[1] For the Chippewa Indians see OJIBWAY, 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 Mamore 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. [Greek: cheir], hand, and [Greek: manteia],
divination), the art of telling the character or fortune of persons by
studying the lines of the palms of the hands (see PALMISTRY).
CHIRON, or CHEIRON, 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
constellation _Sagittarius_ (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's _Realencyclopaedie_ and W.H. Roscher's
_Lexikon der Mythologie_; W. Mannhardt, _Wald- und Feldkulte_ (1904).
CHIROPODIST (an invented word from Gr. [Greek: cheir], hand, and [Greek:
pous], 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.
[Illustration: FIG. 1.--Skeleton and Wing-Membranes of the Noctule Bat
(_Pipistrellus noctula_).
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 never exceeds 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 some _Pteropodidae_ and a few _Nycteridae_ and
_Emballonuridae_; in _Pteropus leucopterus_ alone 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 of _Phyllostomatidae_.
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 the _Rhinolophidae_ the milk-teeth are absorbed
before birth. The permanent teeth exhibit great variety, sometimes even
in the same family, as in _Phyllostomatidae_, whilst in other families,
as _Rhinolophidae_, 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 the _Rhinolophidae_ alone they form a
symphysis. Only in the _Molossinae_ is 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 the _Rhinolophidae_, 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. In
_Rhinopoma_ and _Megaderma_ a 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).
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 the _Emballonuridae_, and also in the
_Molossinae_, 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.
[Illustration: FIG. 2.--Head and Neck of _Epomophorus 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.]
[Illustration: FIG. 3.--Frontal Sac and Nose-Leaf in Male and Female
Masked Bat (_Phyllorhina larvata_). From Dobson.]
Bats are divisible into two suborders, Megachiroptera and
Microchiroptera.
_Megachiroptera_.
Fruit-eating bats.
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 slowly 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 family
_Pteropodidae_, the first representatives of which are the African
epauletted bats, forming the genus _Epomophorus_. In this the dental
formula is _i._ 2/2 (or 1/2), _c._ 1/1, _p._ 2/3, _m._ 1/2. 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 genus
_Pteropus_, of which the dental formula is _i._ 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.
[Illustration: FIG. 4.--Head of a Flying-Fox or Fruit-Bat (_Pteropus
personatus_). From Gray.]
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, as
_Styloctenium wallacei_. In _Roussettus_ (or _Cynonycteris_) the
dentition is as in _Pteropus_, 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 of _Pteropus_, except that it includes
Africa and does not reach farther east than New Ireland. _R.
aegyptiacus_ inhabits 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. _Harpyionycteris_ and
_Scotonycteris_, respectively from the Philippines and West Africa,
are represented by a single species each; but of _Cynopterus_, which
is mainly confined to the Indo-Malay countries, there are some
half-score different kinds. The dentition is _i._ 2/[2 or 1], _c._
1/1, _p._ 3/3, _m._ 3/3, the muzzle is shorter than in _Roussettus_,
with the upper lip grooved in front as in _Pteropus_, while the tail
and fur resemble those of the former genus. These bats are extremely
voracious, a specimen of the Indian _C. marginatus_ having eaten a
banana twice its own weight in three hours. Among several Austro-Malay
genera, such as _Ptenochirus_ and _Balionycteris_, the tube-nosed bats
of the genus _Gelasinus_ (or _Harpyia_) are remarkable for the
conformation of the nostrils (fig. 5). _Cephalotes_, with one species,
ranging from Celebes to the Solomon group, has the dentition _i._ 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 in _Gelasinus_, 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.
_Leipenyx_ is an allied West African genus with one species.
[Illustration: FIG. 5.--Head of Papuan Tube-Nosed Bat (_Gelasinus
major_). From G.E. Dobson.]
The foregoing belong to the typical subfamily _Pteropodinae_, while
the remainder represent a second group, _Carponycterinae_ (or
_Macroglossinae_), 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 long
as the forearm. The dentition is _i._ 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 dentition _i._ 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 a _Roussettus_, 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_) and
_Melonycteris_, 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 in _Eonycteris_. _C. minimus_ is 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 as _Cynopterus marginatus_, and
extends eastwards through the Malay Archipelago as far as New Ireland,
where it is associated with _Melonycteris melanops_, distinguished by
its larger size and the total absence of the tail. An allied small
_Carpopycteris_ inhabits India. _Trygenycteris_ (_Megaloglossus_)
_woermanni_, of West Africa, is the only member of the group occurring
west of the Himalaya. _Callinycteris_ of Celebes, with the dentition
_i._ 2/2, _c._ 1/1, _p._ 2/2, _m._ 3/3, has a short tail and no
index-claws, while _Nesonycteris_ of the Solomons, with the dentition
_i._ 2/1, _c._ 1/1, _p._ 3/3, _m._ 3/3, differs by the absence of the
tail.
_Microchiroptera._
Insect-eating bats.
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 the 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 be
mentioned.
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 the _Rhinolophidae_, 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 acute W-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.
[Illustration: FIG. 6.--Head of Mitred Horseshoe Bat (_Rhinolophus
mitratus_). From Dobson.]
The _Rhinolophidae_ are 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 forms _R. luctus_ is the largest,
and inhabits elevated hill-tracts in India and Malaysia; _R.
hipposiderus_ of Europe, extending into south England and Ireland, is
one of the smallest; and _R. ferrum-equinum_ represents 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.
[Illustration: FIG. 7.--Head of Squirrel Leaf-Bat (_Phyllorhina
calcarata_). From Dobson.]
[Illustration: FIG. 8.--Head of Persian Leaf-Bat. (_Triaenops
persicus_). From Dobson.]
In the second subfamily, _Hippo-siderinae_ (formerly called
_Phyllorhinae_), 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, Cloeeotis, Rhinonycteris, Triaenops, Anthops_ and
_Coelops_ represent 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 from
_Rhinolophus_ in 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 by _R. aurantia_ from Australia, and _Triaenops_. by _T.
persicus_ from 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.
_Cloeeotis_ is represented by a single East African species, and
_Anthops_ by one from the Solomon Islands characterized by the
nose-leaf covering the whole front of the face.
False vampires.
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 of _Myadermatinae_. It differs by
the presence of a small tragus in 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 common
to 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 in _Megaderma_, 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 of _N. javanica_, the species are limited to Africa.
[Illustration: FIG. 9.--The False Vampire (_Megaderma gigas_). From
Dobson.]
Vampires.
According to the classification followed by Dr G.E. Dobson, the
extensive family of New World bats known as _Phyllostomatidae_ was
widely sundered from the two preceding groups; but in Prof. Max
Weber's system they are placed next one 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 World _Rhinolophidae_ and
_Nycteridae_.
The _Phyllostomatidae_ are 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 in _Nycteridae_ and _Rhinolophidae_).
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 is _i._ 2/2, _c._ 1/1,
_p._ 2/3, _m._ 3/3.
[Illustration: 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 the _Rhinolophidae_, 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 groups
_Vampyreae_ and _Glossophageae_ appear to live on a mixed diet of
insects and fruits, and the _Desmodonteae_, 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 of _Otopterus_ reaches
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. In
_Chilonycteris_ the 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 in _Mormops_ (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 of _Chilonycteris_, 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 genus
_Noctilio_, which presents characters common to this and the following
family, to which latter it is often referred. The typical _N.
leporinus_ is 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
subfamily _Phyllostomatinae_, characterized by the presence of a
distinct nose-leaf and the warty chin. The clitoris is imperforate,
whereas it is perforated in the _Mormopsinae_. 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, the _Vampyreae_, 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 with W-shaped cusps, usually well developed.
Nearly all the _Vampyreae_ appear 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,
and _Otopterus waterhousei_ appears 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_ (or _Macrotus_)
and _Dolichophyllum_ (or _Macrophyllum_), the first represented by _L.
aurita_, characterized by an extraordinary long nose-leaf, and
peculiarly large ears and tragus. In the second subsection are
included _Vampyrus_, _Chrotopterus_, _Tonatia_ (_Lophostoma_)
_Micronycteris_, _Glyphonycteris_, _Trachyops_, _Phylloderma_,
_Phyllostoma_, _Anthorhina_ (_Tylostoma_), _Mimon_, _Hemiderma_
(_Carollia_) and _Rhinophylla_; 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 to _Vampyrus 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
resembles _Glossophaga soricina_, and forms a connecting link between
this and the next group. _Rhinophylla pumilio_ is the smallest species
of the family; further distinguished by the absence of a tail, the
narrowness of its molars, which do not form W-shaped cusps, and the
small size of the last upper molar, characters connecting it and the
group with the _Stenodermateae_. Both in _Hemiderma_ and _Rhinophylla_
the zygomatic arch is incomplete.
[Illustration: 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 narrow W-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. In _Glossophaga_ and _Phyllonycteris_
the upper incisors form a continuous row between the canines. In
_Monophyllus_ and _Leptonycteris_ (_Ischnoglossa_) they are separated
into pairs by a narrow interval in front; while in _Lonchoglossa_,
_Glossonycteris_ and _Choeronycteris_ they 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 in _Phyllonycteris_, _Glossonycteris_ and _Choeronycteris_.
The typical species is _Glossophaga soricina_, which, as already
mentioned, closely resembles _Hemiderma brevicauda_, 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 are _Lonchophylla_, _Rhithronycteris_,
_Hylonycteris_ and _Lychonycteris_, each with a single species (in
1904).
[Illustration: 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 in
_Brachyphylla_ and _Centurio_); 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 in _Sturnira_), molars with concave or flat crowns margined
externally by raised cutting-edges. Although the _Stenodermateae_ are
generally easily distinguished from the _Vampyreae_ by 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. _Artibeus_ includes the frugivorous _A. 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 the _Pteropodidae_ more 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 from _Stenoderma_ and 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 of _Centurio_, included in the _Stenodermateae_
were _Artibeus_ (with several sub-genera), _Vampyrops_ (also with
subgenera), _Mesophylla_, _Chiroderma_, _Stenoderma_ (with 3
subgenera), _Ectophylla_, _Ametrida_ (with 2 sub-genera), _Pygoderma_,
_Sturnira_ and _Brachyphylla_.
[Illustration: 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 having _i._ 1/2, 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, and _Diphylla_, 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. (See
VAMPIRE.)
Free-tailed bats.
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 family _Emballonuridae_, present
the following distinctive features. The 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.
[Illustration: FIG. 14.--Ear of _Emballonura raffrayana_. From
Dobson.]
The family may be divided into two subfamilies, of which the
_Emballonurinae_ is characterized by the incomplete premaxillae, the
presence of only one phalange in the index finger, and the short tail.
The dental formula is generally _i._ 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
genus _Emballonura_ presents 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_, with _i._ 1/3, the extremity of the
muzzle broad, and the forehead concave, has two species from East
Africa and the Seychelles. _Rhynchonycteris_ is distinguished from
_Coleura_ by 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. _Saccopteryx_ has _i._ 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 American _Peropteryx_ and the Brazilian _Cormura_. 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 in
_T. 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_, with _i._ 1/3, _c._ 1/1, _p._ 3/2,
_m._ 3/3, resemble _Taphozous_ in 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.
[Illustration: FIG. 15.--Heads of Tomb-Bat (_Taphozous longimanus_),
showing relative development of throat-sacs in male and female. From
Dobson.]
The second subfamily of the _Emballonuridae_, _Rhinopomatinae_, is
represented only by the genus _Rhinopoma_, 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 formula
_i._ 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 the _Nycteridae_ on the one hand
and to the _Emballonuridae_ on the other. These bats, distinguished
from all other Microchiroptera by the presence of two phalanges in the
index finger and 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.
Typical bats.
[Illustration: FIG. 16.--Skull of Mouse-tailed Bat (_Rhinopoma
microphyllum_). X2. (From Dobson.)]
The last group, according to the system adopted by Prof. Max Weber, is
that of the _Vespertilionidae_, which includes such typical bats as
the pipistrelle, the noctule, and the long-eared species. By Mr G.S.
Miller[1] the first section of the family--_Natalinae_--is regarded as
of family rank, while the last section, or _Molossinae_, is included
by Dr G.E. Dobson in the _Emballonuridae_, 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 in
_Thyroptera_) 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 the
_Molossinae_.
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 is _i._ 2/3, _c._ 1/1,
_p._ {2 or 3}/3, _m._ 3/3. In general appearance these bats recall the
more typical _Vespertilionidae_, although the form of the muzzle is
suggestive of the _Mormopsinae_ among the _Phyllostomatidae_. 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 typical
_Vespertilionidae_, the second of these is elongated and in
_Thyroptera_ divided into two, as in _Phyllostomatidae_.
[Illustration: FIG. 17.--Head of _Chilonatalus micropus_. x2. (From
Dobson.)]
The first two genera, _Furipterus_ and _Amorphochilus_, 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 genera
_Natalus_ and _Chilonatalus_ (fig. 17), in which the premolars are
3/3: in general appearance these bats are very like the Old World
vespertilionine genus _Cerivoula_, except for the short triangular
tragus. Lastly, _Thyroptera_ includes two species distinguished by an
additional phalange in the middle finger and by accessory
clinging-organs attached to the extremities. In _Thyroptera 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.
[Illustration: FIG. 18.--Suctorial Disks in _Thyroptera tricolor_,
_a_, side, and _b_, concave surface, of thumb disk; _c_, foot with
disk, and calcar with projections (all much enlarged). (From Dobson.)]
The second or typical subfamily, _Vespertilioninae_, includes all the
remaining members of the family with the exception of the aberrant
_Molossinae_. The upper incisors are in proximity to the canines; the
premaxillae widely separated; the ears medium or large; the dental
formula is _i._ 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, the _Plecoteae_, 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_,
with _i._ 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_, with _i._ 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_, and _B. darjelingensis_ from the
Himalaya. _Otonycteris_, _i._ 1/3, _pm._, 1/2, connecting this group
with the _Vespertilioneae_, is represented by _O. 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; and _Antrozous_, _i._ 1/2, _p._ 1/2, distinguished from
all the other members of the subfamily by having but two lower
incisors, and from other _Plecoteae_ by the separate ears; the two
species inhabit California. The sixth genus, _Euderma_, is also
represented by a Californian species.
[Illustration: FIG. 19.--Head of _Scotophilus emarginatus_. (From
Dobson.)]
[Illustration: FIG. 20.--Head of _Cerivoula hardwickei_. (From
Dobson.)]
The second group _Vespertilioneae_, with about thirteen genera,
includes the great majority of the species; and a large number of
these may be classed under _Vespertilio_, which is divisible into
subgenera, differing from one another in the number of premolars, and
often ranked as separate genera. One group is represented by _V._
(_Histiolus_) _magellanicus_, a species remarkable for its extreme
southern range, its relatives being also South American. A second
group, with _p._ 1/2, includes the British serotine, _V._
(_Eptesicus_) _serotinus_, of Europe and northern Asia, and
represented in North America by the closely allied _V._ (_E._)
_fuscus_. In the typical group, which includes the Old World _V.
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, with
_p._ 2/2. The only other group that need be mentioned is one
represented by the North American _V._ (_Lasionycteris_)
_noctivagans_, with _p._ 2/3. The African _Laeephotes_, the Chinese
_Ia_, and the Papuan _Philetor_ are allied genera, each with a single
species. _Chalinolobus_ and _Glauconycteris_ have the same general
dental character as _Vespertilio_, but are distinguished by the
presence of a lobe projecting from the lower lip near the gape; the
former, with _p._ 2/2, is represented by five Australasian species,
one of which extends into New Zealand; while the latter, with _p._
1/2, is African. The species of _Glauconycteris_ are noticeable for
their peculiarly thin membranes traversed by distinct reticulations
and parallel lines. _Scotophilus_, with _i._ 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 of _Vespertilio_ in
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.
temmincki_ is 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 as
_Scotophilus_, 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 American _N.
humeralis_ (_crepuscularis_), a bat scarcely larger than the
pipistrelle. The hairy-membraned bats of the genus _Lasiurus_
(_Atalapha_), with _i._ 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 into
_Lasiurus_ proper and _Dasypterus_, is further characterized by the
presence of four teats in the female, and by the general production
of three or four offspring at a birth. _Rhogeessa_ and _Tomopeas_ are
allied tropical American types. _Murina_, with the subgenus
_Harpiocephalus_, has _i._ 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 allied
_M. hilgendorfi_ is 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 to
_Vespertilio_ the genus _Myotis_ (divisible into several subgenera),
with _i._ 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 British _M. bechsteini_
and _M. nattereri_ are examples of this group. _Cerivoula_
(_Kerivoula_), which also has _p._ 3/3, is distinguished by the
parallel upper incisors and the large second upper premolar. There are
numerous African and Indo-Malayan species, of which _C. 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 genus
_Miniopterus_, with _i._ 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 typical _M. schreibersi_ ranges from Europe,
southern Asia, and Africa to Japan and Australasia.
[Illustration: FIG. 21.--Head of Mastiff-bat (_Molossus glaucinus_).
(From Dobson.)]
[Illustration: FIG. 22.--Head of _Nyctinomops macrotis_. (From
Dobson.)]
The last subfamily is that of the _Molossinae_, included by Dobson in
the family _Emballonuridae_. In this group the premaxillae are in
contact or but very slightly separated; the ears are large, with the
tragus small; the dental formula is _i._ 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 the _Emballonuridae_, 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 genus
_Chiromeles_, with _i._ 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 by _C. 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 genus _Molossus_ (fig. 21) includes
the mastiff-bats, characterized by the dental formula _i._ 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
by _Eomops_, with one species; while _Nyctinomops_ includes 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 spread
_Nyctinomus_, with _i._ 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 in _Molossus_, in
many of the species (fig. 22) showing vertical wrinkles. _N.
toeniotis_ (or _cestonii_), 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. brasiliensis_ is common in tropical America, and extends as far
north as California.
Myzopoda and Mystacops.
[Illustration: 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_ (or _Myxopoda_) _aurita_ of Madagascar, and _Mystacops_ (or
_Mystacina_) _tuberculatas_ of New Zealand, the latter of which is
believed to be well-nigh, if not entirely, exterminated. Their
systematic position and affinities are somewhat uncertain; but in the
opinion of O. Thomas[2] the former should typify a separate family,
_Myzopodidae_, in which the latter may also find a place. From all
other bats _Myzopoda_ is 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 subfamily _Mormopsinae_ of the _Phyllostomatidae_,
and on the other to the _Natalinae_ among the _Vespertilionidae_; both
these groups being regarded by him as of family rank.
_Mystacops_ resembles _Myzopoda_ in 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 the _Emballonuridae_. 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.
_Extinct Bats_.
Palaeontology tells us nothing with regard to the origin of the
Chiroptera, all the known fossil species, some of which date back to the
Oligocene, being more or less closely allied to existing types, and
therefore of comparatively little interest. The origin of the order from
primitive insectivorous mammals must have taken place at least as early
as the Lower Eocene. It is, however, noteworthy that several of the
earlier extinct species appear to be related to the _Rhinolophidae_,
which is the most generalized family of the order. Remains of
_Pteropodidae_ belonging to existing genera occur in the caves of
tropical countries in the eastern hemisphere; and the skeleton of an
extinct generic type, _Archaeopteropus_, has been obtained from the
Miocene lignite of Italy, which indicates a form to a certain extent
transitional in character between typical fruit-bats and the
insectivorous bats. The tail, for instance, which in most modern
fruit-bats is rudimentary, with only three or four vertebrae, in the
fossil has eight complete vertebrae; while the teeth of the extinct
form are distinctly cusped. Whether, however, the tail is longer than in
the existing _Notopteris_ of Fiji and New Guinea, or whether the molars
are more distinctly cusped than is the case with the Solomon Island
_Pteropus_ (_Pteralopex_), is not stated. Still, the fact that the
Miocene fruit-bat does show certain signs of approximation to the
insectivorous (and more generalized) section of the order is of
interest. Of the Oligocene forms, _Pseudorhinolophus_ of Europe is
apparently a member of the _Rhinolophidae_; but the affinities of
_Alastor_ and _Vespertiliavus_, which are likewise European, are more
doubtful, although the latter may be related to _Taphozous_. The North
American _Vespertilio_ (_Vesperugo_) _anemophilus_ and the European _V.
aquensis_ and _V. parisiensis_ are, on the other hand, members of the
_Vespertilionidae_, the last being apparently allied to the serotine
(_V. serotinus_).
AUTHORITIES.--The above article is based to some extent on the article
in the 9th edition of this work by G.E. Dobson, whose British Museum
"Catalogue" is, however, now obsolete. Professor H. Winge's
"Jordfundae og nulevende Flagermus (Chiroptera)," published in _E.
Mus. Lundi_ (Copenhagen, 1892), contains much valuable information;
and for _Pteropodidae_ Dr P. Matschie's _Megachiroptera_ (Berlin,
1899), should be consulted. For the rest the student must refer to
namerous papers by G.M. Allen, K. Andersen, F.A. Jentink, G.S. Miller,
T.S. Palmer, A.G. Rehn, O. Thomas and others, in various English and
American zoological serials, all of which are quoted in the volumes of
the _Zoological Record_. (R. L.*)