in that they are unlike those of the permanent series. They are slender, with pointed recurved cusps, and are soon shed, but exist for a short time with the permanent teeth. In theRhinolophidaethe milk-teeth are absorbed before birth. The permanent teeth exhibit great variety, sometimes even in the same family, as inPhyllostomatidae, whilst in other families, asRhinolophidae, the resemblance between the dentition of species differing in many respects is remarkable. In all they are provided with well-developed roots, and their crowns are acutely tuberculate, with more or less well-defined W-shaped cusps, in the insectivorous species, or variously hollowed out or longitudinally grooved in the frugivorous kinds.
The shoulder-girdle varies but slightly, the clavicle being long, strong and curved; and the scapula large, oval and triangular, with a long curved coracoid process. The humerus, though long, is scarcely two-thirds the length of the radius; and the rudimentary ulna is welded with the radius. A sesamoid bone exists in the tendon of the triceps muscle. The upper row of the carpus consists of the united scaphoid, lunar and cuneiform bones.
The “hand” has five digits, the first, fourth and fifth of which consist each of a metacarpal and two phalanges; but in the second and third the number of phalanges is different in certain families. The first digit terminates in a claw, most developed in the frugivorous species, in most of which the second digit is also clawed, although in other bats this and the remaining digits are unarmed.
In the weak pelvis the ilia are long and narrow, while in most species the pubes of opposite sides are loosely united in front in males, and widely separated in females; in theRhinolophidaealone they form a symphysis. Only in theMolossinaeis there a well-developed fibula; in the rest this bone is either very slender or cartilaginous and ligamentous in its upper third, or reduced to a small bony process above the heel, or absent. The foot consists of a short tarsus, and of slender, laterally compressed toes, with much-curved claws.
Although the brain is of a low type, probably no animals possess so delicate a sense of touch as Chiroptera. In ordinary bats tactile organs exist, not only in the bristles on the sides of the muzzle, but in the sensitive structures forming the wing-membranes and ears, while in many species leaf-like expansions surrounding the nasal apertures or extending backwards behind them are added. These nose-leaves are made up partly of the extended and thickened integument of the nostrils, and partly of the glandular eminences occupying the sides of the muzzle, in which in other bats the sensitive bristles are implanted.
In no mammals are the ears so developed or so variable in form; in most insectivorous species they are longer than the head, while in the long-eared bat their length nearly equals that of the head and body. The form is characteristic in each of the families; in most the “earlet,” or tragus, is large, in some cases extending nearly to the outer margin of the conch; its office appears to be to intensify and prolong the waves of sound by producing undulations in them. In theRhinolophidae, the only family of insectivorous bats wanting the tragus, the auditory bullae reach their greatest size, and the nasal appendages their highest development. In frugivorous bats the ear is simple and but slightly variable. In all bats the ears are extremely mobile, each independently at will.
The oesophagus is narrow, especially in blood-sucking vampires. The stomach presents two types of structure, corresponding respectively to the two divisions of the order, Megachiroptera and Microchiroptera; in the former the pyloric extremity is, with one exception, elongated and folded upon itself, in the latter simple; an exceptional type is met with in the blood-suckers, where the cardiac extremity is elongated, forming a long appendage. The intestine is comparatively short, varying from one and a half to four times the length of the head and body; longest in the frugivorous, shortest in the insectivorous species. InRhinopomaandMegadermaa small caecum has been found. The liver is characterized by the great size of the left lateral lobe, which occasionally equals half that of the whole organ; the right and left lateral fissures are usually very deep; in Megachiroptera the spigelian lobe is, with one exception, ill defined or absent, and the caudate is generally large; but in Microchiroptera the former lobe is large, while the caudate is small. The gall-bladder is generally well developed.
In most species the hyoids are simple, consisting of a chain of slender, long, cylindrical bones connecting the basi-hyoid with the skull, while the pharynx is short, and the larynx shallow with feebly developed vocal cords, and guarded by a short pointed epiglottis. In the African epauletted bats,Epomophorus, the pharynx is long and capacious, the aperture of the larynx far removed from the fauces, and, opposite to it, opens a canal, leading from the nasal chambers, and extending along the back of the pharynx; the laryngeal cavity is spacious and its walls are ossified; the hyoids are unconnected, except by muscle with the skull; while the cerato-hyals and epi-hyals are cartilaginous and expanded, entering into the formation of the walls of the pharynx, and (in males of some species) supporting the orifices of a pair of air-sacs communicating with the pharynx (fig. 2).
Fig. 2.—Head and Neck ofEpomophorus franqueti(adult male). From Dobson. The anterior (a.ph.s) and posterior (p.ph.s) pharyngeal sacs are opened from without, the dotted lines indicating the points where they communicate with the pharynx;s, thin membranous partition in middle line between the anterior pharyngeal sacs of opposite sides;s.m, sterno-mastoid muscle separating the anterior from the posterior sac.
Fig. 2.—Head and Neck ofEpomophorus franqueti(adult male). From Dobson. The anterior (a.ph.s) and posterior (p.ph.s) pharyngeal sacs are opened from without, the dotted lines indicating the points where they communicate with the pharynx;s, thin membranous partition in middle line between the anterior pharyngeal sacs of opposite sides;s.m, sterno-mastoid muscle separating the anterior from the posterior sac.
The extent and shape of the wings generally depend on the form of the bones of the fore-limbs, and on the presence or absence of the tail. The wings consist of an “antebrachial membrane,” which extends from the point of the shoulder along the humerus and more or less of the fore-arm to the base of the thumb, the metacarpal bone of which is partially or wholly included in it; the “wing-membrane” spread out between the elongated fingers, and extending along the sides of the body to the posterior extremities, generally reaching to the feet; and the “interfemoral membrane,” the most variable of all, which is supported between the extremity of the body, the legs and the calcar (fig. 1). The antebrachial and wing membranes are most developed in species fitted only for aerial locomotion which when at rest hang with the body enveloped in the wings;but in theEmballonuridae, and also in theMolossinae, which are the best fitted for terrestrial progression, the antebrachial membrane is reduced to a small size, and not developed along the fore-arm, leaving the thumb quite free, while the wing-membrane is narrow and folded in repose under the forearm. The relative development of the interfemoral membrane has been referred to in connexion with the caudal vertebrae. Its small size in the frugivorous and blood-sucking species, which do not require it, is easily understood. Scent-glands and pouches opening on the surface of the skin are developed in many species, but in most cases more so in males than in females (fig. 3). As rule, bats produce only a single offspring at a birth, which for some time is carried about by the female parent clinging to the fur of her breast; but certain North American bats commonly give birth to three or four young ones at a time, which are carried about in the same manner.
Bats are divisible into two suborders, Megachiroptera and Microchiroptera.
Megachiroptera.The first of these comprises the fruit-eating species, which are generally of large size, with the crowns of the cheek-teeth smooth and marked with a longitudinal groove. The bony palate is continued behind the last molar, narrowing slowlyFruit-eating bats.backwards; there are three phalanges in the index finger, the third phalange being terminated generally by a claw; the sides of the ear form a ring at the base; the tail, when present, is inferior to (not contained in) the interfemoral membrane; the pyloric extremity of the stomach is generally much elongated; and the spigelian lobe of the liver is ill-defined or absent, while the caudate is well developed. This group is limited to the tropical and sub-tropical parts of the Eastern Hemisphere.Fig. 4.—Head of a Flying-Fox or Fruit-Bat (Pteropus personatus). From Gray.All the members of this suborder are included in the single familyPteropodidae, the first representatives of which are the African epauletted bats, forming the genusEpomophorus. In this the dental formula isi.2/2 (or ½),c.1/1,p.2/3,m.½. Tail short or absent, when present free from the interfemoral membrane; second finger with a claw; premaxillae united in front. The species are strictly limited to Africa south of the Sahara, and are distinguished by the large and long head, expansible and often folded lips, and the white tufts of hair on the margins of the ears. The males are provided with glandular pouches, situated in the skin of the side of the neck near the point of the shoulder, which are rudimentary or absent in females. In the males they are lined with glandular membrane, from which long coarse yellowish hairs project to form conspicuous epaulet-like tufts on the shoulders. The males often have a pair of air-sacs extending outwards on each side from the pharynx beneath the integument of the neck, in the position shown in fig. 2. These bats appear to live principally on figs, the juicy contents of which their voluminous lips and capacious mouths enable them to swallow without loss. The huge and ugly West African hammer-headed bat,Hypsignathus monstrosus, represents an allied genus distinguished by the absence of shoulder-pouches, and the presence of leaf-like expansions of skin on the front of the muzzle, and of distinct cusps on the outer sides of the cheek-teeth. The great majority of the bats of this group, commonly known as “flying-foxes,” are included in the typical genusPteropus, of which the dental formula isi.2/2,c.1/1,p.3/3,m.2/3. All are of large size, and the absence of a tail, the long pointed muzzle, and the woolly fur covering the neck render their recognition easy. One of the species,P. edulis, inhabiting Java, measures 5 ft. across the fully extended wings, and is the largest member of the order.The range of the genus extends from Madagascar through the Seychelles to India, Ceylon, Burma, the Malay Archipelago, Japan, New Guinea, Australia and Polynesia. Although two species inhabit the Comoro Islands, scarcely 200 m. from the mainland, not one is found in Africa; while the common Indian species is closely allied to the Madagascar flying-fox. The Malay Archipelago and Australia form the headquarters of these bats, which in some places occur in countless multitudes. The colonies exhale a strong musky odour, and when awake the occupants utter a loud incessant chatter. Wallace’s fruit-bat of Celebes and Macassar has been made the type of a separate genus, asStyloctenium wallacei. InRoussettus(orCynonycteris) the dentition is as inPteropus, but the tail is short, and the fur of the nape of the neck not different from that of the back: its distribution accords with that ofPteropus, except that it includes Africa and does not reach farther east than New Ireland.R. aegyptiacusinhabits the chambers of the Great Pyramid and other deserted buildings in Egypt, and is probably the species figured in Egyptian frescoes.Boneia, with two species, from Celebes, differs in having only two upper incisors.HarpyionycterisandScotonycteris, respectively from the Philippines and West Africa, are represented by a single species each; but ofCynopterus, which is mainly confined to the Indo-Malay countries, there are some half-score different kinds. The dentition isi.2/(2 or 1),c.1/1,p.3/3,m.3/3, the muzzle is shorter than inRoussettus, with the upper lip grooved in front as inPteropus, while the tail and fur resemble those of the former genus. These bats are extremely voracious, a specimen of the IndianC. marginatushaving eaten a banana twice its own weight in three hours. Among several Austro-Malay genera, such asPtenochirusandBalionycteris, the tube-nosed bats of the genusGelasinus(orHarpyia) are remarkable for the conformation of the nostrils (fig. 5).Cephalotes, with one species, ranging from Celebes to the Solomon group, has the dentitioni.1/1,c.1/1,p.2/3,m.2/3, premaxillae not united in front, nostrils simple, muzzle short, index finger without a claw, tail short. As inGelasinus, the wing-membrane arises from the middle line of the back, to which it is attached by a longitudinal thin process of skin; the wings are naked, but the back covered with hair.Leipenyxis an allied West African genus with one species.Fig. 5.—Head of Papuan Tube-Nosed Bat (Gelasinus major). From G.E. Dobson.The foregoing belong to the typical subfamilyPteropodinae, while the remainder represent a second group,Carponycterinae(orMacroglossinae), characterized by having the facial part of the skull produced, the molar teeth narrow, and scarcely raised above the gum, and the tongue exceedingly long, attenuated in the anterior third, and armed with long recurved papillae near the tip. The single representative of the first genus,Notopteris macdonaldi, inhabiting Fiji, New Guinea and the New Hebrides, is distinguished from other bats of this family by the length of its tail, which is nearly as longas the forearm. The dentition isi.2/1,c.1/1,p.2/3,m.2/2, while the index finger has no claw, and the wings arise from the spine.Eonycteris, with the dentitioni.2/2,c.1/1,p.3/3,m.2/3, is also represented by a single species,E. spelaea, from Tenasserim, Burma, and the Malay Peninsula and Islands, which has somewhat the appearance of aRoussettus, but the absence of a claw in the index finger and the presence of the characteristic tongue and teeth at once distinguish it.Carponycteris(Macroglossus) andMelonycteris, the former with several and the latter with a single species, are closely allied Indo-Malay and Papuan genera, the index finger in both having a claw, but the number of the teeth being the same as inEonycteris.C. minimusis the smallest known species of the suborder, much smaller than the serotine bat of Europe, with the fore-arm scarcely longer than that of the long-eared bat. It is nearly as common in certain parts of Burma asCynopterus marginatus, and extends eastwards through the Malay Archipelago as far as New Ireland, where it is associated withMelonycteris melanops, distinguished by its larger size and the total absence of the tail. An allied smallCarpopycterisinhabits India.Trygenycteris(Megaloglossus)woermanni, of West Africa, is the only member of the group occurring west of the Himalaya.Callinycterisof Celebes, with the dentitioni.2/2,c.1/1,p.2/2,m.3/3, has a short tail and no index-claws, whileNesonycterisof the Solomons, with the dentitioni.2/1,c.1/1,p.3/3,m.3/3, differs by the absence of the tail.Microchiroptera.The second and larger suborder, the Microchiroptera, includes all the insectivorous species, the majority of which are of relatively small size as compared with the Megachiroptera. In these bats, with a few specialized exceptions, the crowns of theInsect-eating bats.cheek-teeth are surmounted by sharp cusps, divided by transverse grooves. In the skull the bony palate narrows abruptly and is not continued backwards laterally behind the last molar; there is one rudimentary phalange (rarely two or none) in the index finger, which is never terminated by a claw; the outer and inner sides of the ear commence interiorly from separate points of origin; the tail, when present, is contained in the interfemoral membrane, or appears on its upper surface; the stomach, except in the blood-sucking group, is simple; and the spigelian lobe of the liver large, and the caudate generally small.The bats included in this suborder are so numerous in genera (to say nothing of species) that only some of the more important types can bementioned.Brief references have already been made to the manner in which in many or most of these bats the tail aids in the capture of prey. From the observations of C. Oldham, it appears that these bats, when walking, carry the tail downwards and forwards, so that the membrane connecting this organ with the hind-legs forms a kind of pouch or bag. If a large insect be encountered the bat seizes it with a snatch, and slightly spreading its folded wings and pressing them on the ground in order to steady itself, brings its feet forwards so as to increase the capacity of the tail-pouch, into which, by bending its neck and thrusting its head beneath the body, it pushes the insect. Although the latter, especially if large, will often struggle violently, when once in the pouch it but rarely escapes, from which it is subsequently extracted and devoured. It is assumed that the same method of capture is employed when on the wing; and a naturalist who has observed the long-eared bat picking moths off willows states that the bat always hovers when taking off the moth, and bends up the tail so as to form a receptacle for the insect as it drops.Fig. 6.—Head of Mitred Horseshoe Bat (Rhinolophus mitratus). From Dobson.In theRhinolophidae, Horse-shoe and Leaf-nosed bats of the Old World, the nose-leaf is developed and surrounds the nasal apertures, which are situated in a depression on the upper surface of the muzzle so as to look upwards; the ears are large and generally separate, without trace of a tragus or earlet; the premaxillae are rudimentary, suspended from the nasal cartilages, and support a single pair of small incisors; the molars have acuteW-shaped cusps; the skull is large, and the nasal bones which support the nose-leaf much expanded vertically and laterally. In females a pair of teat-like appendages are found in front of the pubis; and the long tail extends to the margin of the interfemoral membrane. The middle finger has two phalanges, but the index is rudimentary. The fibula is rudimentary.TheRhinolophidaeare the most highly organized of insectivorous bats, in which the osseous and cutaneous systems reach the fullest development. Compared with theirs, the bones of the extremities and the wings of other bats appear coarsely formed, and their teeth seem less perfectly fitted to crush the hard bodies of insects. The complicated nasal appendages reach their highest development, and the differences in their form afford characters in the discrimination of the species, which resemble one another closely in dentition and the colour of the fur.Fig. 7.—Head of Squirrel Leaf-Bat (Phyllorhina calcarata). From Dobson.In the first subfamily,Rhinolophinae, the first toe has two, and the other toes three phalanges each; and the ilio-pectineal spine is not connected by bone with the antero-inferior surface of the ilium. In the horseshoe bats,Rhinolophus, the dentition is i. 1/2, c. 1/1, p. 2/3, m. 3/8, the nose-leaf has a central process behind and between the nasal orifices, with the posterior extremity lanceolate, and the antitragus large. Among the numerous formsR. luctusis the largest, and inhabits elevated hill-tracts in India and Malaysia;R. hipposiderusof Europe, extending into south England and Ireland, is one of the smallest; andR. ferrum-equinumrepresents the average size of the species, which are mainly distinguished from one another by the form of the nose-leaf. The last-named species extends from England to Japan, and southward to the Cape of Good Hope, but is represented by a number of local races. When sleeping, the horseshoe bats, at least in some instances, suspend themselves head downwards, with the wings wrapped round the body after the manner of fruit bats. The posture of ordinary bats is quite different, and while the lesser horseshoe (R. hipposiderus) alights from the air in an inverted position, other bats, on first coming to rest, do so with the head upwards, and then reverse their position.Fig. 8.—Head of Persian Leaf-Bat. (Triaenops persicus). From Dobson.In the second subfamily,Hippo-siderinae(formerly calledPhyllorhinae), the toes are equal and include two phalanges each, while the ilio-pectineal spine is united by a bony isthmus with a process derived from the antero-inferior surface of the ilium.Hipposiderus, Clöeotis, Rhinonycteris, Triaenops, AnthopsandCoelopsrepresent this subfamily.Hipposiderus(Phyllorhina), with many species, ranging over Asia, Africa and Australasia, and the dental formula i. 1/2, c. 1/1, p. 2/2, or 1/2, m. 3/3, differs fromRhinolophusin the form of the nose-leaf, which is not lanceolate behind (fig. 6), and is unprovided with a central process covering the nostrils; the largest species,H. armiger, appears to be the most northerly, having been taken at Amoy in China, and in the Himalaya at an elevation of 5500 ft. Many are provided with a frontal sac behind the nose-leaf, rudimentary in females (see fig. 7), which can be everted at pleasure; the sides of this sac secrete a waxy substance, and its extremity supports a tuft of straight hairs.Rhinonycteris, represented byR. aurantiafrom Australia, andTriaenops. byT. persicusfrom Persia and other species from Africa and Madagascar, are closely allied genera.Triaenops(fig. 8) is characterized by the remarkable form of its nasal appendages and ears, and the presence of a bony projection from the upper extremity of the second phalange of the fourth finger.Coelops(C. Frithi), from the Bengal Sanderbans, Java and Siam is distinguished by the peculiar form of its nose-leaf and the length of the metacarpal bone of the index finger, as well as by the shortness of the calcar and interfemoral membrane.Clöeotisis represented by a single East African species, andAnthopsby one from the Solomon Islands characterized by the nose-leaf covering the whole front of the face.The next family,Nycteridae, which is also Old World, is a small one, nearly allied to the last, in which it is included by Prof. Max Weber as a subfamily under the name ofMyadermatinae. It differs by the presence of a small tragus inFalse vampires.the ears, which are united at their bases; and by the nasal chamber not being inflated. The premaxillae are either small and separated in front, or rudimentary; and the first phalange of the middle finger when in repose is laid back on the metacarpus. There are only pectoral teats.Of the two genera,Megaderma, as represented by the five species of false vampires, is distinguished by the absence of ossified premaxillae and upper incisors (i. 0/2, p. (2 or 1)/2), the cylindrical narrow muzzle surmounted by an erect nose-leaf the base of which conceals the nasal orifices, the immense joined ears with large bifid tragus, and the great extent of the interfemoral membrane, in the base of which the short tail is concealed.M. gigas(fig. 9), from central Queensland, is the largest species of the genus, and of the suborder.M. lyra, common in India (fore-arm 2.7 in.), has been caught in the act of sucking the blood, while flying, from a small bat which it afterwards devoured. The range of the genus includes Africa, the Indo-Malay countries and Australasia.Nycteris, which is commonto Africa and the Malay Peninsula and Islands, has ossified premaxillae and upper incisors (i.2/3,p.1/2), and a long tail; but lacks a nose-leaf. As inMegaderma, the frontal bones are deeply hollowed and expanded laterally, the muzzle presents a similar cylindrical form, and the lower jaw also projects; but, instead of a nose-leaf, the face is marked by a deep longitudinal sharp-edged groove extending from the nostrils to the band connecting the base of the large ears; the sides of this depression being margined as far back as the eyes by small horizontal cutaneous appendages. With the exception ofN. javanica, the species are limited to Africa.Fig. 9.—The False Vampire (Megaderma gigas). From Dobson.According to the classification followed by Dr G.E. Dobson, the extensive family of New World bats known asPhyllostomatidaewas widely sundered from the two preceding groups; but in Prof. Max Weber’s system they are placed next oneVampires.another—an arrangement which has the great advantage of bringing together all the bats furnished with nose-leaves. It is indeed probable that the vampires, as the members of the present family may be collectively termed, are the New World representatives of the Old WorldRhinolophidaeandNycteridae.ThePhyllostomatidaeare characterized by the presence of a nose-leaf, or of lappets on the chin, but the nostrils are not directed upwards. The ethmoturbinal bones of the nasal cavity form simple plates (much as in the two preceding families). The premaxillae are always well developed, with their palatal portions forming a suture and denning the boundaries of distinct palatine foramina (in place of being rudimentary, as inNycteridaeandRhinolophidae). The large ears have a tragus. The middle finger has three phalanges, and the index one. There is an incomplete fibula. The tail may be either long or short. Generally the dentition isi.2/2,c.1/1,p.2/3,m.3/3.Fig. 10.—Head of Blainville’s Vampire (Mormops blainvillei). From Dobson.All the bats of this family may be readily recognized by the presence of a well-developed third phalange in the middle finger, associated either with a distinct nose-leaf, or with central upper incisors, or with both. Unlike theRhinolophidae, their eyes are generally large and the tragus is well developed, maintaining almost the same form throughout the species, however much the other parts of the body may vary. Their fur is of a dull colour, and the face and back are often marked with white streaks. A few species, probably all those with the tail and interfemoral membrane well developed, feed principally on insects, while the greater number of the species of the groupsVampyreaeandGlossophageaeappear to live on a mixed diet of insects and fruits, and theDesmodonteae, of which two species are known, are true blood-suckers, and have their teeth and intestinal tract specially modified in accordance with their habits. The group is practically limited to the tropical and subtropical parts of Central and South America, although one species ofOtopterusreaches California. In the first subfamily,Mormopsinae(Lobostominae), the nostrils open by simple apertures at the extremity of the muzzle in front, not margined by a distinct nose-leaf; while, in compensation, the chin is furnished with expanded leaf-like appendages. The tail is short. It includes two genera. InChilonycteristhe crown of the head is moderately elevated above the face-line, and the basi-cranial axis is almost in the same plane as the facial, while inMormops(fig. 10) the crown of the head is greatly elevated above the face-line, and the basi-cranial axis is nearly at right angles to the facial;i.2/2,p.2/3, in both genera. As regards the species ofChilonycteris, the most striking feature is the occurrence of a rufous and a dark brown phase in each. In some the two phases are very marked, but in others they are connected by intermediate shades. Here may be mentioned the two species of tropical American hare-lipped bats, forming the genusNoctilio, which presents characters common to this and the following family, to which latter it is often referred. The typicalN. leporinusis a bat of curious aspect, with strangely folded lips, erect skin-processes on the chin, and enormous feet and claws. The two middle incisors are close together, and so large as to conceal the small outer ones, while in the lower jaw there are but two small incisors; the premolars numbering 1/2. These bats live near the coast, and feed on small crabs and fishes.Most of the remaining members of the family are included in the subfamilyPhyllostomatinae, characterized by the presence of a distinct nose-leaf and the warty chin. The clitoris is imperforate, whereas it is perforated in theMormopsinae. The incisors are generally 2/2 (occasionally 2/1), and the molars well developed. The subfamily is divided into a number of groups or sections. The first of them, theVampyreae, is characterized as follows: Muzzle long and narrow in front, the distance between the eyes generally less than (rarely equal to) that from the eye to the extremity of the muzzle; nose-leaf horseshoe-shaped in front, lanceolate behind; interfemoral membrane well developed; tail generally distinct, rarely absent; inner margin of the lips not fringed;i.2/2 or 2/1,p.2/2 or 2/3; molars withW-shaped cusps, usually well developed.Nearly all theVampyreaeappear to be insectivorous, so that the term cannot be considered indicative of habits; but a few, if not all, probably supplement their insect diet with fruit.Vampyrus spectrum(the largest bat in the New World) is said to be wholly frugivorous, andOtopterus waterhouseiappears to prey occasionally on smaller bats. The genera may be arranged in two subgroups according as the tail is produced to the margin of the interfemoral membrane or perforates it to appear on its upper surface. In the first division are included three genera,Lonchorhina,Otopterus(orMacrotus) andDolichophyllum(orMacrophyllum), the first represented byL. aurita, characterized by an extraordinary long nose-leaf, and peculiarly large ears and tragus. In the second subsection are includedVampyrus,Chrotopterus,Tonatia(Lophostoma)Micronycteris,Glyphonycteris,Trachyops,Phylloderma,Phyllostoma,Anthorhina(Tylostoma),Mimon,Hemiderma(Carollia) andRhinophylla; all, with the exception of the last, distinguished chiefly by the form of the skull and the presence or absence of the second lower premolar.Phyllostoma hastatum, next in point of size toVampyrus spectrum, is a well-known species in South America;P. elongatum(fig. 11) differs in its smaller size and larger nose-leaf.Hemiderma brevicauda, a small species, closely resemblesGlossophaga soricina, and forms a connecting link between this and the next group.Rhinophylla pumiliois the smallest species of the family; further distinguished by the absence of a tail, the narrowness of its molars, which do not formW-shaped cusps, and the small size of the last upper molar, characters connecting it and the group with theStenodermateae. Both inHemidermaandRhinophyllathe zygomatic arch is incomplete.Fig. 11.—Head of Lesser Javelin Vampire (Phyllostoma elongatura).The next subsection,Glossophageae, presents the following distinctive features: Muzzle long and narrow; tongue long and extensible, attenuated towards the tip, and beset with long filiform recurved papillae; lower lip with a wide groove above, and in front margined by small warts; nose-leaf small; tail short or none;i.2/2,p.2/3 or 3/3 or 2/2,m.3/3 or 2/3 or 2/2; teeth narrow; molars with narrowW-shaped cusps, sometimes indistinct or absent; lower incisors small or deciduous. The species included in this group represent some ten genera, distinguished principally by differences in the form and number of the teeth, and the presence or absence of the zygomatic, arch of the skull. InGlossophagaandPhyllonycteristhe upper incisors form a continuous row between the canines. InMonophyllusandLeptonycteris(Ischnoglossa) they are separated into pairs by a narrow interval in front; while inLonchoglossa,GlossonycterisandChoeronycteristhey are widely separated and placed in pairs near the canines. In the first four of these genera the lower incisors are present (at least to a certain age), in the last three they are deciduous even in youth. The zygomatic arch is wanting inPhyllonycteris,GlossonycterisandChoeronycteris. The typical species isGlossophaga soricina, which, as already mentioned, closely resemblesHemidermabrevicauda, both in form and dentition. Its long brush-tipped tongue (which it possesses in common with other species of the group) is used to lick out the pulpy contents of fruits having hard rinds. The food of the species of this group appears to consist of both fruit and insects, and the long tongue may be used for extracting the latter from the deep corollas of flowers. Other genera areLonchophylla,Rhithronycteris,HylonycterisandLychonycteris, each with a single species (in 1904).Fig. 12.—Head of Long-tongued Vampire (Choeronycteris mexicana), showing brush-tipped tongue. From Dobson.The third group,Stenodermateae, presents the following characteristics:—Muzzle very short and generally broad in front, the distance between the eyes nearly always exceeding (rarely equalling) the distance from the eye to the extremity of the muzzle; nose-leaf short, horseshoe-shaped in front, lanceolate behind (except inBrachyphyllaandCenturio); interfemoral membrane concave behind; tail none; inner margin of the lips fringed with conical papillae;i.2/2 or 2/1,p.2/2,m.3/3 or 2/3 or 2/2; cheek-teeth broad (except inSturnira), molars with concave or flat crowns margined externally by raised cutting-edges. Although theStenodermateaeare generally easily distinguished from theVampyreaeby the shortness and breadth of the muzzle and the form of the cheek-teeth, certain species of the latter resemble the former in external appearance, agreeing almost absolutely in the form of the nose-leaf, the ears and the tragus, and the warts on the chin. These resemblances show that, while the form of the teeth and jaws has become modified to suit the food, the external characters have remained much the same, and indicate the common origin of the two sections. The food of these bats appears to be wholly or in great part fruit. The species are divided into some eleven genera, mostly distinguished by the form of the skull and teeth.Artibeusincludes the frugivorousA. perspicillatus.Stenoderma achradophilum, found in Jamaica and Cuba, with the last, from which it is scarcely distinguishable externally except by its much smaller size, differs in the absence of the horizontal plate of the premaxillae on the palate.Sturnira lilium, while agreeing with these in the form of the nose-leaf and ears, differs from all the species of the family in its longitudinally-grooved molars, which resemble those of thePteropodidaemore closely than those of any other bats; and the presence of tufts of long differently-coloured hairs over glands in the sides of the neck is another character in common with that group.Centurio senex(fig. 13) is the type of a small genus distinguished fromStenodermaand other genera of this group by the absence of a distinct nose-leaf. Some naturalists make this genus the type of a distinct subgroup,Centurioneae. Up to 1904 the genera, exclusive ofCenturio, included in theStenodermateaewereArtibeus(with several sub-genera),Vampyrops(also with subgenera),Mesophylla,Chiroderma,Stenoderma(with 3 subgenera),Ectophylla,Ametrida(with 2 sub-genera),Pygoderma,SturniraandBrachyphylla.Fig. 13.—Head of Masked Vampire (Centurio senex). From Dobson.The third subfamily,Desmodontieae, is represented only by the blood-sucking bats, and distinguished by havingi.½, of which the upper pair are cutting, the rudimentary molars, the very short interfemoral membrane, and the blood-sucking habit. They are further characterized as follows: Muzzle short and conical; nose-leaf distinct;p.2/3,m.1/1 or 0/0; upper incisors occupying the whole space between the canines; premolars narrow, with sharp-edged longitudinal crowns; molars rudimentary or absent; stomach elongated, and intestiniform. There are two genera,Desmodus, without calcar or molars, andDiphylla, with a short calcar and a single rudimentary molar on each side—restricted to Central and South America.Desmodus rufus, the commoner species, is a little larger than the noctule bat, and abundant in certain parts of South America, where it is troublesome owing to its attacks upon domestic animals, sucking their blood and leaving them weakened from repeated bleedings. (SeeVampire.)The fourth family of bats, unlike any of the three previous ones, has a cosmopolitan distribution. These free-tailed bats, as they are conveniently called, constituting the familyEmballonuridae, present the following distinctive features. TheFree-tailed bats.nostrils are of normal form and without a nose-leaf. The premaxillae have their palatal portion imperfectly developed, and united by a slender process with the maxillae. The ears are large, with a small tragus. The middle finger has two phalanges, and the index generally a single one. The fibula is incomplete. The tail is generally short, and always partly free from the interfemoral membrane. There is generally only a single pair of upper incisors, separated by gaps from the canines, and from one another in the middle line.The distinctive feature of these bats is the free tail-tip, which pierces the interfemoral membrane to appear on its upper surface, and may project beyond its margin. As a rule, these bats may also be recognized by the peculiar form of the muzzle, which is obliquely truncated, the nostrils projecting more or less in front beyond the lower lip, by the first phalange of the middle finger being folded in repose forwards on the upper surface of the metacarpal bone, and by the upper incisors. Although cosmopolitan, these bats rarely extend north or south of the thirtieth parallels of latitude.Fig. 14.—Ear ofEmballonura raffrayana. From Dobson.The family may be divided into two subfamilies, of which theEmballonurinaeis characterized by the incomplete premaxillae, the presence of only one phalange in the index finger, and the short tail. The dental formula is generallyi.1/3 (sometimes 2/3 or 1/2),c.1/1,p.2/3,m.3/3. This subfamily may be further subdivided into subgroups or sections of which the first,Embalionurae, is characterized by the slender tail perforating the interfemoral membrane, so as to appear on its upper surface; the legs long, with a slender fibula; the incisors weak; and the premolars 2/2. The typical genusEmballonurapresents the following features:i.2/3, extremity of the muzzle more or less produced beyond the lower lip, forehead flat. The genus contains several species, inhabiting islands from Madagascar through the Malay Archipelago and Siam to the Navigator Islands.Coleura, withi.1/3, the extremity of the muzzle broad, and the forehead concave, has two species from East Africa and the Seychelles.Rhynchonycterisis distinguished fromColeuraby the produced extremity of the muzzle. The single species,R. naso, from Central and South America, is common in the vicinity of streams, where it is usually found during the day resting on the vertical faces of rocks, or on trunks of trees growing over water; it escapes notice owing to the greyish colour of the fur of the body and of small tufts on the antebrachial membrane counterfeiting the weathered surfaces of rocks and bark. As evening approaches it appears on the wing, flying close to the water.Saccopteryxhasi.1/3 and the antibrachial membrane with a pouch opening on its upper surface; it contains several species from Central and South America. This sac is developed only in the male and in the female is rudimentary. In adult males a valvular longitudinal opening occupies the upper surface of the membrane leading into a small pouch, the interior of which is lined with a glandular membrane secreting an unctuous reddish substance with a strong ammoniacal odour. Allied genera are the tropical AmericanPeropteryxand the BrazilianCormura. The various species of tomb-bats (Taphozous) inhabit the tropical and subtropical parts of all the eastern hemisphere except Polynesia, and are distinguished by the cartilaginous premaxillaries, the deciduous pair of upper incisors, and the presence of only two pairs of lower incisors. Most of the species have a glandular sac (fig. 15) between the angles of the lower jaw, more developed in males than in females, in some species absent in the latter. An open throat-sac is wanting inT. melanopogon, but about its position are the openings of small pores, the secretion from which probably causes the hairs to grow long, forming the black beard found in many males. The three tropical American white bats,Diclidurus, withi.1/3,c.1/1,p.3/2,m.3/3, resembleTaphozousin the form of the head and ears, but, besides other characters, differ from all other bats in possessing a pouch, opening off the centre of the interior surface of the interfemoral membrane; the extremity of the tail enters this, and perforates its base.Fig. 15.—Heads of Tomb-Bat (Taphozous longimanus), showing relative development of throat-sacs in male and female. From Dobson.The second subfamily of theEmballonuridae,Rhinopomatinae, is represented only by the genusRhinopoma, with several species ranging from Egypt through Arabia to India, Burma and Sumatra. The premaxillae (fig. 16) are complete; the index finger has two phalanges; the tail is very long and mouselike; and the dental formulai.1/2,c.1/1,p.1/2,m.2/3. Dr G.E. Dobson has remarked that these mouse-tailed bats might be elevated to the rank of a family, for it is difficult to determine their affinities, a kind of cross relationship attaching them to theNycteridaeon the one hand and to theEmballonuridaeon the other. These bats, distinguished from all other Microchiroptera by the presence of two phalanges in the index fingerand the long and slender tail projecting far beyond the narrow interfemoral membrane, inhabit the subterranean tombs in Egypt and deserted buildings generally from north-east Africa to Burma and Sumatra.Fig. 16.—Skull of Mouse-tailed Bat (Rhinopoma microphyllum). ×2. (From Dobson.)The last group, according to the system adopted by Prof. Max Weber, is that of theVespertilionidae, which includes such typical bats as the pipistrelle, the noctule, and the long-eared species. By Mr G.S. Miller1the first section of theTypical bats.family—Natalinae—is regarded as of family rank, while the last section, orMolossinae, is included by Dr G.E. Dobson in theEmballonuridae, from the typical forms of which its members differ widely in tail-structure. In this extended sense the family, which has a cosmopolitan distribution, may be defined as follows:—The nostrils are normal and without a nose-leaf. The ethmoturbinal bones of the nasal chamber are involuted. The palatine processes of the premaxillae do not form a suture. The ear is mostly large, with a tragus. The middle finger (except inThyroptera) has two phalanges. The fibula is usually rudimentary. The tail is long and does not perforate the interfemoral membrane. The incisors are generally 2/3 or 1/2, but may be reduced to 1/1 in theMolossinae.Fig. 17.—Head ofChilonatalus micropus. ×2. (From Dobson.)In the first subfamily,Natalinae, which is exclusively tropical American, the other upper incisors are separated from one another and from the canines; palatine processes of the premaxillae are at least partially developed; and the dental formula isi.2/3,c.1/1,p.(2 or 3)/3,m.3/3. In general appearance these bats recall the more typicalVespertilionidae, although the form of the muzzle is suggestive of theMormopsinaeamong thePhyllostomatidae. Again, while the form of the skull is vespertilione, the relation of the vomer to the front end of the premaxillae is of the phyllostomine type. The molars and incisors are likewise vespertilione, whereas the premolars are as distinctly phyllostomine. Finally, while the third, or middle, finger normally has two phalanges, as in typicalVespertilionidae, the second of these is elongated and inThyropteradivided into two, as inPhyllostomatidae.Fig. 18.—Suctorial Disks inThyroptera tricolor,a, side, andb, concave surface, of thumb disk;c, foot with disk, and calcar with projections (all much enlarged). (From Dobson.)The first two genera,FuripterusandAmorphochilus, each have a single species, the latter being distinguished from the former by the wide separation of the nostrils and the backward prolongation of the palate. In both the crown of the head is elevated, the thumb and first phalange of the middle finger are very short, and the premolars are 2/3. The same elevation of the crown characterizes the generaNatalusandChilonatalus(fig. 17), in which the premolars are 3/3: in general appearance these bats are very like the Old World vespertilionine genusCerivoula, except for the short triangular tragus. Lastly,Thyropteraincludes two species distinguished by an additional phalange in the middle finger and by accessory clinging-organs attached to the extremities. InThyroptera tricolor, i.2/3,p.3/3, from Brazil, these have the appearance of small, circular, stalked, hollow disks (fig. 18), resembling miniature sucking-cups of cuttle-fishes, and are attached to the inferior surfaces of the thumbs and the soles of the feet. By their aid the bat is able to maintain its hold when creeping over smooth vertical surfaces.Fig. 19.—Head ofScotophilus emarginatus. (From Dobson.)The second or typical subfamily,Vespertilioninae, includes all the remaining members of the family with the exception of the aberrantMolossinae. The upper incisors are in proximity to the canines; the premaxillae widely separated; the ears medium or large; the dental formula isi.2/3 (or 1/3),c.1/1,p.3/3 (2/3, 2/2, or 1/2),m.3/3; and the fibula very small and imperfect. All the members of this large cosmopolitan group are closely allied, and differ chiefly by external characters. They may be divided into subgroups. In the first of these, thePlecoteae, of which the long-eared bat (Plecotus auritus) is the type, the crown of the head is but slightly raised above the face-line, the upper incisors are close to the canines, and the nostrils are margined behind by grooves an the upper surface of the muzzle, or by rudimentary nose-leaves; the ears being generally very large and united. Of the six genera,Plecotus, withi.2/3,p.2/3, has three species:—one the long-eared European bat referred to above;P. macrotis, restricted to North America, is distinguished by the great size of the glandular prominences of the sides of the muzzle, which meet in the centre above and behind the nostrils; the third species being also American. The second,Barbastella, withi.2/3,p.2/2, distinguished by its dentition and by the outer margin of the ear being carried forwards above the mouth and in front of the eye, includes the European barbastelle bat,B. barbastellus, andB. darjelingensisfrom the Himalaya.Otonycteris,i.1/3,pm., 1/2, connecting this group with theVespertilioneae, is represented byO. hemprichii, from North Africa and the Himalaya, and an Arabian species. The next two genera are distinguished by the presence of a rudimentary nose-leaf:Nyctophilus,i.1/3,p.1/2, with three species from Australasia; andAntrozous,i.1/2,p.1/2, distinguished from all the other members of the subfamily by having but two lower incisors, and from otherPlecoteaeby the separate ears; the two species inhabit California. The sixth genus,Euderma, is also represented by a Californian species.Fig. 19.—Head ofCerivoula hardwickei. (From Dobson.)The second groupVespertilioneae, with about thirteen genera, includes the great majority of the species; and a large number of these may be classed underVespertilio, which is divisible into subgenera, differing from one another in the number of premolars, and often ranked as separate genera. One group is represented byV.(Histiolus)magellanicus, a species remarkable for its extreme southern range, its relatives being also South American. A second group, withp.1/2, includes the British serotine,V.(Eptesicus)serotinus, of Europe and northern Asia, and represented in North America by the closely alliedV.(E.)fuscus. In the typical group, which includes the Old WorldV. murinus, one species,V. borealis, ranges to the Arctic circle. The European noctule,V.(Pierygistes)noctula, and Leisler’s bat,V.(P.)leisleri, represent another group; and the common pipistrelle,V.(Pipistrellus)pipistrellus, yet another, withp.2/2. The only other group that need be mentioned is one represented by the North AmericanV.(Lasionycteris)noctivagans, withp.2/3. The AfricanLäephotes, the ChineseIa, and the PapuanPhiletorare allied genera, each with a single species.ChalinolobusandGlauconycterishave the same general dental character asVespertilio, but are distinguished by the presence of a lobe projecting from the lower lip near the gape; the former, withp.2/2, is represented by five Australasian species, one of which extends into New Zealand; while the latter, withp.1/2, is African. The species ofGlauconycterisare noticeable for their peculiarly thin membranes traversed by distinct reticulations and parallel lines.Scotophilus, withi.1/3,p.1/2, includes several species, restricted to the tropical and subtropical regions of the eastern hemisphere, though widely distributed within these limits. These bats, though approaching certain species ofVespertilioin many points, are distinguished by the single (in place of two) pair of unicuspidate upper incisors separated by a wide space and placed close to the canines, by the small transverse first lower premolar crushed in between the canine and second premolar, and, generally, by their conical, nearly naked, muzzles and thick leathery membranes.S. temminckiis the commonest bat in India, and appears often before the sun has touched the horizon.S. gigas, from equatorial Africa, is the largest species.Nycticejus, with the same dental formula asScotophilus, is distinguished, by the first lower premolar not being crushed in between the adjoining teeth, and the comparatively greater size of the last upper molar. It includes only the North AmericanN. humeralis(crepuscularis), a bat scarcely larger than the pipistrelle. The hairy-membraned bats of the genusLasiurus(Atalapha), withi.1/3,p.2/2 or 1/2, are also limited to the New World, and generally characterized by the interfemoral membrane being more or less covered with hair and by the peculiar form of the tragus, which is expanded above and abruptly curved inwards. In those species which have two upper premolars the first is extremely small and internal to the tooth-row. The genus, which is divided intoLasiurusproper andDasypterus, is further characterized by the presence of four teats in the female, and by the general productionof three or four offspring at a birth.RhogëessaandTomopeasare allied tropical American types.Murina, with the subgenusHarpiocephalus, hasi.2/3,p.2/2, and includes several small bats distinguished by the prominent tube-like nostrils and hairy interfemoral membrane.M. suilla, from Java, the Malay and neighbouring islands, is a well-known species, and the closely alliedM. hilgendorfiis from Japan. The remaining species are from the Himalaya, Tibet and Ceylon; and apparently restricted to the hill-tracts of the countries in which they are found. Next toVespertiliothe genusMyotis(divisible into several subgenera), withi.2/3,p.3/3, includes the largest number of species, and has rather a wider geographical distribution in both hemispheres, one species being recorded from the Navigator Islands. The species may be recognized by the peculiar character of the pairs of upper incisors on each side, the cusps of which diverge from each other, by the large number of premolars, of which the second upper is always small, and by the oval elongated ear and narrow tragus. The BritishM. bechsteiniandM. nattereriare examples of this group.Cerivoula(Kerivoula), which also hasp.3/3, is distinguished by the parallel upper incisors and the large second upper premolar. There are numerous African and Indo-Malayan species, of whichC. picta, from India and Indo-Malay, is characterized by its brilliant orange fur, and membranes variegated with orange and black. The genus includes delicately formed insectivorous, tropical, forest-haunting bats, whose colouring approximates them to the ripe bananas among which they often pass the daytime.Another subgroup,Minioptereae, is represented solely by the genusMiniopterus, withi.2/3,p.2/3. The incisors are separated from one another in front and from the canines; the first phalange of the middle finger is very short, the crown of the head elevated, and the tail long. The genus is represented by some half-dozen Old World species, among which the typicalM. schreibersiranges from Europe, southern Asia, and Africa to Japan and Australasia.Fig. 21.—Head of Mastiff-bat (Molossus glaucinus). (From Dobson.)Fig. 22.—Head ofNyctinomops macrotis. (From Dobson.)The last subfamily is that of theMolossinae, included by Dobson in the familyEmballonuridae. In this group the premaxillae are in contact or but very slightly separated; the ears are large, with the tragus small; the dental formula isi.1/1 (1/2 or 1/3),c.1/1,p.1/2 (2/2),m.3/3; and the fibula is strongly developed. In their blunt muzzles and many other features these bats undoubtedly resemble theEmballonuridae, from the typical members of which they differ by the production of the thick tail far beyond the margin of the interfemoral membrane. They are further characterized by their broad and stout feet, in which the first, and in most cases also the fifth, toe is thicker than the rest, and furnished with long bent hairs; and by the presence of callosities at the base of the thumbs, and a single pair of large upper incisors occupying the centre of the space between the canines. The feet are free from the wing-membrane, which folds up under the fore-arm and legs; the interfemoral membrane is retractile, being movable backwards and forwards along the tail; this power of varying its superficial extent confers on these bats great dexterity in changing the direction of flight. All are able to walk or crawl well, and spend much of their time on trees. The genusChiromeles, withi.1/1,c.1/1,p.1/2,m.3/3, the first hind-toe much larger than and separate from the others, and the widely sundered ears, is represented byC. torquata, a large bat of peculiar aspect, inhabiting the Indo-Malay countries. This species is nearly naked, a collar only of thinly spread hairs half surrounding the neck, and is remarkable for its enormous throat-sac and nursing-pouches. The former consists of a semicircular fold of skin forming a pouch round the neck beneath, concealing the orifices of subcutaneous pectoral glands which discharge an oily fluid of offensive smell. The nursing-pouch is formed on each side by an extension of a fold of skin from the side of the body to the inferior surfaces of the humerus and femur. In the anterior part of this pouch the teat is placed. The typical genusMolossus(fig. 21) includes the mastiff-bats, characterized by the dental formulai.1/1 or 1/2,p.1/2 or 2/2; and by the upper incisors being close together in front. The genus is restricted to the tropical and subtropical regions of the New World.M. obscurus, a small species common in tropical America, inhabits the hollow trunks of palms and other trees and the roofs of houses. The males and females live apart (as is the case in most if not all bats). In West Africa the mastiff-bats are represented byEomops, with one species; whileNyctinomopsincludes a number of tropical American species more nearly related to the next genus, in which some of them (fig. 22) were formerly included. The widely spreadNyctinomus, withi.1/3 or 1/2,p.2/2 or 1/2, and the upper incisors separate in front, includes numerous species inhabiting the tropical and subtropical parts of both hemispheres. The lips of the bats of this genus are even more expansible than inMolossus, in many of the species (fig. 22) showing vertical wrinkles.N. toeniotis(orcestonii), one of the largest species, alone extends into Europe, as far north as Switzerland.N. johorensis, from the Malay Peninsula, is remarkable for the extraordinary form of its ears.N. brasiliensisis common in tropical America, and extends as far north as California.Fig. 23.—Thumb and leg and foot of New Zealand bat (Mystacops tuberculatus), enlarged. (From Dobson.)Here may be conveniently noticed two very rare and aberrant bats,Myzopoda(orMyxopoda)auritaof Madagascar, andMystacops(orMystacina)tuberculatasof New Zealand, the latter of which is believed to be well-nigh, if not entirely, exterminated.Myzopoda and Mystacops.Their systematic position and affinities are somewhat uncertain; but in the opinion of O. Thomas2the former should typify a separate family,Myzopodidae, in which the latter may also find a place. From all other batsMyzopodais distinguished by the presence of a peculiar mushroom-shaped organ at the base of the large ear, and by the union of the tragus with the latter, on the inner base of which it forms a small projection. There are three phalanges in the middle finger; and the whole inferior surface of the thumb supports a large sessile horseshoe-shaped adhesive pad, with the circular margin directed forwards and notched along its edge, while a smaller pad occupies part of the sole of the hind-foot. Mr Thomas regards this bat as related on the one hand to the subfamilyMormopsinaeof thePhyllostomatidae, and on the other to theNatalinaeamong theVespertilionidae; both these groups being regarded by him as of family rank.MystacopsresemblesMyzopodain having three phalanges to the middle finger, but differs in that the tail perforates the interfemoral membrane to appear on its upper surface in the manner characteristic of theEmballonuridae. The greater part of the wing-membrane is exceedingly thin, but a narrow portion along the fore-arm, the sides of the body, and the legs, is thick and leathery, and beneath this thickened portion the wings are folded. Other peculiarities of structure are found in the form of the claws of the thumbs and toes, each of which has a small heel projecting from its concave surface near the base, also in the sole of the foot and inferior surface of the leg, as shown in fig. 23. The plantar surface, including the toes, is covered with soft and very lax, deeply wrinkled skin, and each toe is marked by a central longitudinal groove with short grooves at right angles to it. The lax wrinkled integument is continued along the inferior flattened surface of the ankle and leg. These peculiarities appear to be related to climbing habits in the species.
Megachiroptera.
The first of these comprises the fruit-eating species, which are generally of large size, with the crowns of the cheek-teeth smooth and marked with a longitudinal groove. The bony palate is continued behind the last molar, narrowing slowlyFruit-eating bats.backwards; there are three phalanges in the index finger, the third phalange being terminated generally by a claw; the sides of the ear form a ring at the base; the tail, when present, is inferior to (not contained in) the interfemoral membrane; the pyloric extremity of the stomach is generally much elongated; and the spigelian lobe of the liver is ill-defined or absent, while the caudate is well developed. This group is limited to the tropical and sub-tropical parts of the Eastern Hemisphere.
All the members of this suborder are included in the single familyPteropodidae, the first representatives of which are the African epauletted bats, forming the genusEpomophorus. In this the dental formula isi.2/2 (or ½),c.1/1,p.2/3,m.½. Tail short or absent, when present free from the interfemoral membrane; second finger with a claw; premaxillae united in front. The species are strictly limited to Africa south of the Sahara, and are distinguished by the large and long head, expansible and often folded lips, and the white tufts of hair on the margins of the ears. The males are provided with glandular pouches, situated in the skin of the side of the neck near the point of the shoulder, which are rudimentary or absent in females. In the males they are lined with glandular membrane, from which long coarse yellowish hairs project to form conspicuous epaulet-like tufts on the shoulders. The males often have a pair of air-sacs extending outwards on each side from the pharynx beneath the integument of the neck, in the position shown in fig. 2. These bats appear to live principally on figs, the juicy contents of which their voluminous lips and capacious mouths enable them to swallow without loss. The huge and ugly West African hammer-headed bat,Hypsignathus monstrosus, represents an allied genus distinguished by the absence of shoulder-pouches, and the presence of leaf-like expansions of skin on the front of the muzzle, and of distinct cusps on the outer sides of the cheek-teeth. The great majority of the bats of this group, commonly known as “flying-foxes,” are included in the typical genusPteropus, of which the dental formula isi.2/2,c.1/1,p.3/3,m.2/3. All are of large size, and the absence of a tail, the long pointed muzzle, and the woolly fur covering the neck render their recognition easy. One of the species,P. edulis, inhabiting Java, measures 5 ft. across the fully extended wings, and is the largest member of the order.
The range of the genus extends from Madagascar through the Seychelles to India, Ceylon, Burma, the Malay Archipelago, Japan, New Guinea, Australia and Polynesia. Although two species inhabit the Comoro Islands, scarcely 200 m. from the mainland, not one is found in Africa; while the common Indian species is closely allied to the Madagascar flying-fox. The Malay Archipelago and Australia form the headquarters of these bats, which in some places occur in countless multitudes. The colonies exhale a strong musky odour, and when awake the occupants utter a loud incessant chatter. Wallace’s fruit-bat of Celebes and Macassar has been made the type of a separate genus, asStyloctenium wallacei. InRoussettus(orCynonycteris) the dentition is as inPteropus, but the tail is short, and the fur of the nape of the neck not different from that of the back: its distribution accords with that ofPteropus, except that it includes Africa and does not reach farther east than New Ireland.R. aegyptiacusinhabits the chambers of the Great Pyramid and other deserted buildings in Egypt, and is probably the species figured in Egyptian frescoes.Boneia, with two species, from Celebes, differs in having only two upper incisors.HarpyionycterisandScotonycteris, respectively from the Philippines and West Africa, are represented by a single species each; but ofCynopterus, which is mainly confined to the Indo-Malay countries, there are some half-score different kinds. The dentition isi.2/(2 or 1),c.1/1,p.3/3,m.3/3, the muzzle is shorter than inRoussettus, with the upper lip grooved in front as inPteropus, while the tail and fur resemble those of the former genus. These bats are extremely voracious, a specimen of the IndianC. marginatushaving eaten a banana twice its own weight in three hours. Among several Austro-Malay genera, such asPtenochirusandBalionycteris, the tube-nosed bats of the genusGelasinus(orHarpyia) are remarkable for the conformation of the nostrils (fig. 5).Cephalotes, with one species, ranging from Celebes to the Solomon group, has the dentitioni.1/1,c.1/1,p.2/3,m.2/3, premaxillae not united in front, nostrils simple, muzzle short, index finger without a claw, tail short. As inGelasinus, the wing-membrane arises from the middle line of the back, to which it is attached by a longitudinal thin process of skin; the wings are naked, but the back covered with hair.Leipenyxis an allied West African genus with one species.
The foregoing belong to the typical subfamilyPteropodinae, while the remainder represent a second group,Carponycterinae(orMacroglossinae), characterized by having the facial part of the skull produced, the molar teeth narrow, and scarcely raised above the gum, and the tongue exceedingly long, attenuated in the anterior third, and armed with long recurved papillae near the tip. The single representative of the first genus,Notopteris macdonaldi, inhabiting Fiji, New Guinea and the New Hebrides, is distinguished from other bats of this family by the length of its tail, which is nearly as longas the forearm. The dentition isi.2/1,c.1/1,p.2/3,m.2/2, while the index finger has no claw, and the wings arise from the spine.Eonycteris, with the dentitioni.2/2,c.1/1,p.3/3,m.2/3, is also represented by a single species,E. spelaea, from Tenasserim, Burma, and the Malay Peninsula and Islands, which has somewhat the appearance of aRoussettus, but the absence of a claw in the index finger and the presence of the characteristic tongue and teeth at once distinguish it.Carponycteris(Macroglossus) andMelonycteris, the former with several and the latter with a single species, are closely allied Indo-Malay and Papuan genera, the index finger in both having a claw, but the number of the teeth being the same as inEonycteris.C. minimusis the smallest known species of the suborder, much smaller than the serotine bat of Europe, with the fore-arm scarcely longer than that of the long-eared bat. It is nearly as common in certain parts of Burma asCynopterus marginatus, and extends eastwards through the Malay Archipelago as far as New Ireland, where it is associated withMelonycteris melanops, distinguished by its larger size and the total absence of the tail. An allied smallCarpopycterisinhabits India.Trygenycteris(Megaloglossus)woermanni, of West Africa, is the only member of the group occurring west of the Himalaya.Callinycterisof Celebes, with the dentitioni.2/2,c.1/1,p.2/2,m.3/3, has a short tail and no index-claws, whileNesonycterisof the Solomons, with the dentitioni.2/1,c.1/1,p.3/3,m.3/3, differs by the absence of the tail.
Microchiroptera.
The second and larger suborder, the Microchiroptera, includes all the insectivorous species, the majority of which are of relatively small size as compared with the Megachiroptera. In these bats, with a few specialized exceptions, the crowns of theInsect-eating bats.cheek-teeth are surmounted by sharp cusps, divided by transverse grooves. In the skull the bony palate narrows abruptly and is not continued backwards laterally behind the last molar; there is one rudimentary phalange (rarely two or none) in the index finger, which is never terminated by a claw; the outer and inner sides of the ear commence interiorly from separate points of origin; the tail, when present, is contained in the interfemoral membrane, or appears on its upper surface; the stomach, except in the blood-sucking group, is simple; and the spigelian lobe of the liver large, and the caudate generally small.
The bats included in this suborder are so numerous in genera (to say nothing of species) that only some of the more important types can bementioned.
Brief references have already been made to the manner in which in many or most of these bats the tail aids in the capture of prey. From the observations of C. Oldham, it appears that these bats, when walking, carry the tail downwards and forwards, so that the membrane connecting this organ with the hind-legs forms a kind of pouch or bag. If a large insect be encountered the bat seizes it with a snatch, and slightly spreading its folded wings and pressing them on the ground in order to steady itself, brings its feet forwards so as to increase the capacity of the tail-pouch, into which, by bending its neck and thrusting its head beneath the body, it pushes the insect. Although the latter, especially if large, will often struggle violently, when once in the pouch it but rarely escapes, from which it is subsequently extracted and devoured. It is assumed that the same method of capture is employed when on the wing; and a naturalist who has observed the long-eared bat picking moths off willows states that the bat always hovers when taking off the moth, and bends up the tail so as to form a receptacle for the insect as it drops.
In theRhinolophidae, Horse-shoe and Leaf-nosed bats of the Old World, the nose-leaf is developed and surrounds the nasal apertures, which are situated in a depression on the upper surface of the muzzle so as to look upwards; the ears are large and generally separate, without trace of a tragus or earlet; the premaxillae are rudimentary, suspended from the nasal cartilages, and support a single pair of small incisors; the molars have acuteW-shaped cusps; the skull is large, and the nasal bones which support the nose-leaf much expanded vertically and laterally. In females a pair of teat-like appendages are found in front of the pubis; and the long tail extends to the margin of the interfemoral membrane. The middle finger has two phalanges, but the index is rudimentary. The fibula is rudimentary.
TheRhinolophidaeare the most highly organized of insectivorous bats, in which the osseous and cutaneous systems reach the fullest development. Compared with theirs, the bones of the extremities and the wings of other bats appear coarsely formed, and their teeth seem less perfectly fitted to crush the hard bodies of insects. The complicated nasal appendages reach their highest development, and the differences in their form afford characters in the discrimination of the species, which resemble one another closely in dentition and the colour of the fur.
In the first subfamily,Rhinolophinae, the first toe has two, and the other toes three phalanges each; and the ilio-pectineal spine is not connected by bone with the antero-inferior surface of the ilium. In the horseshoe bats,Rhinolophus, the dentition is i. 1/2, c. 1/1, p. 2/3, m. 3/8, the nose-leaf has a central process behind and between the nasal orifices, with the posterior extremity lanceolate, and the antitragus large. Among the numerous formsR. luctusis the largest, and inhabits elevated hill-tracts in India and Malaysia;R. hipposiderusof Europe, extending into south England and Ireland, is one of the smallest; andR. ferrum-equinumrepresents the average size of the species, which are mainly distinguished from one another by the form of the nose-leaf. The last-named species extends from England to Japan, and southward to the Cape of Good Hope, but is represented by a number of local races. When sleeping, the horseshoe bats, at least in some instances, suspend themselves head downwards, with the wings wrapped round the body after the manner of fruit bats. The posture of ordinary bats is quite different, and while the lesser horseshoe (R. hipposiderus) alights from the air in an inverted position, other bats, on first coming to rest, do so with the head upwards, and then reverse their position.
In the second subfamily,Hippo-siderinae(formerly calledPhyllorhinae), the toes are equal and include two phalanges each, while the ilio-pectineal spine is united by a bony isthmus with a process derived from the antero-inferior surface of the ilium.Hipposiderus, Clöeotis, Rhinonycteris, Triaenops, AnthopsandCoelopsrepresent this subfamily.Hipposiderus(Phyllorhina), with many species, ranging over Asia, Africa and Australasia, and the dental formula i. 1/2, c. 1/1, p. 2/2, or 1/2, m. 3/3, differs fromRhinolophusin the form of the nose-leaf, which is not lanceolate behind (fig. 6), and is unprovided with a central process covering the nostrils; the largest species,H. armiger, appears to be the most northerly, having been taken at Amoy in China, and in the Himalaya at an elevation of 5500 ft. Many are provided with a frontal sac behind the nose-leaf, rudimentary in females (see fig. 7), which can be everted at pleasure; the sides of this sac secrete a waxy substance, and its extremity supports a tuft of straight hairs.Rhinonycteris, represented byR. aurantiafrom Australia, andTriaenops. byT. persicusfrom Persia and other species from Africa and Madagascar, are closely allied genera.Triaenops(fig. 8) is characterized by the remarkable form of its nasal appendages and ears, and the presence of a bony projection from the upper extremity of the second phalange of the fourth finger.Coelops(C. Frithi), from the Bengal Sanderbans, Java and Siam is distinguished by the peculiar form of its nose-leaf and the length of the metacarpal bone of the index finger, as well as by the shortness of the calcar and interfemoral membrane.Clöeotisis represented by a single East African species, andAnthopsby one from the Solomon Islands characterized by the nose-leaf covering the whole front of the face.
The next family,Nycteridae, which is also Old World, is a small one, nearly allied to the last, in which it is included by Prof. Max Weber as a subfamily under the name ofMyadermatinae. It differs by the presence of a small tragus inFalse vampires.the ears, which are united at their bases; and by the nasal chamber not being inflated. The premaxillae are either small and separated in front, or rudimentary; and the first phalange of the middle finger when in repose is laid back on the metacarpus. There are only pectoral teats.
Of the two genera,Megaderma, as represented by the five species of false vampires, is distinguished by the absence of ossified premaxillae and upper incisors (i. 0/2, p. (2 or 1)/2), the cylindrical narrow muzzle surmounted by an erect nose-leaf the base of which conceals the nasal orifices, the immense joined ears with large bifid tragus, and the great extent of the interfemoral membrane, in the base of which the short tail is concealed.M. gigas(fig. 9), from central Queensland, is the largest species of the genus, and of the suborder.M. lyra, common in India (fore-arm 2.7 in.), has been caught in the act of sucking the blood, while flying, from a small bat which it afterwards devoured. The range of the genus includes Africa, the Indo-Malay countries and Australasia.Nycteris, which is commonto Africa and the Malay Peninsula and Islands, has ossified premaxillae and upper incisors (i.2/3,p.1/2), and a long tail; but lacks a nose-leaf. As inMegaderma, the frontal bones are deeply hollowed and expanded laterally, the muzzle presents a similar cylindrical form, and the lower jaw also projects; but, instead of a nose-leaf, the face is marked by a deep longitudinal sharp-edged groove extending from the nostrils to the band connecting the base of the large ears; the sides of this depression being margined as far back as the eyes by small horizontal cutaneous appendages. With the exception ofN. javanica, the species are limited to Africa.
According to the classification followed by Dr G.E. Dobson, the extensive family of New World bats known asPhyllostomatidaewas widely sundered from the two preceding groups; but in Prof. Max Weber’s system they are placed next oneVampires.another—an arrangement which has the great advantage of bringing together all the bats furnished with nose-leaves. It is indeed probable that the vampires, as the members of the present family may be collectively termed, are the New World representatives of the Old WorldRhinolophidaeandNycteridae.
ThePhyllostomatidaeare characterized by the presence of a nose-leaf, or of lappets on the chin, but the nostrils are not directed upwards. The ethmoturbinal bones of the nasal cavity form simple plates (much as in the two preceding families). The premaxillae are always well developed, with their palatal portions forming a suture and denning the boundaries of distinct palatine foramina (in place of being rudimentary, as inNycteridaeandRhinolophidae). The large ears have a tragus. The middle finger has three phalanges, and the index one. There is an incomplete fibula. The tail may be either long or short. Generally the dentition isi.2/2,c.1/1,p.2/3,m.3/3.
All the bats of this family may be readily recognized by the presence of a well-developed third phalange in the middle finger, associated either with a distinct nose-leaf, or with central upper incisors, or with both. Unlike theRhinolophidae, their eyes are generally large and the tragus is well developed, maintaining almost the same form throughout the species, however much the other parts of the body may vary. Their fur is of a dull colour, and the face and back are often marked with white streaks. A few species, probably all those with the tail and interfemoral membrane well developed, feed principally on insects, while the greater number of the species of the groupsVampyreaeandGlossophageaeappear to live on a mixed diet of insects and fruits, and theDesmodonteae, of which two species are known, are true blood-suckers, and have their teeth and intestinal tract specially modified in accordance with their habits. The group is practically limited to the tropical and subtropical parts of Central and South America, although one species ofOtopterusreaches California. In the first subfamily,Mormopsinae(Lobostominae), the nostrils open by simple apertures at the extremity of the muzzle in front, not margined by a distinct nose-leaf; while, in compensation, the chin is furnished with expanded leaf-like appendages. The tail is short. It includes two genera. InChilonycteristhe crown of the head is moderately elevated above the face-line, and the basi-cranial axis is almost in the same plane as the facial, while inMormops(fig. 10) the crown of the head is greatly elevated above the face-line, and the basi-cranial axis is nearly at right angles to the facial;i.2/2,p.2/3, in both genera. As regards the species ofChilonycteris, the most striking feature is the occurrence of a rufous and a dark brown phase in each. In some the two phases are very marked, but in others they are connected by intermediate shades. Here may be mentioned the two species of tropical American hare-lipped bats, forming the genusNoctilio, which presents characters common to this and the following family, to which latter it is often referred. The typicalN. leporinusis a bat of curious aspect, with strangely folded lips, erect skin-processes on the chin, and enormous feet and claws. The two middle incisors are close together, and so large as to conceal the small outer ones, while in the lower jaw there are but two small incisors; the premolars numbering 1/2. These bats live near the coast, and feed on small crabs and fishes.
Most of the remaining members of the family are included in the subfamilyPhyllostomatinae, characterized by the presence of a distinct nose-leaf and the warty chin. The clitoris is imperforate, whereas it is perforated in theMormopsinae. The incisors are generally 2/2 (occasionally 2/1), and the molars well developed. The subfamily is divided into a number of groups or sections. The first of them, theVampyreae, is characterized as follows: Muzzle long and narrow in front, the distance between the eyes generally less than (rarely equal to) that from the eye to the extremity of the muzzle; nose-leaf horseshoe-shaped in front, lanceolate behind; interfemoral membrane well developed; tail generally distinct, rarely absent; inner margin of the lips not fringed;i.2/2 or 2/1,p.2/2 or 2/3; molars withW-shaped cusps, usually well developed.
Nearly all theVampyreaeappear to be insectivorous, so that the term cannot be considered indicative of habits; but a few, if not all, probably supplement their insect diet with fruit.Vampyrus spectrum(the largest bat in the New World) is said to be wholly frugivorous, andOtopterus waterhouseiappears to prey occasionally on smaller bats. The genera may be arranged in two subgroups according as the tail is produced to the margin of the interfemoral membrane or perforates it to appear on its upper surface. In the first division are included three genera,Lonchorhina,Otopterus(orMacrotus) andDolichophyllum(orMacrophyllum), the first represented byL. aurita, characterized by an extraordinary long nose-leaf, and peculiarly large ears and tragus. In the second subsection are includedVampyrus,Chrotopterus,Tonatia(Lophostoma)Micronycteris,Glyphonycteris,Trachyops,Phylloderma,Phyllostoma,Anthorhina(Tylostoma),Mimon,Hemiderma(Carollia) andRhinophylla; all, with the exception of the last, distinguished chiefly by the form of the skull and the presence or absence of the second lower premolar.Phyllostoma hastatum, next in point of size toVampyrus spectrum, is a well-known species in South America;P. elongatum(fig. 11) differs in its smaller size and larger nose-leaf.Hemiderma brevicauda, a small species, closely resemblesGlossophaga soricina, and forms a connecting link between this and the next group.Rhinophylla pumiliois the smallest species of the family; further distinguished by the absence of a tail, the narrowness of its molars, which do not formW-shaped cusps, and the small size of the last upper molar, characters connecting it and the group with theStenodermateae. Both inHemidermaandRhinophyllathe zygomatic arch is incomplete.
The next subsection,Glossophageae, presents the following distinctive features: Muzzle long and narrow; tongue long and extensible, attenuated towards the tip, and beset with long filiform recurved papillae; lower lip with a wide groove above, and in front margined by small warts; nose-leaf small; tail short or none;i.2/2,p.2/3 or 3/3 or 2/2,m.3/3 or 2/3 or 2/2; teeth narrow; molars with narrowW-shaped cusps, sometimes indistinct or absent; lower incisors small or deciduous. The species included in this group represent some ten genera, distinguished principally by differences in the form and number of the teeth, and the presence or absence of the zygomatic, arch of the skull. InGlossophagaandPhyllonycteristhe upper incisors form a continuous row between the canines. InMonophyllusandLeptonycteris(Ischnoglossa) they are separated into pairs by a narrow interval in front; while inLonchoglossa,GlossonycterisandChoeronycteristhey are widely separated and placed in pairs near the canines. In the first four of these genera the lower incisors are present (at least to a certain age), in the last three they are deciduous even in youth. The zygomatic arch is wanting inPhyllonycteris,GlossonycterisandChoeronycteris. The typical species isGlossophaga soricina, which, as already mentioned, closely resemblesHemidermabrevicauda, both in form and dentition. Its long brush-tipped tongue (which it possesses in common with other species of the group) is used to lick out the pulpy contents of fruits having hard rinds. The food of the species of this group appears to consist of both fruit and insects, and the long tongue may be used for extracting the latter from the deep corollas of flowers. Other genera areLonchophylla,Rhithronycteris,HylonycterisandLychonycteris, each with a single species (in 1904).
The third group,Stenodermateae, presents the following characteristics:—Muzzle very short and generally broad in front, the distance between the eyes nearly always exceeding (rarely equalling) the distance from the eye to the extremity of the muzzle; nose-leaf short, horseshoe-shaped in front, lanceolate behind (except inBrachyphyllaandCenturio); interfemoral membrane concave behind; tail none; inner margin of the lips fringed with conical papillae;i.2/2 or 2/1,p.2/2,m.3/3 or 2/3 or 2/2; cheek-teeth broad (except inSturnira), molars with concave or flat crowns margined externally by raised cutting-edges. Although theStenodermateaeare generally easily distinguished from theVampyreaeby the shortness and breadth of the muzzle and the form of the cheek-teeth, certain species of the latter resemble the former in external appearance, agreeing almost absolutely in the form of the nose-leaf, the ears and the tragus, and the warts on the chin. These resemblances show that, while the form of the teeth and jaws has become modified to suit the food, the external characters have remained much the same, and indicate the common origin of the two sections. The food of these bats appears to be wholly or in great part fruit. The species are divided into some eleven genera, mostly distinguished by the form of the skull and teeth.Artibeusincludes the frugivorousA. perspicillatus.Stenoderma achradophilum, found in Jamaica and Cuba, with the last, from which it is scarcely distinguishable externally except by its much smaller size, differs in the absence of the horizontal plate of the premaxillae on the palate.Sturnira lilium, while agreeing with these in the form of the nose-leaf and ears, differs from all the species of the family in its longitudinally-grooved molars, which resemble those of thePteropodidaemore closely than those of any other bats; and the presence of tufts of long differently-coloured hairs over glands in the sides of the neck is another character in common with that group.Centurio senex(fig. 13) is the type of a small genus distinguished fromStenodermaand other genera of this group by the absence of a distinct nose-leaf. Some naturalists make this genus the type of a distinct subgroup,Centurioneae. Up to 1904 the genera, exclusive ofCenturio, included in theStenodermateaewereArtibeus(with several sub-genera),Vampyrops(also with subgenera),Mesophylla,Chiroderma,Stenoderma(with 3 subgenera),Ectophylla,Ametrida(with 2 sub-genera),Pygoderma,SturniraandBrachyphylla.
The third subfamily,Desmodontieae, is represented only by the blood-sucking bats, and distinguished by havingi.½, of which the upper pair are cutting, the rudimentary molars, the very short interfemoral membrane, and the blood-sucking habit. They are further characterized as follows: Muzzle short and conical; nose-leaf distinct;p.2/3,m.1/1 or 0/0; upper incisors occupying the whole space between the canines; premolars narrow, with sharp-edged longitudinal crowns; molars rudimentary or absent; stomach elongated, and intestiniform. There are two genera,Desmodus, without calcar or molars, andDiphylla, with a short calcar and a single rudimentary molar on each side—restricted to Central and South America.Desmodus rufus, the commoner species, is a little larger than the noctule bat, and abundant in certain parts of South America, where it is troublesome owing to its attacks upon domestic animals, sucking their blood and leaving them weakened from repeated bleedings. (SeeVampire.)
The fourth family of bats, unlike any of the three previous ones, has a cosmopolitan distribution. These free-tailed bats, as they are conveniently called, constituting the familyEmballonuridae, present the following distinctive features. TheFree-tailed bats.nostrils are of normal form and without a nose-leaf. The premaxillae have their palatal portion imperfectly developed, and united by a slender process with the maxillae. The ears are large, with a small tragus. The middle finger has two phalanges, and the index generally a single one. The fibula is incomplete. The tail is generally short, and always partly free from the interfemoral membrane. There is generally only a single pair of upper incisors, separated by gaps from the canines, and from one another in the middle line.
The distinctive feature of these bats is the free tail-tip, which pierces the interfemoral membrane to appear on its upper surface, and may project beyond its margin. As a rule, these bats may also be recognized by the peculiar form of the muzzle, which is obliquely truncated, the nostrils projecting more or less in front beyond the lower lip, by the first phalange of the middle finger being folded in repose forwards on the upper surface of the metacarpal bone, and by the upper incisors. Although cosmopolitan, these bats rarely extend north or south of the thirtieth parallels of latitude.
The family may be divided into two subfamilies, of which theEmballonurinaeis characterized by the incomplete premaxillae, the presence of only one phalange in the index finger, and the short tail. The dental formula is generallyi.1/3 (sometimes 2/3 or 1/2),c.1/1,p.2/3,m.3/3. This subfamily may be further subdivided into subgroups or sections of which the first,Embalionurae, is characterized by the slender tail perforating the interfemoral membrane, so as to appear on its upper surface; the legs long, with a slender fibula; the incisors weak; and the premolars 2/2. The typical genusEmballonurapresents the following features:i.2/3, extremity of the muzzle more or less produced beyond the lower lip, forehead flat. The genus contains several species, inhabiting islands from Madagascar through the Malay Archipelago and Siam to the Navigator Islands.Coleura, withi.1/3, the extremity of the muzzle broad, and the forehead concave, has two species from East Africa and the Seychelles.Rhynchonycterisis distinguished fromColeuraby the produced extremity of the muzzle. The single species,R. naso, from Central and South America, is common in the vicinity of streams, where it is usually found during the day resting on the vertical faces of rocks, or on trunks of trees growing over water; it escapes notice owing to the greyish colour of the fur of the body and of small tufts on the antebrachial membrane counterfeiting the weathered surfaces of rocks and bark. As evening approaches it appears on the wing, flying close to the water.Saccopteryxhasi.1/3 and the antibrachial membrane with a pouch opening on its upper surface; it contains several species from Central and South America. This sac is developed only in the male and in the female is rudimentary. In adult males a valvular longitudinal opening occupies the upper surface of the membrane leading into a small pouch, the interior of which is lined with a glandular membrane secreting an unctuous reddish substance with a strong ammoniacal odour. Allied genera are the tropical AmericanPeropteryxand the BrazilianCormura. The various species of tomb-bats (Taphozous) inhabit the tropical and subtropical parts of all the eastern hemisphere except Polynesia, and are distinguished by the cartilaginous premaxillaries, the deciduous pair of upper incisors, and the presence of only two pairs of lower incisors. Most of the species have a glandular sac (fig. 15) between the angles of the lower jaw, more developed in males than in females, in some species absent in the latter. An open throat-sac is wanting inT. melanopogon, but about its position are the openings of small pores, the secretion from which probably causes the hairs to grow long, forming the black beard found in many males. The three tropical American white bats,Diclidurus, withi.1/3,c.1/1,p.3/2,m.3/3, resembleTaphozousin the form of the head and ears, but, besides other characters, differ from all other bats in possessing a pouch, opening off the centre of the interior surface of the interfemoral membrane; the extremity of the tail enters this, and perforates its base.
The second subfamily of theEmballonuridae,Rhinopomatinae, is represented only by the genusRhinopoma, with several species ranging from Egypt through Arabia to India, Burma and Sumatra. The premaxillae (fig. 16) are complete; the index finger has two phalanges; the tail is very long and mouselike; and the dental formulai.1/2,c.1/1,p.1/2,m.2/3. Dr G.E. Dobson has remarked that these mouse-tailed bats might be elevated to the rank of a family, for it is difficult to determine their affinities, a kind of cross relationship attaching them to theNycteridaeon the one hand and to theEmballonuridaeon the other. These bats, distinguished from all other Microchiroptera by the presence of two phalanges in the index fingerand the long and slender tail projecting far beyond the narrow interfemoral membrane, inhabit the subterranean tombs in Egypt and deserted buildings generally from north-east Africa to Burma and Sumatra.
The last group, according to the system adopted by Prof. Max Weber, is that of theVespertilionidae, which includes such typical bats as the pipistrelle, the noctule, and the long-eared species. By Mr G.S. Miller1the first section of theTypical bats.family—Natalinae—is regarded as of family rank, while the last section, orMolossinae, is included by Dr G.E. Dobson in theEmballonuridae, from the typical forms of which its members differ widely in tail-structure. In this extended sense the family, which has a cosmopolitan distribution, may be defined as follows:—The nostrils are normal and without a nose-leaf. The ethmoturbinal bones of the nasal chamber are involuted. The palatine processes of the premaxillae do not form a suture. The ear is mostly large, with a tragus. The middle finger (except inThyroptera) has two phalanges. The fibula is usually rudimentary. The tail is long and does not perforate the interfemoral membrane. The incisors are generally 2/3 or 1/2, but may be reduced to 1/1 in theMolossinae.
In the first subfamily,Natalinae, which is exclusively tropical American, the other upper incisors are separated from one another and from the canines; palatine processes of the premaxillae are at least partially developed; and the dental formula isi.2/3,c.1/1,p.(2 or 3)/3,m.3/3. In general appearance these bats recall the more typicalVespertilionidae, although the form of the muzzle is suggestive of theMormopsinaeamong thePhyllostomatidae. Again, while the form of the skull is vespertilione, the relation of the vomer to the front end of the premaxillae is of the phyllostomine type. The molars and incisors are likewise vespertilione, whereas the premolars are as distinctly phyllostomine. Finally, while the third, or middle, finger normally has two phalanges, as in typicalVespertilionidae, the second of these is elongated and inThyropteradivided into two, as inPhyllostomatidae.
The first two genera,FuripterusandAmorphochilus, each have a single species, the latter being distinguished from the former by the wide separation of the nostrils and the backward prolongation of the palate. In both the crown of the head is elevated, the thumb and first phalange of the middle finger are very short, and the premolars are 2/3. The same elevation of the crown characterizes the generaNatalusandChilonatalus(fig. 17), in which the premolars are 3/3: in general appearance these bats are very like the Old World vespertilionine genusCerivoula, except for the short triangular tragus. Lastly,Thyropteraincludes two species distinguished by an additional phalange in the middle finger and by accessory clinging-organs attached to the extremities. InThyroptera tricolor, i.2/3,p.3/3, from Brazil, these have the appearance of small, circular, stalked, hollow disks (fig. 18), resembling miniature sucking-cups of cuttle-fishes, and are attached to the inferior surfaces of the thumbs and the soles of the feet. By their aid the bat is able to maintain its hold when creeping over smooth vertical surfaces.
The second or typical subfamily,Vespertilioninae, includes all the remaining members of the family with the exception of the aberrantMolossinae. The upper incisors are in proximity to the canines; the premaxillae widely separated; the ears medium or large; the dental formula isi.2/3 (or 1/3),c.1/1,p.3/3 (2/3, 2/2, or 1/2),m.3/3; and the fibula very small and imperfect. All the members of this large cosmopolitan group are closely allied, and differ chiefly by external characters. They may be divided into subgroups. In the first of these, thePlecoteae, of which the long-eared bat (Plecotus auritus) is the type, the crown of the head is but slightly raised above the face-line, the upper incisors are close to the canines, and the nostrils are margined behind by grooves an the upper surface of the muzzle, or by rudimentary nose-leaves; the ears being generally very large and united. Of the six genera,Plecotus, withi.2/3,p.2/3, has three species:—one the long-eared European bat referred to above;P. macrotis, restricted to North America, is distinguished by the great size of the glandular prominences of the sides of the muzzle, which meet in the centre above and behind the nostrils; the third species being also American. The second,Barbastella, withi.2/3,p.2/2, distinguished by its dentition and by the outer margin of the ear being carried forwards above the mouth and in front of the eye, includes the European barbastelle bat,B. barbastellus, andB. darjelingensisfrom the Himalaya.Otonycteris,i.1/3,pm., 1/2, connecting this group with theVespertilioneae, is represented byO. hemprichii, from North Africa and the Himalaya, and an Arabian species. The next two genera are distinguished by the presence of a rudimentary nose-leaf:Nyctophilus,i.1/3,p.1/2, with three species from Australasia; andAntrozous,i.1/2,p.1/2, distinguished from all the other members of the subfamily by having but two lower incisors, and from otherPlecoteaeby the separate ears; the two species inhabit California. The sixth genus,Euderma, is also represented by a Californian species.
The second groupVespertilioneae, with about thirteen genera, includes the great majority of the species; and a large number of these may be classed underVespertilio, which is divisible into subgenera, differing from one another in the number of premolars, and often ranked as separate genera. One group is represented byV.(Histiolus)magellanicus, a species remarkable for its extreme southern range, its relatives being also South American. A second group, withp.1/2, includes the British serotine,V.(Eptesicus)serotinus, of Europe and northern Asia, and represented in North America by the closely alliedV.(E.)fuscus. In the typical group, which includes the Old WorldV. murinus, one species,V. borealis, ranges to the Arctic circle. The European noctule,V.(Pierygistes)noctula, and Leisler’s bat,V.(P.)leisleri, represent another group; and the common pipistrelle,V.(Pipistrellus)pipistrellus, yet another, withp.2/2. The only other group that need be mentioned is one represented by the North AmericanV.(Lasionycteris)noctivagans, withp.2/3. The AfricanLäephotes, the ChineseIa, and the PapuanPhiletorare allied genera, each with a single species.ChalinolobusandGlauconycterishave the same general dental character asVespertilio, but are distinguished by the presence of a lobe projecting from the lower lip near the gape; the former, withp.2/2, is represented by five Australasian species, one of which extends into New Zealand; while the latter, withp.1/2, is African. The species ofGlauconycterisare noticeable for their peculiarly thin membranes traversed by distinct reticulations and parallel lines.Scotophilus, withi.1/3,p.1/2, includes several species, restricted to the tropical and subtropical regions of the eastern hemisphere, though widely distributed within these limits. These bats, though approaching certain species ofVespertilioin many points, are distinguished by the single (in place of two) pair of unicuspidate upper incisors separated by a wide space and placed close to the canines, by the small transverse first lower premolar crushed in between the canine and second premolar, and, generally, by their conical, nearly naked, muzzles and thick leathery membranes.S. temminckiis the commonest bat in India, and appears often before the sun has touched the horizon.S. gigas, from equatorial Africa, is the largest species.Nycticejus, with the same dental formula asScotophilus, is distinguished, by the first lower premolar not being crushed in between the adjoining teeth, and the comparatively greater size of the last upper molar. It includes only the North AmericanN. humeralis(crepuscularis), a bat scarcely larger than the pipistrelle. The hairy-membraned bats of the genusLasiurus(Atalapha), withi.1/3,p.2/2 or 1/2, are also limited to the New World, and generally characterized by the interfemoral membrane being more or less covered with hair and by the peculiar form of the tragus, which is expanded above and abruptly curved inwards. In those species which have two upper premolars the first is extremely small and internal to the tooth-row. The genus, which is divided intoLasiurusproper andDasypterus, is further characterized by the presence of four teats in the female, and by the general productionof three or four offspring at a birth.RhogëessaandTomopeasare allied tropical American types.Murina, with the subgenusHarpiocephalus, hasi.2/3,p.2/2, and includes several small bats distinguished by the prominent tube-like nostrils and hairy interfemoral membrane.M. suilla, from Java, the Malay and neighbouring islands, is a well-known species, and the closely alliedM. hilgendorfiis from Japan. The remaining species are from the Himalaya, Tibet and Ceylon; and apparently restricted to the hill-tracts of the countries in which they are found. Next toVespertiliothe genusMyotis(divisible into several subgenera), withi.2/3,p.3/3, includes the largest number of species, and has rather a wider geographical distribution in both hemispheres, one species being recorded from the Navigator Islands. The species may be recognized by the peculiar character of the pairs of upper incisors on each side, the cusps of which diverge from each other, by the large number of premolars, of which the second upper is always small, and by the oval elongated ear and narrow tragus. The BritishM. bechsteiniandM. nattereriare examples of this group.Cerivoula(Kerivoula), which also hasp.3/3, is distinguished by the parallel upper incisors and the large second upper premolar. There are numerous African and Indo-Malayan species, of whichC. picta, from India and Indo-Malay, is characterized by its brilliant orange fur, and membranes variegated with orange and black. The genus includes delicately formed insectivorous, tropical, forest-haunting bats, whose colouring approximates them to the ripe bananas among which they often pass the daytime.
Another subgroup,Minioptereae, is represented solely by the genusMiniopterus, withi.2/3,p.2/3. The incisors are separated from one another in front and from the canines; the first phalange of the middle finger is very short, the crown of the head elevated, and the tail long. The genus is represented by some half-dozen Old World species, among which the typicalM. schreibersiranges from Europe, southern Asia, and Africa to Japan and Australasia.
The last subfamily is that of theMolossinae, included by Dobson in the familyEmballonuridae. In this group the premaxillae are in contact or but very slightly separated; the ears are large, with the tragus small; the dental formula isi.1/1 (1/2 or 1/3),c.1/1,p.1/2 (2/2),m.3/3; and the fibula is strongly developed. In their blunt muzzles and many other features these bats undoubtedly resemble theEmballonuridae, from the typical members of which they differ by the production of the thick tail far beyond the margin of the interfemoral membrane. They are further characterized by their broad and stout feet, in which the first, and in most cases also the fifth, toe is thicker than the rest, and furnished with long bent hairs; and by the presence of callosities at the base of the thumbs, and a single pair of large upper incisors occupying the centre of the space between the canines. The feet are free from the wing-membrane, which folds up under the fore-arm and legs; the interfemoral membrane is retractile, being movable backwards and forwards along the tail; this power of varying its superficial extent confers on these bats great dexterity in changing the direction of flight. All are able to walk or crawl well, and spend much of their time on trees. The genusChiromeles, withi.1/1,c.1/1,p.1/2,m.3/3, the first hind-toe much larger than and separate from the others, and the widely sundered ears, is represented byC. torquata, a large bat of peculiar aspect, inhabiting the Indo-Malay countries. This species is nearly naked, a collar only of thinly spread hairs half surrounding the neck, and is remarkable for its enormous throat-sac and nursing-pouches. The former consists of a semicircular fold of skin forming a pouch round the neck beneath, concealing the orifices of subcutaneous pectoral glands which discharge an oily fluid of offensive smell. The nursing-pouch is formed on each side by an extension of a fold of skin from the side of the body to the inferior surfaces of the humerus and femur. In the anterior part of this pouch the teat is placed. The typical genusMolossus(fig. 21) includes the mastiff-bats, characterized by the dental formulai.1/1 or 1/2,p.1/2 or 2/2; and by the upper incisors being close together in front. The genus is restricted to the tropical and subtropical regions of the New World.M. obscurus, a small species common in tropical America, inhabits the hollow trunks of palms and other trees and the roofs of houses. The males and females live apart (as is the case in most if not all bats). In West Africa the mastiff-bats are represented byEomops, with one species; whileNyctinomopsincludes a number of tropical American species more nearly related to the next genus, in which some of them (fig. 22) were formerly included. The widely spreadNyctinomus, withi.1/3 or 1/2,p.2/2 or 1/2, and the upper incisors separate in front, includes numerous species inhabiting the tropical and subtropical parts of both hemispheres. The lips of the bats of this genus are even more expansible than inMolossus, in many of the species (fig. 22) showing vertical wrinkles.N. toeniotis(orcestonii), one of the largest species, alone extends into Europe, as far north as Switzerland.N. johorensis, from the Malay Peninsula, is remarkable for the extraordinary form of its ears.N. brasiliensisis common in tropical America, and extends as far north as California.
Here may be conveniently noticed two very rare and aberrant bats,Myzopoda(orMyxopoda)auritaof Madagascar, andMystacops(orMystacina)tuberculatasof New Zealand, the latter of which is believed to be well-nigh, if not entirely, exterminated.Myzopoda and Mystacops.Their systematic position and affinities are somewhat uncertain; but in the opinion of O. Thomas2the former should typify a separate family,Myzopodidae, in which the latter may also find a place. From all other batsMyzopodais distinguished by the presence of a peculiar mushroom-shaped organ at the base of the large ear, and by the union of the tragus with the latter, on the inner base of which it forms a small projection. There are three phalanges in the middle finger; and the whole inferior surface of the thumb supports a large sessile horseshoe-shaped adhesive pad, with the circular margin directed forwards and notched along its edge, while a smaller pad occupies part of the sole of the hind-foot. Mr Thomas regards this bat as related on the one hand to the subfamilyMormopsinaeof thePhyllostomatidae, and on the other to theNatalinaeamong theVespertilionidae; both these groups being regarded by him as of family rank.
MystacopsresemblesMyzopodain having three phalanges to the middle finger, but differs in that the tail perforates the interfemoral membrane to appear on its upper surface in the manner characteristic of theEmballonuridae. The greater part of the wing-membrane is exceedingly thin, but a narrow portion along the fore-arm, the sides of the body, and the legs, is thick and leathery, and beneath this thickened portion the wings are folded. Other peculiarities of structure are found in the form of the claws of the thumbs and toes, each of which has a small heel projecting from its concave surface near the base, also in the sole of the foot and inferior surface of the leg, as shown in fig. 23. The plantar surface, including the toes, is covered with soft and very lax, deeply wrinkled skin, and each toe is marked by a central longitudinal groove with short grooves at right angles to it. The lax wrinkled integument is continued along the inferior flattened surface of the ankle and leg. These peculiarities appear to be related to climbing habits in the species.