The aural anatomy of bats

by

Ade Pye

Institute of Laryngology & Otology, University of London, Great Britain

The fine structure of the ears

of 62 species of bats out on the cochleain the horizontal modiolar plane.
from 13 families has been studied by means of These are: the height and width of the cochlea,

serial sections. The bats alive in the number of half-turns, the width and thickness
were caught
West Central and North of the basilar membrane, the height of the cells of
Britain, Indies, Panama,
Africa and with fixa- Claudius and the size of the These
were intra-vitally perfused spiral ligament.
of the chosen because showed
tive in order to obtain perfect preservation measurements were they
internal ear. Where possible, up to six specimens the greatest variations among the Chiroptera.
of each fixed for this and all Nine species of bats representing all the
species were study super-

the ears were sectioned in the horizontal plane. families have been chosen for discussion in this

This will briefly some of the results paper. These are set out in figure 1 with their
paper survey

obtained. classification and other information.

The of bats varied in size, shape In all Microchiroptera the cochlea is relatively
pinnae are

and structure and have not been studied in detail. large compared with other mammals and this is

In many and especially surprising when the of the ani-

bats, e.g. Rhinolophus Hipposideros weight

they are mobile, the movements being correlated mal is taken into account. It consists of from five

with emission Flinn & to seven half-turns. In the cochlea

pulse (Pye, Pye, 1962; Pye Megachiroptera
usual- is smaller and only four half-turns in
& Roberts, 1970). These mobile pinnae are are
present
others immobile and all the 13 examined. The
ly simple in shape, are may species greatest height

be and have leaflets inside the of the cochlea is recorded in

complex or tragi Rhinolophus fumiga-
main part of the ear. tus and this is enormous for an
animal which

Emphasis has been placed on the structure of weighs only 20 g (fig. 1). Also both species of

Hinch- Rhinolophus have the widest cochleae of

the middle ear (Pye & Hinchcliffe, 1968; all the

cliffe & Pye, 1969; and unpublished) and especial- bats studies and the greatest number of half-turns.

ly the of the cochlea 1966a and b, Generally in the Microchiroptera the basilar
structure
(Pye,
cavity is membrane is relatively the base of
1967, and unpublished). The middle ear
very
narrow at

medium in size, but the intra-aural muscles are the cochlea and increases in width towards the

large, especially in the echolocating Microchirop- apex. This is shown by Noctilio, Carollia, Natalus

The stapedius muscle is large and fan-shap- An is found in

tera. and Pipistrellus (fig. 2). exception

ed, having a Paaw's cartilage at the junction with the Rhinolophidae, where the membrane is wider

the tendon. This is in Micro- the first and narrowest the second turn,
cartilage present at turn at

chiroptera, but absent in all Megachiroptera. The increasing from there towards the 2).
apex (fig.
stapedius muscle is in the same bony cavity as In Hipposideros a slight decrease occurs at the

the facial whereas other half-turn. The

nerve (Vllth), in many second overall increase in width of

mammals, such as it is in cavity. the membrane from the base to the doubles
man, separate apex

The stapedial artery persists in all adults, contrary in Noctilio and Carollia, while in Natalus it in-

to
many other mammals, where it is present only creases only by a half. In Rhinolophus this in-

during development. The ossicles are medium in crease is even less, probably due to the fact that

size. the membrane decreases in width from the first

A of has been carried the second and only then increases

variety measurements to turn
again
68 PROCEEDINGS 2ND INTERNATIONAL BAT RESEARCH CONFERENCE

PRINCIPAL HEIGHT NUMBER OF

FREQUENCY WEIGHT OF COCHLEA 1/2 TURNS

NAME CLASSIFICATION PULSE TYPE

None 2.9mm. 4
Pteropus giganteus Suborder None 500g.

V
r

12 Og. 4
Rousettus aegyptiacus Megachiroptera Damped click 12 kHz. 2.8mm.

Superfamily

— Emballonuroidea FM., CF. 60 kHz. 55g. 2.6mm. 6

Most primitive
_

5
Carollia perspicillata Phyikjstomatoidea FM.-Short 80 kHz. 18g. 1.9mm.

Intermediate

CF.-Long 85 kHz. 25g. 2.7mm. 7

Rhinolophoidea

7
CF.-Long 45 kHz. 20g. 3.1mm.
Rhinolophus fumigatus
�

Intermediate

caffer CF. 150 kHz. 8g. 2.2mm. 6

Hlpposideros

135 kHz. 5g. 1.8mm. 5

Natalus tumldlrostris Vespertilionoidea FM.

FM. 45 kHz. 7g. 1.7mm. 5

Most advanced f
CF.

Fig. 1. Some data on the bats reviewed in this paper. FM . . . frequency modulated. CF . .. constant frequency.

towards the apex. It is still a question of specula- ary spiral lamina, for instance in Rhinolophoidea.
tion the width of the membrane is narrowest None of the Megachiroptera echolocation,
why uses

the second turn. The Megachiroptera have rela- for Rousettus, which produces audible pul-
at except

tively wider membranes throughout the cochlear ses

by clicking its tongue (Mohres & Kulzer, 1955)
turns and the increase in width towards the apex is and because of this mechanism is the
only Old

gradual (fig. 2). World fruit bat to be able to live in caves. Of the

In and Natalus the basilar echolocating Microchiroptera, the Rhinolophidae

Rhinolophoidea mem-

brane is thickened at the base of the cochlea. It stand out by producing long, constant frequency
be as much as 50 in thickness at the first pulses, which are also directional in output. As
may p.

half-turn, decreasing towards the apex and dis- has already been described, these are also the bats

appearing by the third or fourth half-turns. In which have a more modified cochlea than those

other is less well defined and in which produce short, frequency modulated pulses.
species thickening
the basilar membrane is thicker at the Natalus is the only other bat in this selection to
Pteropus

apex than at the base. produce directional pulses, which are short in

The cells may be the base of

supporting tall at duration and are
frequency modulated. To some

the cochlea in Microchiroptera, e.g. in Rhinolo- extent Natalus has a more modified cochlea than,
phus fumigatus, Hipposideros caffer and Natalus for example, Carollia, which produces non-direc-

tumidirostris. In all tional, short, frequency modulated

cases they decrease in height pulses. Noctilio

towards the apex, while in Megachiroptera they re- produces both frequency modulated and constant

main about the same height throughout the cochlea frequency pulses, depending on its hunting beha-

(fig- 2). viour. It catches fish from just under the surface of

In Microchiroptera the spiral ligament is large water, but it is not known to what extent echoloca-

in size, especially the base of the cochlea and tion plays part in this
at a
(Suthers, 1965).
there may be extra ossification, called the second- In a computer correlation of cochlear dimen-
 BIJDRAGEN TOT DE DIERKUNDE, 40 (1) -
1970 69

Fig. 2. Histological measurement. BM.W. . .

the width of the basilar membrane. CC . . . the heigth of the cells of

Claudius.

sions with acoustic (Hinchcliffe & type of pulses produced by the animal. The struc-
measures Pye,
1968) it was established that the highest frequency ture of the middle ear in the Microchiroptera is

of sounds emitted quite distinct from that of the Megachiroptera

was significantly inversely cor- or

related with the width the most other but shows rather little varia-
measurements at sec- mammals,

ond, third and fourth half-turns. The highest fre- tion within the sub-order.

quency was also significantly inversely correlated In conclusion it can be said that the Megachi-
with the widths of the basilar membrane. This is have cochlea and
roptera a fairly unspecialised

exemplified by echolocation. Of the Micro-

Natalus and Hipposideros which only one genus uses

produce the highest frequency sounds in this se- chiroptera, those bats which produce long, con-

lection and also have the narrowest basilar mem- stant frequency pulses or which are directional in

their signal output, have the modified coch-

branes at the base of the cochlea (figs. 1 and
2). more

The highest frequency was also inversely related leae. It is. hoped to continue this work not only by

to the mean weights of the animals, which is looking at more types of bats, but also by using

again exemplified by Natalus and Hipposideros. acoustic traumatisation experiments to determine

These bats with the the frequency of the cochlea in bats.

two together Pipistrellus were range

smallest bats studied in the whole survey, while

the largest. This of correlations

Pteropus was set

is the correlation of ACKNOWLEDGEMENTS

supported by high species

weight and the width of the basilar membrane Thanks due The of Clinical Photo-
are to: Department
found graphy at the Institute of Laryngology and Otology for
by us.

preparing the illustrations; Dr. J. D. of King's Col-

associate the Pye
At it difficult to
present seems
lege, London, for the analysis of the recordings,
tape
structure of the external ears with either the varia-
from which the acoustic data have been taken. This

tions found in the structure of the cochlea or the research has been sponsored by the Wellcome Trust.
70 PROCEEDINGS 2ND INTERNATIONAL BAT RESEARCH CONFERENCE

REFERENCES

R. & A. PYE, 1968. The cochlea in Chirop- 1967. The structure of the cochlea in Chiroptera, 3.
HINCHCLIFFE, —,

J. int. Audiol, 7: Microchiroptera: J. 121 :

tera: a quantitative approach. Phyllostomatoidea. Morph.

259—266. 241—254.

& 1969. in the middle of the

—

—, Variations ear
PYE, A. & R. HINCHCLIFFE, 1968. Structural variations in

Mammalia. J. Zool., London, 157 : 277 —288. the mammalian middle Med. biol. 18:
ear.
Ulust.,
MOHRES, F. P. & E. KULZER, 1955. tlber die Orientierung 122—127.

der Flughunde (Chiroptera-Pteropodidae). Z. vergl.

PYE, J. D., M. FLINN, & A. PYE, 1962. Correlated orien-
Physiol., 38 : 1 —29.
tation sounds and ear movements in Horseshoe bats.
1966a. The structure of the cochlea in Chirop-
PYE, A.,
Nature, London, 196: 1186—1188.
tera, 1. Microchiroptera : Emballonuroidea and Rhi-
PYE, J. D. & L. H. ROBERTS, 1970. Ear movements in a
nolophoidea. J. Morph., 118: 495—510.

hipposiderid bat. Nature, London, 225 : 285—286.

1966b. The structure of the cochlea in Chiroptera,
—,

SUTHERS, R. 1965. Acoustic orientation by fish-catch-

2. Megachiroptera and Vespertilionoidea of the Micro- A.,

J. Morph., 119 101—120. ing bats. J. Zool., 158: 319—348.

chiroptera. :
exp.