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ART |Octobe:, = Price 28. 6. net.
[7 AYHISTORY OF

"
BY
GERALD E. H. BARRETT-HAMILTON
B.A.(CANTAB.), M.R.LA., F.Z.S.
\WITH TWENTY-SEVEN FULL-PAGE PLATES. IN COLOUR, FIFTY-FOUR IN
, BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND

FIFTY SMALLER ILLUSTRATIONS

DRAWN BY

EDWARD A. WILSON

B.A., M.B. (CANTAB,)

be 2NS10 7
GURNEY AND JACKSON
10 PATERNOSTER ROW, LONDON, E.C.
1910

in Pranaretior:
lalate sia

A NEW AND REVISED EDITION OF
YARRELL, NEWTON, AND SAUNDERS’

HisTORY OF BRITISH BIRDS —

WILLIAM EAGLE CLARKE, F.R.S.E., F.LS.

Keeper of the Natural History Department, The Royal Scottish Museum; Member of the
British Association Committee on the Migration of Birds as Observed on the British
and Irish Coasts; Corresponding Fellow of the American Ornithologists’ Union;
Correspondirender Mitglied des Ornithologischen Vereins in Wien;

Membre Honoraire du Bureau Central Ornithologique Hongrois ;

Member of the British Ornithologists’ Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES
SPECIALLY EXECUTED BY

MISS LILIAN MEDLAND

HE publication of Yarrell’s “History of British Birds” was
commenced in 1837 and completed in 1843. Its outstanding
merits were at once recognised, and a Second Edition was

called for in 1845, followed by a third in 1856.

From the issue of the Original Edition down to the present
day, Varrell’s “History of British Birds” has generally and
deservedly been regarded as the standard authority on British
ornithology.

In the year 1871 a Fourth Edition was begun, under the
masterly editorship of Professor Newton—the greatest British
ornithologist of all time. Unfortunately Professor Newton’s
official engagements at the University of Cambridge only allowed
him to complete the first two volumes; and in 1882 Mr Howard
Saunders was selected to edit the remaining volumes, a task
which he successfully accomplished to the entire satisfaction of
ornithologists in 1885. |

The many excellences of this last edition advanced the work
more than ever in the public and in scientific favour. To its
stimulating influence is to be mainly attributed the marvellous and
unprecedented activity which has resulted in those extraordinary
advances made in all branches of British ornithology during
recent years—advances which have rendered it essential that a
new work based upon this classical and comprehensive founda-
tion should be issued. itera a

During the period alluded to, a considerable number of new ve
and interesting species have been added to our avifauna. The

[Continued on Page 3 of oon

A HISTORY OF
BRITISH MAMMALS

BY
GERALD E. H. BARRETT-HAMILTON

B.A.(CANTAB.), M.R.I.A., F.Z.S.

WITH TWENTY-SEVEN FULL-PAGE PLATES IN COLOUR, FIFTY-FOUR 1N
BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND
FIFTY SMALLER ILLUSTRATIONS

DRAWN BY

EDWARD A. WILSON

B.A., M.B. (CANTAB.)

IN THREE VOLS.

VOL. I—BATS

Seen Y AND JACKSON
10 PATERNOSTER ROW, LONDON, E.C.

I9IO
Temporary Title Page. |

recrR EW ORD.

peor OPin iD THOMAS, F:R.S. of the” British

Museum of Natural History, the well-known authority

on the mammals of the world, has kindly written the
following appreciation of the forthcoming work :—

“In my official capacity at the British Museum I have
constantly been asked for the name of a trustworthy book
on the subject of British Mammals, and I have had to put
off the enquirers with the promise of the present work, to the
appearance of which every mammalogist, technical or amateur,
has long been looking forward.

“To produce such a work is a difficult, though highly
interesting task, which can only be efficiently completed in
a large Museum. For this no one could be better suited
than the author of the present work, who has for the last
fifteen years worked constantly at the National Museum, not
only on the British, but on all the European and Asiatic
mammalia, and has written on them a considerable number
of valuable scientific papers. He is therefore fully in
touch with the literature of the subject, acquainted with all
its recent advances, and in agreement with the principles
of modern science. He has also had experience in the
collecting and observing of mammals in the field—at home,
in continental Europe, on the Seal Islands of Bering’s Sea,
in Morocco, Kamchatka, and on the veldt of South Africa.”

lil

PRELIMINARY ANNOUNCEMENT.

FOR many years Tomes and Alston’s edition of Bell’s

FIistory of British Quadrupeds has been the standard
work on recent British Mammals, and although several attempts
have been made to oust it from its place, not one can be said
to have been entirely successful. Even the elaborately illustrated
work of Mr J. G. Millais is, by its very price and size, placed
outside the reach of the ordinary reader.

INCREASING INTEREST IN THE STuDy oF BritisH Zoo.ocy.
—The number of students of this branch of British natural
history has greatly increased of late years, and their progress
has been much impeded by the want of some more recent
work than Bell’s. In no other branch has the growth of
knowledge or the change of view developed at a greater rate,
and yet there is no work in existence offering in convenient
form an authoritative summary of the information now avail-
able, with an indication of the lines upon which future research
is most needed.

THe Desicn or THE Worx.—The present work is
designed to meet the above want. Its author has had its
production before him for over twenty years, and the book
has been in his hands a labour of love. During this period
he has been in constant touch (or correspondence) with all
the leading local naturalists, and received from them much
valuable assistance. He has consulted and critically examined
all printed references to his subject in scientific and periodical
literature, and made use of and referred to all notes of genuine
value.

It was at first intended to produce merely a new edition of
Bell’s book, but as the work proceeded this was found im-

possible, so great has been the advance of knowledge, so complete
a2

vi PRELIMINARY ANNOUNCEMENT

the change of view between 1874 (the date of Bell’s last edition)
and 1910. A few of Bell's paragraphs may, indeed, be utilized,
wholly or partially, but the work as a whole will be found so
completely altered as to leave hardly any trace of connection
between the old and the new.

It may with truth be stated that no such work has ever
been written, or attempted, in the English language. Its
publication is bound to give an immense stimulus to the study

of British Mammals.

Drvrsion into Sections.—The work will consist of three
sections, dealing respectively with the Bats, the Land Mammals,
and the Marine Mammals. Besides introductory chapters to
the various orders, each genus and species will be the subject
of an article complete in itself and divided into two portions,
the one treating of the habits and life history, the other explain-
ing the technical aspects of each animal.

Hasirs and Lire Hrisrory.—The habits will be fully
traced, thus making the popular side of the subject very
complete. The literary and historical aspect is not neglected,
the author having made it a point to quote from the best British
writers, with a view to illustrate the work of his predecessors
and fellow-workers.

TecunicaAL Aspects.—The technical portions of each
article will be ample, and contain authoritative details and ex-
planations of matters which, although they admittedly appeal
rather to the museum worker than to the field naturalist, yet
must often be consulted by the latter, and are not at present
available in any single work. Much of this part of the book
will be based on original work of the author, and it goes without
saying that a treatise of this character could emanate only from
one who has probably had, in his own particular line, a unique
experience of combined work both in the field and in the
museum, and whose studies on European Mammals have given

PRELIMINARY ANNOUNCEMENT Vii

him an exceptional knowledge and understanding of those
which occur in the British Islands.

DisTRiBuTION, ETC.—The local names are being treated
somewhat fully, and it is intended to include a full account
of the geographical relationships and distribution, individual
variation in colour, size, or form, with descriptions, measure-
ments, and keys for distinguishing all doubtful species.

ILLustRaTIoNs.——_No expense shall be spared in regard to
illustrations. These are being drawn by Dr Edward A.
Wilson, a trained naturalist as well as a gifted artist, who is
already well known for his beautiful pictures of Antarctic life.
The large numbers of explanatory or quasi-technical drawings
will be unique. The reproduction of the coloured and black
and white drawings into half-tone and line blocks is being
entrusted to Messrs Hislop & Day, Edinburgh.

TypoGrapHy AND Paprer.—The type chosen for both parts
of the work has been carefully selected, the smaller being
used in the technical and the larger in the popular section.
To ensure permanency, specially manufactured paper made
from pure rags will be employed, that for the text being
supplied by Messrs Alex. Cowan & Sons, Edinburgh, and that
for the plates by Messrs John Dickinson & Co., London.

Form or Pusiication.—The work will be published in
about twenty-four monthly parts, at the price of 2s. 6d. each
net. Part I. will be ready on 18th October 1910. When
completed, the whole will form three volumes extra royal
8vo, bound in buckram, gilt top, fore and under edges uncut.
In issuing the Parts it is impracticable to arrange the coloured
and black and white plates in proper order, but a detailed list
of instructions for placing will be inserted in the concluding
Part, which will also contain Title Pages, General Introduction,

Contents, and Index.

CONTENTS

VOLUME I.—Ftriyinc Mammats.

ORDER CHIROPTERA, or BATS.

VESPERTILIONID&, or TYPICAL INSECT-EATING BATS.

» Genus WVyctalus. Genus //yotis—continued.
- Noctule Bat. Whiskered Bat.
Leisler’s Bat. Bechstein’s Bat.

Natterer’s Bat.
[ Notch-eared Bat (non-British). |
[Mouse-eared Bat (non-British).]

S Genus Pipistrellus.
~Common Bat, or Pipistrelle.

Genus Vespertilio.

Berctine Bat. Genus Plecotus.

‘[Parti-coloured Bat (non-British). } Long-eared Bat.

of,

‘ Genus JZyotzs. Genus Larbastella.
Daubenton’s, or the Water Bat. Barbastelle Bat.
[Rough-legged Water Bat (non-

British). ] [The Hoary Bat (non-British). ]

RHINOLOPHID, or HORSESHOE-NOSED BATS.

: } Genus Rhinolophus. Genus Rhinolophus—continued.

Greater Horseshoe Bat. Lesser Horseshoe Bat.
ea ix

VOLUME I1.—Lanp Mammats.

ORDER INSECTIVORA.

Genus /irinaceus.

Hedgehog.

Genus Jalpa.
Mole.

Genus Sorer
Common Shrew.
Pygmy Shrew.

~ Genus Veomys.
Water Shrew.

ORDER RODENTIA. |

Genus Lepus.

Brown Hare.
Scottish Varying or Blue Hare.
Irish Hare.

Genus Oryctolagus.
Rabbit.

Genus Sczurus.

British Squirrel.

Genus Muscardinus.

Dormouse.

Genus Wicromys.

Harvest Mouse.

Genus Afodemus.

Long-tailed Field Mouse.

| Hebridean Field Mouse.

Fair Island Field Mouse. ly ;
De Winton’s Yellow-necked Field
Mouse. a:

St Kilda Field Mouse.

Genus Jus.
House Mouse.
St Kilda Mouse.

Faroe Island Mouse.

Genus Epimys.
Black Rat.

Brown Rat.

CONTENTS xi

Votume II.—Lanpn Mammats—continued.

ORDER RODENTIA—continued.

Genus Arvicola. Genus Microtus—continued,

Water Vole. Sanday Island Vole.

Westray Isl ?
Genus Wicrotus. estray Island Vole

Short-tailed Field Vole. Genus Evotomys.
Hebridean Vole. Red or Bank Vole.
Orkney Vole. Skomer Vole.

ORDER UNGULATA.

Genus Bos. | Genus Cervas—continued.
[White Park Cattle: domesticated. } | [Fallow Deer: introduced. ]

Genus Cervus. Genus Capreolus.
Red Deer. Roe Deer.

ORDER CARNIVORA.
Sub-order FISSIPEDIA, or LAND CARNIVORA.

Genus Wustela. Genus Felzs.
Weasel. Wild Cat.
Common Stoat.

Jura Island Stoat. Genus Vulpes.

Irish Stoat Roe
ris @at:
f Genus JVeles.
Genus Putorzus. Bad
Polecat. ae:
Genus Jartes. Genus Lutra.

Pine Marten. | Otter.

VOLUME III.—Aovatic MammMats.

ORDER CARNIVORA.

Sub-order PINNIPEDIA, or SEALS.

Genus Phoca.

Common Seal.
Ringed Seal.
Greenland Seal.

Genus Cystophora.
Hooded Seal.

Genus Halicherus.
Great Grey Seal.

Genus £ rignatus.
Bearded Seal.

Genus Odobenus.
Atlantic Walrus.

ORDER CETACEA, oR WHALES.

Sub-order MYSTACOCETI, or WHALEBONE WHALES.

Genus Balena.
Biscay Right Whale.
[Greenland Right Whale. ]

Genus Megaptera.
Hump-backed Whale.

Genus Lalenoptera.
Common Rorqual.
Sibbald’s Rorqual.

Lesser Rorqual.
Rudolphi’s Rorqual.

Sub-order ODONTOCETI, or TOOTHED WHALES.

Genus Physeter.
Sperm Whale, or Cachalot.

Genus /Typeroodon.

Common Bottle-nosed Whale.

Genus Zzphius.
Cuvier’s Whale.

Genus Mesoplodon.
Sowerby’s Beaked Whale.

ORDER CETACEA, oR WHALES—continued.

Sub-order ODONTOCETI, or TOOTHED WHALES—continued.
vi y

ie
i

\
a

ns, Genus Monodon. Genus Globicephala.
i” a
Narwhal. Caing or Pilot Whale.

Genus Phocena.

Genus Delphinapterus.
White Whale.

Common Porpoise.

Genus Delphinus.
‘Il Genus Orcinus. Common Dolphin.

Killer.

. Genus 7urszops.

Genus Pseudorca.
Bottle-nosed Dolphin. _

ig False Killer.
: Genus Lagenorhynchus.
; Genus Grampus. White-sided Dolphin.
Risso’s Grampus. : White-beaked Dolphin.

ia APPENDIX.
Extinct Mammals. | Domestic Mammals.

LIST OF FULL-PAGE PLATES IN /@
COLOUR | |

Noctule Bat. Seasonal changes in British —

Daubenton’s Bat. Squirrel.

Dormouse.

British Picich ice:

Bechstein’s Bat.

Natterer’s Bat.
British House Mice and allies.

British Rats.

Common Shrew.
Pygmy Shrew.
Water Shrew. Orkney Vole.
British Voles (i.).
British Voles (ii.).
Red Deer.

Pine Marten.
Stoat in summer.

Weasel and Stoat, to show |
pelage. | Joe Deer.

Stoat and Weasel in winter. Common Seal.
Scottish and Irish Hares. Bottle-nosed Whale.
British Squirrel. Killer.

Common Dolphin.

LIST OF FULL-PAGE PLATES IN
BLACK AND WHITE

Noctule Bat. Roebuck.

Leisler’s Bat. Polecat.

Pipistrelle Bat. Wild Cat.

Serotine Bat. Fox.

Whiskered Bat. Badger.

Long-eared Bat. Otter:

Barbastelle Bat. Grey Seal.

Greater Horseshoe Bat. Ringed Seal.

Lesser Horseshoe Bat. Harp Seal.
Hedgehog. Bearded Seal.

Mole. Hooded Seal.

Brown Hare. Walrus.

Scottish Hare. Hump-backed Whale.
Rabbit. Sperm Whale.
Harvest Mouse. Porpoise.

Long-tailed Field Mouse. Ca’ing Whale.

Water Vole. Twenty Plates of Whales (semi-
Short-tailed Field Vole. diagrammatic),
Red Deer.

With 250 Line Drawings, including Maps.

CONTENTS OF PART

GENERAL INTRODUCTION TO BATS
(To be completed in forty-eight pages.)

VESPERTILIONIDZ .
Genus WVyctalus
The Noctule, or Great Bat

Leisler’s Bat .

ILLUSTRATIONS

Dormouse. (Coloured.)
Noctule Bats.

Heads of—(1) Wyctalus noctula, (2) ie letslert, and (3) Vi esprtilo
serotinus. Gn i

in
_ Nyctalus leisleri—(1) adult, and (2) Nyctalus noctula, recently born, to
show the relative difference in size, especially of foot and to

Wings of—(1) Vyctalus leislert, (2) Pipistrellus pipistrellus, and (3) Mya 7S
daubentoni.

CHIROPTERA.

BATS:

SuB-ORDER MICROCHIROPTERA.

INSECT-EATING BATS.

CHIROPTERA.
BATS.

SuB-oRDER MICROCHIROPTERA.

INSECT-EATING BATS.!

Tue twelve British species are all included in the great sub-
order of insect-eating bats or MWoecrochiroptera. They fall into
two families—the Vespertiliontde, or typical simple-faced, and
the Rhznolophide, or horseshoe-nosed bats.

History:—British naturalists have not been quick to
distinguish the various species, and even at the present time
there are few who boast expert knowledge of not the least
interesting group of British mammals. Appreciation of dis-
tinctions grew, in fact, so slowly, that for original descriptions
of our native species we are in every case indebted to con-
tinental naturalists, the chief honours being shared by
Daubenton of France and Leisler and Kuhl of Germany.

Of British writers, Merrett, in his Pzzax (1660), and Ray, in
his Syzopszs (1693), made no attempt to diagnose species, but
Albin, in his Matural History of Birds? (1740), gave a place
to three—the ‘small common sort,” the Double-eared (namely,

1 For many references to the literature of these animals, see N. H. Alcock’s
‘ paper on the vascular system of bats in Proc. Zool. Soc. (London), 1898, 58.
Dobson’s Catalogue of the Chiroptera in the Collection of the British Museum is still
the main authority for the order as a whole, but will probably be replaced by degrees
by Knud Andersen’s review of the order now in progress. G. S. Miller’s (jun.)
“The Families and Genera of Bats” (Bull. 57, U.S. National Museum, 29th June
1907), is also invaluable, and has been largely drawn upon for the purposes of
this chapter. Since my account of British Bats was written, I have received W. L,
Hahn’s important American paper, “Some Habits and Sensory Adaptations of Cave-
inhabiting Bats” (Biol. Bull. Marine Biol. Lab., Woods Holl, Mass., xv., June to
November 1908, 135-193).

ili, 95.
3

4 INTRODUCTION

the Long- -eared), and a flying fox, the latter, of course, exotic.
Pennant in his earlier editions (1766 and 1768) recognised only
two, which he called the Short-eared and the [one eared, but
in 1776 he added the Great and the Horseshoe, and substituted
the Common for the Short-eared. He had taken a similar
course in his Synopsis Quadrupedum (1771), wherein these four
are definitely mentioned as British, but the Serotine, Pipistrelle,
and Barbastelle are noticed as occurring in France. His
knowledge of all British species, except the ‘‘ Horseshoe,” had
evidently been derived from Gilbert White, who mentions “the
great large bat” in his twenty-sixth letter, written on the 8th
December 1769, and who, subsequently, in September 1771,
devoted a complete letter to it as Vespertzlzio altwvolans. In his
eleventh letter, dated 9th September 1767, he had noticed the
common Vespertilio murinus* and the Vespertzlro aurzbus, but
at this point his knowledge appears to have ended. More light
came from Montagu, who in 1808 distinguished the two Horse-
shoes, and from Sowerby, who between 1804 and 1806 detected
the Barbastelle as a British species; but Bingley in 1809 and
Pennant in 1812 were able to enumerate only half the species
now catalogued. Montagu’s name wznxutus, given to the Lesser
Horseshoe, still stands as the first technical name attached to
the small British sub-species. Thenceforth progress again
lagged. Donovan in 1820 omitted the Barbastelle; as did
Gray” in 1826, together with Leisler’s and the Water Bat, but
brought up the British list to ten by the help of the Pygmy.
Fleming in 1828 reinstated the Barbastelle and the Water
Bat, the latter under the name of emarginatus ; but it was not
until the year 1835 that the first complete list appeared in
Jenyns’ Manual. This naturalist had previously corrected
a notable error of British students—namely, the confusion
of the Pipistrelle, the Common Bat of England, with the
Mouse-ear, the ‘‘Common Bat”? of continental writers. He
still, however, retained the latter in his list, and added the Parti-
coloured Bat, both animals having undoubtedly entered the

1 Called “the little bat” in letter xxvi. of 8th Dec. 1769.

2“A List of the Species of Vespertilionide found in Great Britain,” Zoological
Journal, ii., 108-110, 1826.
3 Myotis myosotis (Bechstein).

CHIROPTERA F

British area. Two others, the Pygmy Bat of Leach and the
Lesser Long-eared of Jenyns himself, were based on errors
of identification, and have since been reunited with the
Pipistrelle and the typical Long-eared. Jenyns’ list therefore
included sixteen species, amongst which were all those now
recognised.

Two years later Bell, by the separation of the Notch-eared
from Daubenton’s, swelled the British list to its maximum of
seventeen species, but in 1838 MacGillivray, by the suppression
of the Pygmy and Lesser Long-eared, reduced it to fifteen. It
has since dwindled to its present dimensions, the few specimens
of the Mouse-ear and of the Parti-coloured Bat captured
in Britain having been either escaped captives or stragglers
from across the Channel, while the inclusion of the Notch-eared
was an error due to confusion with Daubenton’s.

Classification and Nomenclature:—The present work has
nothing to do with general relationships, and, although care
has been taken to follow the most natural grouping, for the
present purposes the various British genera may be regarded as
isolated and unconnected. Here, again, knowledge has found
her way blindly, step by step, and those now best qualified to
judge reverse Dobson’s dictum of 1878 that the Horseshoes are
the highest of their order, and, amongst British bats, have
transferred the long-winged members of the genus Vyc/adus, the
high-flying swifts of their kind, to the head of their tribe, with
Myotis as the most primitive vespertilionid genus, and the
family Rhznolophide as of still lower organisation.

So late as 1874, three authorities no less weighty than
Alston, Bell, and R. F. Tomes, united in one genus such dis-
similar animals as the thirty-four toothed Noctule and Pipistrelle
with the thirty-two toothed Serotine. The remainder of Bell’s
arrangement was in accordance with modern ideas of relation-
ships, but, the rules of nomenclature not yet having crystallised
into definite shape, his names of genera and species were applied
with what would now be considered deplorable inaccuracy. The
researches of Mr Miller resulted accordingly in much re-sorting
of names, the elimination of Scotophzlus and Vesperugo, the
transference of Vespertilio from the thirty-eight toothed to the
thirty-two toothed bats, and the reinstatement of the long and

6 INTRODUCTION

unjustly forgotten Pterygistes, Pipestrellus, and Myotis. Yet
so great has been, until recently, the general ignorance of the
mammalian literature of the past, that it was left to Dr
Andersen in 1908 to show that the Noctule and its congeners
must be assigned to the genus WVycta/us hitherto associated
with the fruit bats. Thus Myctalus replaces Pterygestes
with almost confusing celerity. The changes effected, although
in themselves sufficiently violent and for a time inconvenient,
are likely to be as permanent as any other system of
nomenclature, and have now been, with exceptions as to
details, accepted by the majority of systematic zoologists.
We thus find the vespertilionid bats apportioned to six
genera, viz. :—/Vyctalus with two species; Pzpzstre//us with
one; Vespertilio with one; Myotzs with four; Bardbastella with
one; and P/ecotus with one. The Rhznolophide, with one
genus (Akznolophus) and two species, remain as before.

The present aspect of our study cannot but at first sight
appear pedantic; and the writer would be fortunate who could
avoid it altogether. But there is no excuse for neglecting details,
even of nomenclature, and each change may be welcomed as one
step more toward such finality as is possible to human institutions.

Structure:—For a proper appreciation of the specific and
generic characters of bats, a thorough acquaintance with the
form of their ears and wings and the shape
and number of their teeth is necessary. In
acquiring this, the diagram (Fig. 1) will be
found useful.

In all British bats except the Rhzxolo-
phide there springs from the inner or anterior
margin of the ear a process called the tragus ;
it has the appearance of a second or inner
ear. At the base of the outer or opposite

pane margin, and especially conspicuous in the
}, antitragus ; Rhinolophide, arises a lobe known as the
as Spee antitragus. The shape and size of these
two is of some importance in classifying and

identifying these animals (Fig. 3), but great caution should be
observed in the examination of these parts in preserved speci-
mens, the ears of which may alter considerably. Ina series of

FIG. 1—DIAGRAM OF A
BatT’s EAR.

CHIROPTERA 9

Fic. 2.—EARS OF BRITISH BATs, EXCEPT Plecotus auritus (diagrammatic).

1. Nyctalus noctula. 5. Myotis daubentoni. 9. Barbastella barbastellus.
2. XN. leisleri. 6. MW. mystacinus. 10. Rhinolophus ferrum-equinum.
3. Pipistrellus pipistrellus. 7. M. bechsteint. 11. &. hipposideros,

4. Vespertilio serotinus. 8. MW, nattereri.

8 INTRODUCTION

six Pipistrelles in the Dublin Museum, the ears have shrunk to
little more than half their natural size, thus completely changing
their appearance.’ Some such circumstance probably accounts
for an error in Dobson’s description of the ears of the
Barbastelle. |

Even more important is the wing (Fig. 3), which consists of
a thin cutaneous membrane. This, in its complete development,

A r

as.

Fic. 3.—WING OF A BAT (diagrammatic).

Membranes :—am, antebrachial; /m, lateral; 2m, interfemoral; ad, dactylopatagium ;
pel, post-calcarial lobe. Ske/eton :—A, humerus; 7, radius; w, ulnar; w, wrist; @1, 2, 3, 4, 5,

digits 1, 2, 3, 4,5; mI, 2,3, 4, 5, metacarpals 1, 2, 3, 4, 5; 7, femur; 2, tibia; 7%, fibula;
Pp, foot ; c, calcar or spur; /, tail.

arises at the neck at each side, and, including within its sub-
stance, somewhat like the ribs of an umbrella, the four limbs and
the tail, forms an uninterrupted parachute or patagium around
the body. The greater portion of this parachute is supported
by the forearms. In order to maintain it there is an immense
development of each hand or manus, in which the five fingers or
digits are always present. The basal joints of the digits are
not homologous with the finger-joints or phalanges of other
1 See Andersen, Anum. and Mag. Nat. Hist., Nov. 1906, 372-373.

PART. i.—November. ‘ Price 2s. 6d. net,
Ws (A HISTORY OF
BRITISH MAMMALS

GERALD E. H. BARRETT-HAMILTON

B.A. (CANTAB,), M.R.LA,, F.Z.S.

WITH TWENTY-SEVEN FULL-PAGE PLATES IN COLOUR, FIFTY-FOUR IN
eh BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND
Besos etait: : FIFTY SMALLER ILLUSTRATIONS

DRAWN BY

EDWARD A. WILSON

B.A., M,B. (CANTAB.)

> GURNEY AND JACKSON
'-10 PATERNOSTER ROW, LONDON, EC,
I9IO

ANS 235

in Preparation | a , i

A NEW AND REVISED EDITION OF
YARRELL, NEWTON, AND SAUNDERS’

History oF BRITISH BIRDS —

EDITED BY

WILLIAM EAGLE CLARKE, F.R.S.E., F.L.S.

Membre Honoraire du Bureau Central Ornithologique Hongrois;

Member of the British Ornithologists’ Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES
SPECIALLY EXECUTED BY

MISS LILIAN MEDLAND

pee publication of Yarrell’s “History of British Birds” was—

commenced in 1837 and completed in 1843. Its outstanding
merits were at once recognised, and a Second Edition was
called for in 1845, followed by a third in 1856.

From the issue of the Original Edition down to the present
day, Yarrell’s “History of British Birds” has generally and
deservedly been regarded as the standard authority on British
ornithology.

In the year 1871 a Fourth Edition was begun, under the
masterly editorship of Professor Newton—the greatest British
ornithologist of all time. Unfortunately Professor Newton's
official engagements at the University of Cambridge only allowed
him to complete the first two volumes; and in 1882 Mr Howard
Saunders was selected to edit the remaining volumes, a task
which he successfully accomplished to the entire satisfaction of
ornithologists in 1885.

During the period alluded to, a considerable number of new
and interesting species have been added to our avifauna. The

[Continued on Page 3 of Cover 3

a Ra IPE es

;
-- bty Vea
er

ae
te

CONTENTS OF PART IL.

RAL INTRODUCTION TO BATS
(70 be completed in forty-eight pages.)

=RTILIONIDA—

Genus Nyctalus—
ay Leisler’s Bat .
r ¥

Genus P2pestrellius : , :
iy The Common Bat, Pipistrelle or Flitter-Mouse

— Genus Vespertilio
‘a

Ent
Tice

ILLUSTRATIONS

Skins.—1 and 2, Irish Hare—Summer and Winter. 3 to 6,
Scottish Blue Hare—(3) Summer; (6) Winter; (4) and (5)

.—Typical Molar Teeth of an Insectivorous Bat (after Miller).

ig. 2.—Young Pipistrelle.

vier ae

CHIROPTERA 9

mammals, but rather with the metacarpals or bones of the
hand. In the first digit, thumb or pollex, there are two
phalanges. Being always partly free from the wing, and
armed with a claw, it is used by the animal, when not flying,
for purposes of climbing, but not, as often stated, in locomotion.
In the remaining digits the phalanges are as follows :—

Digit. Phalanges.

i 1 (Vespertilionide) nil (AeAzn0lophide)
EET. 3 - p. °
IV. 3 %) : r

V. 3 - 2

”)
(The terminal phalanges are often cartilaginous.)

The modifications in length or development of the digits
materially affect the length and strength of the wing and the
powers of flight.

As in other mammals, the hand is attached to a limb
consisting of a wrist or carpus with six small bones, a forearm
composed of a rudimentary ulna and a long curved radius, and
of a humerus or upper arm. These constitute the complete
framework of the wing. The length of the forearm is suff-
ciently constant in individuals of the same species to afford in
many cases a satisfactory guide to their identification. The
humerus is peculiar in the large size of the trochiter and trochin
or tuberculum majus and minus.

The shoulder girdle and sternum are often much modified,
especially in the RAznolophide.

The hind-limb consists of the parts usual in mammals. It
is so rotated outwards by attachment to the wing, that the knee
points backwards. The foot is armed with claws and used for
suspension and progression otherwise than in the air. A
remarkable, elongated, cartilaginous process, the calcar, arises
from the inner side of the ankle-joint, and helps to support
the posterior margin of the interfemoral membrane, or that
portion of the wing which occupies the space between the legs
and the tail, The points of attachment of the interfemoral
membrane to the tail and, especially, to the legs are diagnostic
characters, as is also, when present, a lobe which has its point
of origin over the calcar, and is known as the post-calcarial
lobe.

10 INTRODUCTION

A third and smaller extension of the flying integument, the
antebrachial membrane, is stretched in front of the humerus and
the forearm. The wing itself may be divided, for purposes of
description, into two parts, viz., the lateral membrane, lying
between the leg and fifth digit, and the dactylopatagium, or part
directly borne by the fingers.

A comparative study of the wing in young and old bats
of both sexes, as well as in the various species and genera,
is necessary. Bats are born with a high development of the
organs of attachment, the hind claws and thumbs, but the
wings, which make their appearance late in the development
of the embryo, are still quite small at birth, and, even when the
young begin to fly, differ in their proportions from those of the
adult (Plate II., Fig. 2, p. 16). Immaturity may, however, always
be recognised by the imperfectly ossified joints of the digits,
which when fully adult appear as definite knobs or swellings
traversed only by a single indefinite line (Plate I.). In imma-
ture specimens, on the other hand, the imperfect ossification is
shown by the presence of one or two small bones, the epiphyses,
which lie between the two joint-heads and give the appearance
of at least two distinct transverse lines between them. The
process of ossification is evidently of variable duration, I
have inspected a young Whiskered Bat and Pipistrelle, with
the phalanges fully developed as regards their length, though
not as regards their joints, on 26th July and oth August
respectively; but the process of fusing the epiphyses had
not been completed in some Lesser Horseshoes examined by |
Mr T. A. Coward in January, nor in another specimen of the
same species, which came under my own notice in the March
following.

Amongst adults of British bats, the females are usually the
larger,! but, after deducting all differences due to age and sex,
there still remains a considerable amount of variation in size
and proportions, the cause of which is not understood. It is
not, however, sufficient to lessen the importance of the wing asa
feature of high diagnostic value.

The actual dimensions of the various species are shown

1 This is not a universal characteristic of the Chzvop/era, since in some fruit-eating
bats the male is the larger sex.

CHIROPTERA II

by the following table, from which it will be seen that, as
regards size, they fall into two groups, viz., three large bats—
the Noctule, Serotine, and Greater Horseshoe—in which
the forearm measures about 50 mm., and the remainder,
with a‘forearm varying between 30 and 42 mm. _Leisler’s and
Bechstein’s are the largest, and the Pipistrelle is the smallest of
the latter group.

TABLE OF AVERAGE DIMENSIONS :—

3 3 3
=| =| “ a on i=") a =
Sea a Be | 8 Se | #3 | gs | &
a8 5 a eS & se | 2 B
7 = =
Nyctalus noctula
5 to 7 males = Se erin 50 49°5 aly gs 12 91°5 48°25 38 [360]
7toSfemales . jae 515 | [49] ily 12 92°5 50 40 [875]
N. leislert
10 of both sexes : - | [59°5] 42 [41] 17 8 (72°5] 88°5 31°5 | [800]
Pipistrellus pipistrellus iS
9to12males . = : 42 29°5 31°85 [10°5] 5°65 50 25 25 [208]
5 to 11 females . 6 a 43°35 30°28 30°5 11°36 714 50 25 25 213
Vespertilio serotinus .
7 males ° - a a 73 | 52°8 52°5 20°8 86°5 47 43°5 368
11 of both sexes ( wi W4e2 52°1 2:8 20°8 10°5 86°6 47 44 360
Myotis daubentoni
19 of both sexes : | 45°5 86°5 83°5 16°4 91 59°8 82°2 30°2 2238
M. mystacinus
Ganalesmieiers | «22. @(RI46] 84 [32] 15°5 [7:5] | [53°5] | [29] [28] | [228]
M. bechstetnt
5 : 4 [55] 41 [35] 19°5 | (?)8-10 | 64 83°5 33 [250]
M. nattereri s
15 females . 4 A Bl etsy 39°4 37°6 16°5 8 65°6 BB eI 33 [274]
Plecotus awritus
POtnalos i yoy ees. , eter) 48-2), 287-5 | 41-6. |. 17-8 88 | 622 | 313 | 30 11 F955)
17 females . 5 3 a ee Gers 38°8 42:2 18°3 88 65 82 30°9 f
Barbastella barbastellus (| [250
6males . A c . | [48] 37°6 43°4 18°6 67 6674 84°8 82°5 1 | —260)
Rhinolophus ferrum-equinum
ll males . . ‘ é 63°2 53°6 32 24°5 11°2 83°4 36-4 38°9 334°6
5females . 5 65°8 54°5 33°2 25°3 11°8 85°8 36°2 40°5 [350]
R. hipposideros . . - | (88) B7°5 25:2 17°5 8 [52] 23°5 26°5 | [245]

Note.—The square brackets indicate that the enclosed figures are based on measurements of less
than five specimens.

In order to compare the proportions of the species, the
dimensions have in a second table been reduced to terms of
their respective forearms, the reason for selecting the forearm
being that it is probably the most stable feature of the wing.
Thus in the eleven British bats (omitting Bechstein’s, for
which no series is available), the forearm bears the fairly

12 INTRODUCTION

constant relation to the total expanse of about .14, and varies
only about .o2 on either side of this mean, from .13 in the long-
winged Noctule to.15 or .16 in the short-winged Horseshoes
and in the Water Bat. Another proof that in this respect
variation is not great is to be found in the fact that in three
species of about equal size, but dissimilar proportions, such as
the Noctule, Serotine, and Greater Horseshoe, the forearm is
almost of identical length.

The proportions which the length of the forearm bears to
the total expanse and to the length of the head and body are
approximately as follows :—

Total Head and
expanse. body.
Nyctalus noctula 5 : : 53 6
A letslert - : ; 14 a7
Pipistrellus pipistrellus Sea =A i
Vespertilio serotinus . : 5 14 y)
Myotis daubentont : : : .16 8
»» mystacinus : : : 149 7
» Oechsternz : ‘ ; 16 7
3 wmatiereri : : : ‘ 14 8
Plecotus auritus . : ; : 14 7
Barbastella barbastellus . : 14 By
Rhinolophus Serrum-equinum St ie 5-.16 8
,», Atpposideros J

The following table shows that, of the vespertilionid bats,
Plecotus and Barbastella excel in length of tail, a feature combined
with exceptional length of lower leg and denoting a capacious
interfemoral membrane. The remaining species do not present
many differences in the former respect, but in JVyctalus
noctula and Prpistrellus the lower leg is shortest. The foot is
longest in JZyotzs daubentont and LN. xnoctula, and shortest in
Larbastella. The expanse is greatest in Vyctalus, Prpistrellus,
Vespertilio, and Myotzs natterert in the order named, and
least in MZ. daubentont. A narrow wing is indicated in
Nyctalus, and less so in Vespertzlio and Larbastella, by the
elongated third metacarpal, considerably longer in each case
than the fifth, The converse is the case in Pzfzstrel/us and
M. nattereri, in which the two metacarpals are about equal.
Larbastella appears to possess the most ample wing, every part
of it being exceptionally developed, although the foot is small.

CHIROPTERA 13

TABLE SHOWING RELATION OF GENERAL DIMENSIONS TO LENGTH
OF FOREARM :—

2 5, = =
Es an 50 3 3s 3
5 3 pele Sepa Caer taaaes
2) 5 i) = 3 nA
& ra a fe] S <3)
) =
4
|
f Nyctalus noctula
7males . : n 0 100 99 35 24 183 96 76 [720]
N. leislert
Both sexes : 5 A 100 [97] 40 19 {172] 91 75 (714)
Pipistrellus pipistrellus
12males . . ‘ . 100 108 [85] 19 169 84 84 [705]
11 females. S 5 : 100 100 37 23 165 82 82 703
Vespertilio serotinus
10, both sexes . F 100 100 39 20 166 90 84 690
Myotis daubentont
13 males . : 5 | or “ P
Bias | kf] 00 91 44 25 163 88 83 613
M. mystacinus
6males . . . O 100 aid 45 [22] [157] [85] [82]
M. bechsteint =
5 < 5 . C 2 100 [$5] 47 [19] 154 81 80 [605]
M. natterert :
15 females. é A : 100 95 42 20 166 84 84 [695]
Plecotus auritus
10 males . . 7 ‘ 100 111 46 23 166 83 80 ) [680]
17 females. 5 z 5 100 109 47 21 167 82 79 f
Barbastella barbastellus ; : of
6males . 4 F : 100 114 49 18 176 92 86 [678]
Average \ 2 2 8 81 688
(excluding M. bechsteint) J au = = Ae u
Rhinolophus ferrum-equinum
ll males . e i ° : 100 60 46 21 155 66 72 624
5 females . . G + 100 61 46 22 157 67 74 [642]
R. hipposideros . c : 100 67 47 21 [139] 63 71 [653]

The few figures available for J/. dechsternt indicate an
exceptional bat ; short-tailed, but with long lower leg and broad
short wing.

The Rhznolophide are markedly different, being characterised
by an exceptionally short tail and third metacarpal. The fifth
metacarpal is also short, but longer than the third. The
expanse is poor, the wing short and broad with poorly
developed interfemoral, but large antebrachial, membrane.
The lower leg is well developed, the foot moderately so, the
longest digit below the average.

Although naturally most at home on the wing, all the

14 INTRODUCTION

vespertilionid bats—statements to the contrary notwithstand-
ing—walk well, the sequence of their movements being that
typical of quadrupedal locomotion. The Horseshoes alone
are unable to make even a pretence of walking, and when
on a flat surface, lie prone and helpless, their feebleness in this
respect, since forearm, leg and foot are all well developed, being
probably due to interference with the action of the forearm
by their well-developed antebrachial membrane.’ In spite
of this, they readily climb backwards, by means of their
feet and thumbs, up an incline, no matter how steep, and
have no difficulty in taking wing from a flat surface, springing
with surprising agility into the air, even from narrow or cramped
surroundings, But, although no healthy British bat usually finds
any difficulty in taking flight from a flat surface, it is possible that
some individuals, perhaps after a heavy meal, may at times lack
the necessary vigour; and all require lateral space for a forward
jump and expansion of the wings. In climbing, the free tip of
the tail is, in the vespertilionid bats, used as a kind of extra
limb, and with it the inequalities of a rough surface are sought
and held. In traversing the wires of a cage, the extended tail
acts as a support like the stiffened rectrices of a woodpecker or
tree-creeper, and even in horizontal progression it assists in
throwing forward the body, being brought into contact with the
ground on either side alternately in correspondence with the
action of the feet. There is no evidence that the feet are ever
used to aid in catching or holding the prey, but their vari-
able proportions in the different species, from relatively largest
in the Water Bat and Noctule to particularly small in the
Barbastelle, must have some meaning, and need explanation.
The thumbs might easily be used for holding or securing food,
as they are in the fruit bats and in a berry-eating bat of
Jamaica,’ but I know of no evidence on this point, except in the
case of a captive Natterer’s Bat.

Dentition :—-Young bats, except the Akznxolophide, are born
with a complete milk-dentition, the function of which is to enable
them to hold on to their mothers’ nipples. These teeth differ

1 As in other bats which walk poorly ; see Dobson, Catalogue of Chiroptera, 466.
2 Phyllonycteris sezekorin of Grundlach and Peters, as observed by Osburn, af. cév.
infra, p. 22.

CHIROPTERA 5

both in number and form from those of the permanent dentition.
They are homodont, slender, sharply recurved and cusped, and
cannot by their shape be divided into the ordinary divisions
found in the adult—incisors, canines, premolars, and molars—
but rather resemble the teeth of seals and cetaceans. They are
weak and insignificant, and may persist in the edges of the
alveoli until the permanent teeth are nearly grown;' their
number for each species is not known, but in A/yotzs it is—

ne Bet mc mpm a
—3 I-1I 2-2
a combination which probably represents the maximum in cor-
respondence with the high number of the teeth of the per-
manent dentition in this genus.

The permanent teeth consist, as in other mammals, of four
kinds—incisors, canines, premolars, and molars. The cheek-
teeth are acutely cusped, with a pattern more or less W-shaped.
They vary in the different genera from thirty-eight in JZyotzs to
thirty-two, as in Vespertelio and Rhinolophus. In most genera
one or more of the teeth are so minute as to be functionless,
difficult to find, and indeed they sometimes drop out of the jaw
of the adult animal, so that the total, as given in the technical
formule, is often deceptive to the student. The molars, the
last three teeth in either jaw, are not affected by this reduction,
neither are the canines at the other end of the cheek-series,
nor, except in the Ahzxolophide, the incisors. The teeth,
therefore, which most call for careful study are the premolars,
and they, fortunately, are easily recognised, since they are all
the cheek-teeth between the last three (the molars), and the
first, the conspicuous canine.

If we accept the current view of evolutionists, that the
tendency of mammalian dentition lies towards reduction in the
numbers of individual teeth, then the genus JZyotzs possesses
the lowest type of dentition, the full formula in this case being—

wa 2 va 3-3 3-3
z RTE i ei ee
Sage Lee i eee 5 S.0

1 For further details see L. F. E. Rousseau, “‘Mém. Zool. et anatomique sur la
Chauve-Souris commune,” etc., in Guerin’s ag. de Zool., 1839, pl. 7; also Wilhelm
Leche in Lund’s Universttets Ars-skrift, xil., 1-47, 1876, pls. 1 and 2 (analysed in
Archiv fiir Naturgeschichte, i., 1877, 353-364), and xiv., 1-37, 1878, pls. 1 and 2.

16 INTRODUCTION

This represents a very near approach to the typical complete
dentition of the higher mammals, from which it appears to have
receded, by the loss of one pair of upper incisors and of a pair
of premolars above and below. Further reduction is fore-
shadowed by the small size of the first two upper premolars,
especially the second upper, while the tendency is_ visible,
although less markedly so, in the second lower premolar.
Ascending the scale, we find Plecotus, having dropped a pair of

2 vat
and a total of thirty-six teeth.

2
upper premolars, with paz ——
In Pipistrellus, the next highest, a pair of lower premolars has
vanished (both remaining pairs being quite small), and the
2? total thirty-four. Barbastella and Nyctalus
2-2
have also a total of thirty-four, but in the latter the anterior
upper premolar has become rudimentary, and the corresponding
lower one is much reduced in size. In Vesfertz/io the former
tooth has entirely disappeared, reducing the total to thirty-two,
all of which are functional. In A4zxolophus the number is
—2

formula is pv

. . ° . 2 .
again thirty-two, but in this case the premolars are , with
2
3
the anterior upper and central under often rudimentary: the
= s I—I . .
incisors, , differ in number and arrangement from those
2-2

of the vespertilionid bats; the upper ones are quite minute and
functionless.

The missing teeth required to complete the _ typical
mammalian dentition of forty-four have usually been taken as
upper incisor 1 and premolar 1 in both jaws, the next to dis-
appear being either premolar 2 or 3. Mr Oldfield Thomas,
however, gives good ground for regarding the missing pre-
molar as 2.1 The posterior molar of both jaws is reduced in
size, but it is present in all British species.

The shape and form of the teeth will be best understood
by a reference to the diagrams representing the dentition of
each species. The premolars resemble the simple canines, but
are conspicuously smaller ; the last is always much the largest,
and is double rooted.

' Ann. and Mag. Nat. Fist., April 1908, 346-348,

rivet atl
1 Tea

. | iseories: of very many athens) which were formerly little known,
Rave been fully elucidated, while, speaking generally, an immense
. increase in our knowledge on such important subjects, as Migra-
' tion, Distribution, Habits, Nidification, Plumages, has accrued:
_ And lastly, a new and important branch of study has been instituted
- —namely, the recognition of the various Racial Forms or Sub-
species exhibited by certain birds in the British Islands, on the
Continent, and elsewhere.

A great advance has also been made toward a more satis-

it a factory system of. classification of the Aves—always a difficult
_ subject—and this necessitates departures from the older views.

To bring this Standard Work thoroughly abreast of the most

recent knowledge in all these sh hide is the object of the
| Sane Nige work, _

It should be remarked that while it is not intended to go fully

il into Synonomy, yet, where changes of nomenclature have been
necessary in order to conform with the Law of Priority—the only
method by which complete uniformity in nomenclature can ulti-
mately be attained—the names used in the Fourth Edition of
. Yarrell’s “British Birds” and in Saunders’ “Manual,” and the
_.Trinomial Names of the British Racial Forms, and of those
occurring in Britain as visitors from the Continent, will be quoted,
as will also the Original Name under which the species was
_ described.
ead 6 requesting Mr Eagle Clarke to undertake the duties of
_ Editorship, the Publishers desire to make it known that they are
acting under the advice of the late Mr Howard Saunders, who
placed all his collected notes for a New Edition at Mr Eagle
-Clarke’s disposal for this purpose. That Mr Eagle Clarke is emin-
ently fitted for the work is well-known to all who are interested in

ornithological science. Through his investigations of the subject,

and contributions to its literature, he has long been recognised
aS one of the foremost authorities on all that relates to British

birds. He has studied our native birds in many portions of

the British Islands, and has visited a number of bird-haunts
in various parts of Europe in order to become acquainted
iS ‘their Continental homes with the visitants that seek our
shores.

On the important matter of the Migrations performed by
British Birds, Mr Eagle Clarke’s knowledge is unrivalled—a

material fact, when it is called to mind how little has been said

on this most important subject in any published History of
_ British Birds.

A new and important feature of the New Work will be a

_ Coloured Plate of each species. These will be reproduced in the
_ best style from original drawings specially executed for the work
by Miss Lilian Medland, F.Z.S., an accomplished and well-
Sane Mat vaisilin ls an

GURNEY & JACKSON
| 10 PATERNOSTER ROW, LONDON,’ E.C.

In the Press, to be published shortly i

STUDIES IN BIRD-MIGRATION

BY WILLIAM EAGLE CLARKE, F.RSE. PLS.

Member of the British Association Committee on the Migration of Birds as ie i

Observed on the British and Irish Coasts, and Author of its Final
Reports, 1896-1903, etc.

With Numerous Illustrations and Maps

\VV'TH__the exception of the two initial chapters this

work is entirely original, being the result of the
author’s investigations and personal experiences. These have

extended over many years, during which exceptional opportunities iN
have been enjoyed for acquiring knowledge on Bird-migration — ~

generally, and its British aspects in particular.

In 1884 Mr Eagle Clarke was elected a member of the British — |
Association Committee on the Migration of Birds as observed
on the British Coasts; and on the completion of that great .

enquiry, he was requested by his colleagues to prepare the final
reports on the results obtained—a difficult and arduous task,
which he accomplished in 1903. ae
During the preparation of these reports (five in number), Mr
Eagle Clarke became much impressed with the advantages which
were likely to accrue from placing a trained ornithologist at
a number of the most favourably situated observing-stations
around our coasts. If this could be done, he believed that some
of the difficulties which the phenomena presented might be

solved, and our knowledge regarding the subject generally —

considerably advanced.

This conviction led him to undertake, by the special permis-
sion of the Elder Brethren of the Trinity House and the Commis-
sioners of Northern Lighthouses, a series of personal investigations

at various light-stations, each of which was selected for a special

purpose. In all, Mr Eagle Clarke has resided ho fewer than forty-
two weeks in these isolated and remote observatories; the stations
visited being the Eddystone Lighthouse, the Kentish Knock
Lightship (33 miles off the Essex coast), the lighthouses on the

Flannan Isles and Suleskerry (both lying far out in the Atlantic),

and the lighthouse at Fair Isle (the ‘British Heligoland”). He
also visited the Island of Ushant—an important station —and
Alderney for similar purposes; and is to spend a month or more
of the autumn of 1910 at St Kilda, for the purpose of carrying
the investigations to the outmost fringe of the British area.

With these unrivalled experiences for its foundations, the
book should not only prove a valuable contribution to the subject of
Bird-Migration, but should occupy a place essentially its own in
ornithological literature.

GURNEY & JACKSON
10 PATERNOSTER ROW, LONDON, E.C.

Oliver and Boyd, Printers, Edinburgh.

*
;

WARDS OF TWO HUNDRED AND
oO

LLER ILLUSTRATIONS
)
AC

LTON
FIETY-FOUR IN

PTO

2 i 44
ats

S S
IN COLOUR

CKSO

nt) ae
J

ETT-E
nD

(hh
a
Ht)

PEE ES AMES
'

PLATES

es
Soe ee

AGE

in Preparation

A NEW AND REVISED EDITION OF |
YARRELL, NEWTON, AND SAUNDERS’

HISTORY OF BRITISH Ra

WILLIAM EAGLE CLARKE, F.R.S. E. F. L S,

Keeper of the Natural History Department, The. Royal Scottish Museum ; Modiner of the
British Association Committee on the Migration of Birds as Observed on the British —
and Irish Coasts; Corresponding Fellow of the American Crnithologists’ Union;
Correspondirender Mitglied des Ornithologischen Vereins in Wien;
Membre Honoraire du Bureau Central Ornithologique Hongrois;

Member of the British emp aclontsts: Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES.
SPECIALLY RXECUTED its

MIS “LILIAN MEDLAND,

(Pe publication of Yarrell’s “History of British Birds” was
commenced in 1837 and completed in 1843. Its etn
merits were at once recognised, and a_ Second Edition was
called for in 1845, followed by a third in 1856. Hi
From the issue of the Original Edition down to the ese it
day, Varrell’s “History of British Birds” has generally an
deservedly been regarded as the standard authority on Britis!
ornithology.
In the year 1871 a Fourth Edition was begun, iNew ay
masterly editorship of Professor Newton-—the greatest British
ornithologist of all time. Unfortunately Professor Newton’ ‘
Official engagements at the University of Cambridge only allowed
him to complete the first two volumes ; and in 1882 Mr Howard
Saunders was selected to edit the remaining volumes, | a task |
which he successfully accomplished to the entire Satisfaction of
ornithologists in 1885.
The many excellences of this last. edition advanced the work
more than ever in the public and in scientific favour. To its —
stimulating influence is to be mainly attributed the marvellous and |
unprecedented activity which has resulted in those extraordinary i
advances made in all branches of British. ornithology during: by
recent years—advances which have rendered it essential | ‘that | a
new work based upon this classical and COM DEER EHSI NE | founda.
tion should be issued. Nit
During the period alluded to, a SLE SA ‘Daan! eet new
and interesting species have been added to our Pane iia | a

CONTENTS OF PART III.

GENERAL INTRODUCTION TO BATS
(70 be completed in forty-eight pages.)

VESPERTILIONIDA—

Genus Vespertzlo
The Serotine.
The Parti-coloured Bat

Genus J/yotzs
Daubenton’s, or the Water Bat
[The Rough-legged Water Bat]
The Whiskered Bat .

ILLUSTRATIONS

FULL-PAGE (Coloured and Black and White).

Natterer’s Bat. (Coloured.)

Wings of—(1) Wyctalus noctula,; (2) Vespertilio serotinus
lophus ferrum-equinum.

Serotine Bat.

Heads of—(1) Myotes mystactnus ; (2) Myotis daubentont
natterert, (4) Pzpestrellus pipistrellus.

Whiskered Bat.

FIGURES IN TEXT.

Diagram of Arrangement of Teeth of Vespertelio serotinus.

Front View of Incisors and Canines of Vesperielzo serotinus.

Diagram of Arrangement of Teeth in Genus J/yodzs.
Front View of Incisors and Canines of J/yotzs daubentonz.

Front View of Incisors and Canines of JZyotes mystacinus.

PAGES

17-24

, (3) Rheno-

, (3) Myotes

CHIROPTERA 17

The crown pattern of the upper molars, often reduced in size
from behind forwards, is formed by three main cusps—the inner
anterior (the protocone), the outer anterior (the paracone), and
the outer posterior (the metacone)—and at the extreme outer
edge three small cusps—the anterior (the parastyle), the median
(the mesostyle), and the posterior (the metastyle). The styles
_and cones are connected by conspicuous ridges—the com-
missures—running respectively from parastyle to paracone,
paracone to mesostyle, mesostyle to metacone, and metacone to
metastyle. The result is a conspicuous W-pattern, the varia-
tions in form of which are of much systematic importance; this
pattern is not obscured by the presence of an inconspicuous
inner posterior hypocone. The second molar is usually the
largest, the third always the smallest (Plate II., Fig. 1).

The lower molars correspond to the upper in size, but the
posterior is relatively larger. There are five cusps—the outer
anterior (the protoconid), the inner anterior (the paraconid), the
inner median (the metaconid), the outer posterior (the hypoconid),
and the inner posterior (the ectoconid). These teeth resemble
the outer higher portion of the upper molars reversed with the
two segments of the W disconnected. In the third lower molar
the posterior segment is usually smaller than the anterior. The
most prominent cusps are, in the first two upper molars the
metacone, in the third upper the paracone, in the lower molars
the protoconid’ (Plate II., Fig. 1).

The key to the teeth and skulls at the end of this article
will probably be found useful.

The colours of bats, although not usually bright as com-
pared with those of many other mammals, include as exceptions
perhaps the brightest tints of the whole class. Although toa
great extent creatures of the night, their external coloration is
evidently subject to laws similar to those which govern that of
diurnal mammals.

The general tints are due often, as in Leisler’s Bat, to the
tips of the hairs, from which the bases differ in colour. In
other cases, as in the Noctule, the hairs are almost or entirely

1 For further details see Herluf Winge, “Jordfundne og nulevende Flagermus
(Chiroptera), fra Lagoa Santa, Minas Geraes, Brasilien,” from 2. Museo Lundit, vol.

il., paper I, 1893.
Cc

18 INTRODUCTION

unicoloured. As in the mammalia generally, the under is fre-
quently lighter than the upper side, and in some exotic species
approaches pure white. The ordinary pelage tints are browns
of varying shades, but brilliant yellow, rufous, white, blue-grey,
and black sometimes appear in certain foreign species. In a
few cases there is a conspicuous “piebald” coloration due to
the combination of strongly contrasted shades, as in Scotophzlus
ornatus (Blyth). An African false vampire (Lavza frons, of
Geoffroy) is said to derive the orange colour of its fur from a
powder secreted by a gland.

Bats may follow any local or geographical tendency to
develop certain tints which manifests itself amongst other verte-
brates, as when in dry arid regions they assume the character-
istic ‘‘desert” coloration.’

Many species, such as the Pipistrelle, are characterised by a
wide range of colour variation, amongst which three phases—
brown, rufous, and dusky—are conspicuous, but are connected by
many intermediate types. The last corresponds to melanism in
other mammals, and may be regarded as an almost normal
form of variation. Albinism, on the other hand, occurs as an
abnormality, probably pathological—which may affect the whole
body, including the eyes, wings or ears, or may be restricted to
certain definite regions only.

Sexual differences of coloration are said to exist in the
Serotine, but I cannot find them in this or any other
British bat. In Dobson’s experience females are usually
darker than males, but in certain South African instances the
reverse is said to be the case.” The most conspicuous
examples occur in India, where the female of Azpposzderus
fulvus assumes a vivid coloration during the breeding season.
In Scotophilus kuhlit, a common bat at Calcutta, the under-
side of the pregnant female changes from pale straw colour or
whitish to rich saffron, ‘exceeding that of the canary bird.” ®

The colour distinctions between young and old, and

1 E.g., Thomas’ Rhinolophus denti of Kuruman, South Africa (Ann. Mag. Nat.
fTist., May 1904, 386); and his Vespertilio matschiet of Aden (Ann. Mag. Nat.
fTist., Nov. 1905, 573.

2 E.g., Miniopterus dasythrix of Temminck and J. fraterculus of Thomas and
Schwann ; see Proc. Zool. Soc. (London), 20th Feb. 1906, 161-162.

3 Proc. Zool. Soc. (London), 4th March 1873, 241-252.

CHIROPTERA 19

between adults at different seasons of the year, have been little
studied, and nothing is definitely known as to how the fur is
moulted. Usually the immature pelage is more uniform, due to
absence of grizzled hair tips, but it may be either darker or
lighter than that of the adult. In young Natterer’s and
Whiskered Bats the ventral and dorsal surfaces are more
strongly contrasted in coloration, while the first coat of young
Horseshoes is recognisable by its grey tint.

Secondary sexual characters, other than those of colour,
are commonly found amongst exotic bats, the males of which
exhibit many remarkable glandular structures.1 The measure-
ments given in this work suggest that the wings of females are
on the average larger and the animals themselves heavier than
the males.

Variations other than those of colour are still in need
of attention, and further study may reveal the presence
of local distinctions at present unknown. Meanwhile it
has been shown that both the British Horseshoes are on
the average slightly smaller than those of the neighbouring
continent ; the same may prove to be true of the Long-eared
and, I suspect, also of the Barbastelle. The British Bechstein’s
Bat also presents some differences, but this bat is so little
known that the nature of its variations is obscure.

Abundance and geographical distribution:—Outdoor know-
ledge, no less than museum study, has advanced haltingly,
and all the more so since bats, like cetaceans, are the bane of
some writers on geographical distribution, the possession of
wings in the one case and of swimming equipment in the other
having apparently convinced them that both orders must be
exempt from the laws governing the distribution of other
mammals. This error, combined with ignorance of distinguish-
ing characters, has in the past led to much confusion of species
and incorrect notions as to their relative numbers. Locally
abundant bats such as Leisler’s, and others so common and
widely distributed as the Whiskered, have shared a reputation
for rarity with Bechstein’s, while Natterer’s, Daubenton’s, and
the Barbastelle have been surrounded with much unnecessary
and mysterious obscurity.

1 See also Osburn, of. cit. infra, 77.

20 INTRODUCTION

The truth is that every undoubtedly British species, except
the little-known Bechstein’s, and, perhaps, the Barbastelle,
is now known to be somewhere common. In some of the
southern English counties the numbers occurring in a single
locality and even in the same sleeping place, although not usually
in the same crevice, are surprising. In the artificial caverns at
Henley-on-Thames at least four’ species congregate, while ten
have been taken within the limits of the surrounding demesne.
Again, in Normandy MM. Paul Noel and H. Gadeau de
Kerville found no less than seven species—the two Horseshoes,
Daubenton’s, Natterer’s, with the Notch-eared, Whiskered, and
Mouse-eared—resorting to a single cavern.’

Neglect of bats is a grave error in studying geographical
distribution, since, inasmuch as these creatures are possessed of
the power of surmounting obstacles which to other mammals
must be insuperable, their permanent restriction to definite
regions must be due to causes of fundamental importance.
And, whereas the wings of bats should have enabled them
to cccupy with uniformity the entire extent of the British
Islands, we find in fact that their distribution therein is not less
restricted than that of other mammals.

In considering the distribution of bats, it must be re-
membered that physical features, such as woods, water, and
caves, have a very distinct influence on their abundance.
Probably no species are less particular about their haunts than
the Long-eared and Pipistrelle, and they accordingly may be met
with in almost every part of the kingdom, one or the other
having found its way to the Shetlands and to remote western
Irish islets. Some, especially the Noctule and Leisler’s, and
perhaps the Whiskered, are probably dependent for their
existence on woods, while caves, at least in winter, are no
doubt essential for the complete comfort of the two Horse-
shoes. Daubenton’s, on the other hand, restricts itself to the
neighbourhood of sheltered streams or stretches of water.

The most striking fact in the distribution of British bats is
the progressive westward decrease in species, from Normandy
with fifteen, through Great Britain with twelve, to Ireland with

1 Daubenton’s, Natterer’s, Bechstein’s (once), and Lesser Horseshoe (once).
2 Bull. Soc. Amis. Sct. Nat. (Rouen), 4th April 1901, reprint, 2.

CHIROPTERA 21

only seven. Skandinavia, on the other hand, has nine and
Denmark twelve.’

The second fact is the paucity of species in Scotland.
Exact details are not in every case available, but only the
Pipistrelle and Long-eared are known throughout that country.
Daubenton’s may fairly be classed with them, as it is attached,
often in abundance, to the larger water systems, but for the
rest, all that Scottish naturalists can point to are isolated’
occurrences of the Noctule, Whiskered, and Natterer’s, giving
a total of three regular and three apparently scarce species.
Mr William Taylor’s recent record of the Noctule so far north
as Elgin probably indicates this bat as a regular member of the
Scottish Fauna; and inasmuch as in Skandinavia Natterer’s,
although rare, is known to occur up to 59° north latitude, the
Whiskered to 68°, while Daubenton’s is one of the commonest
bats, and all three are known from Ireland, it is difficult to avoid
the assumption that the range of all three in Scotland is more
extensive than published information would lead us to suppose.

A third noticeable fact is, that amongst the seventeen species
inhabiting the six countries, Normandy, Denmark, Skandinavia,
England, Scotland, and Ireland, no less than seven distinct
variations in distribution occur. Only four, xoctzla,
piprstrellus, daubentont, and auritus are regularly met with
in all five, but two more, mystacenus? and natterert,® differ only
in their apparent absence from Scotland. Two, J/ezsdert and
hipposideros, are absent from Denmark, Skandinavia, and
Scotland: one, barbastellus, only from Scotland and Ireland.
Two are confined to Normandy and a restricted portion of
England: of these, ferrum-eguinum is southern and south-
western, while sevo¢inus is distinctly south-eastern. Lastly,
two, murinus and nzlssonz, are found only in Skandinavia and
Denmark, while two more, myosotzs and emarginatus, are
known only in Normandy. Dasycneme occurs in Normandy
and Denmark, but is doubtfully a native of Skandinavia.
The partial and individually varying British ranges of the
majority of the species is in strong contrast to their wide range

1 | exclude from the Skandinavian fauna, for the purposes of this article, Pzf7-
strellus nathusii and Myotis dasycneme ; the status of either is doubtful.
2 With two Scotch records. 3 With one Scotch record.

22 INTRODUCTION

elsewhere. It is evident that the islands are the meeting-
ground for bats having elsewhere different types of distribution.
Bechsteint and nathusiz are so little known that they may be
excluded from consideration.

Habits:—The general habits of all insectivorous, as com-
pared with frugivorous bats,’ are at first sight very similar, and it
is only of late years that they have attracted much attention from
British naturalists, thanks mainly to Messrs Alcock, Coward,
C. B. Moffat, Charles Oldham, Tomes, and Arthur Whitaker.’

Most species have their peculiar haunts, which they frequent
in preference to, but not to the entire exclusion of, all others.
The narrow-winged Noctule and Leisler’s Bat, the fleetest
of their race, often accompany the swifts to great heights in
wooded districts. These and other strong fliers swoop falcon-
like on their prey when in full flight, but the Long-eared, using
its ears like sensitory tentacles, the Whiskered, and probably
Natterer’s and the Barbastelle, although not incapable of
captures in mid-air, more usually snatch resting insects from
the branches of trees. Daubenton’s, on the contrary, obtains
its food hovering over the surface of water, and is often accom-
panied by others, especially by Natterer’s. The Serotine loves
glades and gardens, feeding around trees, although not actually
amongst the branches. The Pipistrelle, whilst it affects a
variety of situations, is pre-eminently the familiar bat of our
farmyards and houses. The rare Bechstein’s has not been
definitely connected with any special habitat, but the butterfly-
like sailing on expanded wings of the two Horseshoes is quite
characteristic. The latter possess such a wonderful power of
threading their way through intricate places in captivity that it
seems almost impossible to believe that they do not naturally

1 This distinction, like others used for purposes of convenience, is not absolute,
since some of the bats with insectivorous dentition eat fruit (see Dobson’s Catalogue
of Chiroptera, 390, 466, 503, 511, 514, etc.), suck the blood of larger animals (of.
ctt., 155, 466, 486, 549), or catch shrimps, fish (of. czZ., 397), small birds or mice
(A. M. Primrose, also E. Gleadow, Journ. Bombay Nat. Hist. Soc., xvii., 1021-1022).

* Some important allusions to habits are scattered through Dobson’s Caéalogue of
Chiroptera. P.H. Gosse’s Naturalist in Jamaica, 1851, contains original observations
on the bats of that island. His allusions to the crepuscular habits (p. 162) and segrega-
tion of the sexes (p. 294) in some species are probably amongst the earliest published
references to these points in their economy. Another paper of great interest, W.
Osburn’s “ Notes on the Cheiroptera of Jamaica,” in Proc. Zool. Soc. (London), 24th
Jan. 1865, 61-85, deals with fruit-eating bats.

CHIROPTERA 23

hunt amongst branches of trees or it may be in ivy or rocks.
The suggestion is supported by Mr Coward’s observations on
captives of the larger species, which showed a remarkable
aptitude for seeking their prey upon the ground, a habit not
known in any other British bat, and recalling those of the
African Lavias, which also possess a nose-leaf.

All bats love shelter, and are much impeded, or entirely kept
at home, by foul weather, even in summer, at which season
it is probable that exceptional cold or wet causes the death
of many from starvation.” The delicate Lesser Horseshoe is
probably the most susceptible to wind. The larger kinds, and
even the lesser when they have to deal with small insects,
devour them on the wing, usually rejecting the wings, elytra, or
other indigestible portions, which have been noticed falling
thickly from a party of feeding Noctules. As a rule, bats of
small size, such as the Pipistrelle, retire from flight for a few
moments after the capture of a large insect—a course scorned by
such strong flyers as the Noctule or Leisler’s Bat. To help
them in retaining their grip, it appears almost certain that all the
Vespertthonide make more or less frequent use of the interfemoral
membrane as a kind of bag or pouch (Plate VIII., p. 104).
Into this the head is dipped and the struggling captive pressed
against the membrane until it is overcome. The larger the inter-
femoral membrane, the easier is its use as a pouch; and, as if
ready at a moment's notice, its position in flight is generally
with the tip somewhat below the horizontal. The basal portion
is, however, kept taut by the widely separated feet with their
strong spurs or calcaria.

In Natterer’s Bat the interfemoral is weakly developed, is
carried horizontally in flight, and is possibly used infrequently
as a pouch, but in the Horseshoes alone of British bats this
membrane is too small for such a function, and in fact in their
case the tip of the tail is normally carried bent over the back
(Plates XIX. and XX., pp. 228 and 250). They have, however,
an exceptionally developed antebrachial membrane, and when
eating their prey they press it against one of the wings, seldom
withdrawing their head until the meal is finished. The heaps of
refuse which may be found accumulated under their resting

1 Dobson’s Catalogue of Chiroptera, 160. 2 Whitaker, Naturalist, 1907, 418.

24. INTRODUCTION

places, supported by direct observations of captive individuals
by Mr Coward, show that they usually prefer to alight to feed.
The weak interfemoral membrane of the Horseshoes, correlated
as it is with great perfection, but not rapidity, of flight and
remarkable development of nasal sensory organs, is worthy of
more than a passing notice, and is probably, as in the Barba-
stelle, connected with very special habits: in this connection
it should be remembered that the whole sub-order of fruit-eating
bats are unprovided with an interfemoral membrane.

In the whole order the powers of flight’ are distinctly
superior to those of birds, especially, as Mr Hahn observes, in
the power of checking momentum. Their remarkable agility
when hunting, and the hours at which they appear, make bats,
as a rule, safe from the attacks of predatory creatures. But
instances are on record of their pursuit or capture by a stoat,?
or by hawks*® or owls.*| On one occasion a small bat and a
large beetle were observed to fall to the ground together,
having probably come into collision accidentally ;° while Mr
Lionel E. Adams writes me that he once saw a bat fly into a
bicycle. The position of the wings when in action differs, as
Osburn ® pointed out, from that of birds, the arc formed by the
tip scarcely rising above the plane of the body, beneath which
the wings seem to meet in their downward stroke.

The observation of their habits is somewhat complicated
by the fact that sometimes members of a colony may
remain inactive all night. Subject to such exceptions, how-
ever, it is now definitely known that most bats, such as the
Pipistrelle, Daubenton’s, and Long-eared, normally continue
their flight, no doubt with intervals for rest, throughout the
night. Others, represented in Britain by the Noctule and
Leisler’s, take their flying exercise twice a day, contriving to
secure all the food that they need in the course of two headlong
careers of from eighty to one hundred and twenty minutes each,

1 On this subject, see Baron Francis Nopcsa’s “Ideas on the Origin of Flight,”
Proc. Zool. Soc. (London), 19th Feb. 1907, 223-236.

2 George Wolley, Zoologist, 1846, 1204.

3 G. J. Talbot, Fre/d, 10th Oct. 1903, 635; F. J. Montgomery, Fze/d, 26th Sept.
1903, 532. An African hawk is said to feed on bats—see C. J. Andersson’s Birds of
Damara Land, London, 1872, 23, and for the hobby, John Sclater, Zoologist, 1875, 4538 :
in British Columbia the large rainbow trout leaps at and probably catches them—

John Macoun, quoted by E. Thompson Seton, ii., 1181.
4 Zoologist, 1887, 426-427. 6 J. H. Wilmore, Zoologist, 1886, 242. 8 OD. cit.

EDWA

DRAWN BY

RD A. WILSON

BA . M.B. ticaarias')

HG.)

In Preparation.

A NEW AND REVISED EDITION OF
YARRELL, NEWTON, AND SAUNDERS’

HISTORY OF BRITISH BIRDS

EDITED BY

WILLIAM EAGLE CLARKE, F.R.S.E., F.L.S.

Membre Honoraire du Bureau Central Ornithologique Hongrois;

Member of the British Ornithologists’ Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES
SPECIALLY EXECUTED BY

MISS LILIAN MEDLAND

Tae publication of Yarrell’s “History of British Birds” was
commenced in 1837 and completed in 1843. Its outstanding
merits were at once recognised, and a Second Edition was .
called for in 1845, followed by a third in 1856.
From the issue of the Original Edition down to the present
day, Yarrell’s “History of British Birds” has generally and
deservedly been regarded as the standard authority on British
ornithology.

In the year 1871 a Fourth Edition was begun, under the
masterly editorship of Professor Newton—the greatest British
ornithologist of all time. | Unfortunately Professor Newton's
official engagements at the University of Cambridge only allowed
him to complete the first two volumes; and in 1882 Mr Howard
Saunders was selected to edit the remaining volumes, a task

which he successfully accomplished to the entire satisfaction of —

ornithologists in 1885.

The many excellences of this last edition advanced the work
more than ever in the public and in scientific favour. To its
stimulating influence is to be mainly attributed the marvellous and

unprecedented activity which has resulted in those extraordinary
‘advances made in all branches of British ornithology during

recent years—advances which have rendered it essential that a
new work based upon this classical and comprehensive founda-
tion should be issued. ;
During the period alluded to, a considerable number of new
and interesting species have been added to our avifauna. The

[Continued on Page 3 of Cover

arate )
wea ts al WGVivints an
bts Uke ci }
Wn on san ¥
Lin.

Te
i E oe

CONTENTS OF PART IV.

\ GENERAL INTRODUCTION TO BATS
iA y ‘ (To be completed in forty-eight pages.)

| _ VESPERTILIONIDA—
: Genus Myotis—
The Whiskered Bat .
Bechstein’s Bat
Natterer’s Bat
The Notch-eared Bat

The Mouse-eared Bat

Genus Plecotus .

The Long-eared Bat

ILLUSTRATIONS

FULL-PAGE (Coloured and Black and White).

Noctule Bats. (Coloured.)

; Wings of—(1) Myotis mystacinus ; (2) M. bechsteini ; (3) M. natterert.
Heads of—(1) Myotis bechsteint; (2 and 3) Barbastella barbastellus
. (front and side views); (4) Plecotus auritus.

~ Long-eared Bats.

a FIGURES IN TEXT.

"Side View of Skulls and Teeth of—(1) Myctalus leisleri; (2) Myotis
: bechsteini ; (3) M. natterert.

Front View of Incisors and Canines of J/yotes bechstetnt.

Front View of Incisors and Canines of Myotis nattereri.

Diagram of Arrangement of Teeth in Genus P/ecotus.

- Front View of Incisors and Canines of Plecotus aurttus.

ae , aa : “et ihe, ¥ j ie ; ev ,
ih sie Ah =) (s} a ; bt tA ‘os , P
Noa Le heal rare orp a s Te cara ah es . 7)

ae 48

is ve
J )

CHIROPTERA 25

just after sunset and before sunrise. Except when hindered by
stormy weather, their appearances are very regular. Amongst
the earliest to fly are the inappropriately named Serotine and
the Barbastelle, which with the Noctule and Leisler’s, may all
be seen accompanying swallows and swifts on the wing. The
Pipistrelle has been shown to appear in Ireland soon after sunset,
and Daubenton’s Bat, although seldom noticed, is probably also
an early flier. Details regarding the others are wanting. Thus,
although nominally creatures of the night, bats normally appear
abroad long before the onset of darkness, and do not return
home until daylight is far advanced. The times of emergence
and retirement are so adjusted to suit the seasons that, as the
autumn evenings darken, they awake at a progressively earlier
hour, and in the winter, should they fly at all, they often do so in
daylight. Apart from this, midday flights, although probably
due in some instances to disturbance,’ are so frequent that they
can hardly be regarded as extraordinary.” It seems likely,
therefore, that bats can have no inherent objection to light, and
many exotic species* are regularly active in the daytime,
especially those inhabiting shady woods. Again, British bats
are often found asleep in summer in positions where they are
exposed to moderate or even bright light.

The diurnal retreats are no less varied than the nightly
hunting grounds. But here, again, no fixed rule can be rigidly
applied. Most species, besides using the shelter of trees when
on the wing, find a safe refuge in their trunks. Houses, too,
are resorted to where there are convenient cavities ; and caves,
although often so damp that the fur of the bats hibernating in
them is wet, are very much appreciated in winter, probably on
account of their equable temperature.* Indeed, the occurrence of
Horseshoe Bats in any district appears to depend in some degree
on the presence of caves, in which many other kinds also con-

1 See H. W. Newman, Zoologist, 1859, 6317.

* A remarkable instance is related by A. J. Dasent, ze/d, 7th Sept. 1889, 350.

3 Eig, amongst others, Wyctalus azoreum of the Azorean oak woods; Rhyn-
chonycteris naso of Amazonia and Guiana (E. A. Goeldi, /ézs, 1904, 518) ; Lavia frons

of Tropical Africa (P. L. Sclater and J. H. Speke, Proc. Zool. Soc. (London), 8th March
1864, 99).

* Hahn found that for a period of two years’ observations the extreme variation of
temperature in a bat-cave was from about 51° F. in January to 57° F. in September
(op. ctt., 140).

26 INTRODUCTION

gregate, each asa rule keeping to its own area. As stated above,
seven species have been found thus harbouring in Normandy,
and it seems likely that the majority of our British bats resort
at times to caves. On the other hand, there is, so faras I am
aware, no record of the occurrence in British caves at any time of
the Noctule, Leisler’s Bat, or Serotine, while it appears that the
Pipistrelle enters them very rarely. When at rest bats either
hang suspended, head downwards, their feet gripping tightly the
inequalities of wall or ceiling, or they may lie along a slope,
or may wedge themselves into cracks or crevices. One
exotic species, the Flat-headed Bat,’ has the head peculiarly
flattened, as if to enable it to creep into narrow places. In
repose the wings of vespertilionid bats are folded close to
the sides, but almost completely envelop the body in the
Horseshoes. Bats are extremely conservative in their affection
for particular haunts. In Kent’s Hole, Devonshire, where
Montagu caught his Horseshoes, the heap of excrement’ and
bones associated with those of long extinct animals seem to
suggest an immemorial tenancy. Certain situations are again
much favoured, and when a captive bat is introduced into a
room it shows a tendency to suspend itself time after time in
one spot. In spite of this, disturbance by man may cause a
temporary or permanent desertion of a favoured retreat, and
even Kent’s Hole is now little frequented.

The manner of alighting is interesting. The usual method
is to secure a hold with the thumbs, and then shuffle
quickly round into the ordinary position with head downwards.
The Horseshoes are, however, so agile that, turning a somer-
sault in the air, they contrive to get an immediate grip with
their feet. This manceuvre is combined with exceptional power
of obtaining hold, so that these bats can easily suspend them-
selves from such smooth surfaces as a curtain pole, which are
quite impracticable for a Noctule. Only two other species,
Natterer’s and the Barbastelle, have been observed to follow

1 Platymops macmilani of Thomas, Aun. and Mag. Nat. Hist., May 1906, 499-501.

* Osburn, writing of fruit-eating bats in Jamaica (Zoologist, 1859, 6588-89),
mentions deposits “of great extent and many feet in thickness, on the floors of the
caves,” which in one case could not be less than half a mile in length. See also H. L.
Ward’s remarkable account of a Mexican bat-cave in 7vans. Wisconsin Acad. of Sci.,
Acts and Letters, xiv., 11, 634, etc., 1903 (1904).

CHIROPTERA 27

the Horseshoes’ method of alighting, and they only do so occa-
sionally and with but partial success. It is noticeable that all
three have weak thumbs.

Practically all bats are gregarious : even the Whiskered and
Barbastelle, so long supposed to be solitary, have been found in
colonies. In the case of the Noctule, Leisler’s Bat, and the
Serotine, these colonies are not known to include individuals of
other species. The two Horseshoes are moderately exclusive,
but are often encountered in each other’s company in regions
where both occur, and in France the smaller kind is on good
terms with the Notch-eared Bat. Most other small bats do not
object to the presence of an alien, but asa rule each species, and
indeed each individual,’ is segregated, and in hibernation the
most sociable, as the Long-eared and Leisler’s, are often solitary.

In spite, however, of the sociability of the majority, individuals
may often be found living by themselves and frequently changing
their lodgings with the whim of the evening. Itis certain that,
apart from sex, the causes leading to the congregation or the
segregation of bats are still obscure and in need of investigation.

Besides thus changing their domicile, many bats alter their
feeding grounds with the season, since they must perforce
attend the migrations of their prey. Rivers and pools, lanes
and hedges, ivy-covered walls and ruins are all sought at the
times when they most attract insects, a knowledge of the habits
and flights of which is essential to a bat-hunter.? During the
cockchafer flights the Kentish Serotines catch these beetles as
they buzz at low elevations around trees and bushes, but later
in the summer not one will be seen where before they have been
numerous. So the Noctule and Leisler’s Bat vary their hunt-
ing-grounds, and may travel considerable distances from home ;*
although, where food is always abundant, their appearance and
movements are almost as regular as clockwork. Again,
Daubenton’s Bat probably takes a flight in mid-air before

1 In the Mexico bat-cavern each species was disposed “in a band of some feet or
yards in width; then a slight vacant space, followed by a band of another species.”
—Ward, of. cit., 635. 2 F. Norton, Widland Naturalist, 149, 1883.

3 C, Nicolle and C. Comte state that of forty-seven examples of Pipistrellus
kuhliz, marked and liberated at a distance of two kilometres from their retreat, six
were eventually recaptured where originally taken,— Compz. rend. Soc. Biol., Paris, |x.,

738-739, 1906,

28 INTRODUCTION

settling down to the water surface. Others alter their beats
but little throughout the year, but none is more regular than the
Pipistrelle—which may be encountered night after night on the
same restricted course.

On the whole the food appears to consist of such insects as
may happen to be caught; the choice depending, perhaps,
rather on the strata of the air in which insects are flying than
on any preference for particular kinds. The high flight of the
Noctule and its size seem to bring it largely into contact with
the large strong-flying beetles which the Greater Horseshoe
seems to catch before they have risen to high altitudes. The
smaller bats probably rely on flies and small moths for the chief
part of their sustenance. The resort of the Water Bat leads it
to capture caddis-flies, but for the remaining species there is no
observed differentiation. In captivity practically all insects are
accepted except those which are distasteful to insectivorous
animals generally. Messrs E. L. Rollinat and R. Trouessart*
find that whereas the Mouse-eared, Natterer’s, and Daubenton’s
Bats devour cockroaches with avidity, the Serotine, Pipistrelle,
Long-eared, Barbastelle, and Notch-eared show no marked
predilection for them. Bats in general are extremely voracious,
and their powers of consuming insects will be alluded to in
connection with their habits in confinement.

Bats are thirsty creatures and love to lick up water or
milk. In nature they often drink, like swallows,’ on the wing,
and, although there is no reason to suppose that, like the
birds, they seek to wash themselves in the water, they have
been occasionally detected alighting on its surface,’ and on
such occasions have surprised the observer by the agility with
which they rowed themselves along with flapping wings, or at
will resumed their flight. A Long-ear has been observed
settling by some water to drink.

As is well known, bats afford a conspicuous instance of
the phenomenon of hibernation, on which subject the brilliant
researches of the Italian Lazarre Spallanzani,* although under-
taken over one hundred years ago, are still fascinating reading.

1 OD. cit. infra, p. 32. 2 As noticed by Gilbert White.
3 J. G. Millais; C. M. Smith, F2e/d, 20th July 1889, 97; Lord Lilford, Zoo/ogist,
1887, 66 (M/. mystacinus). 4 Op. cit. infra, p. 30.

eOxaifson,
ae

NocTULE BATs. (About 4 natural size.)

CHIROPTERA 20

All British species are liable to enter the lethargic condition
should the temperature fall low enough, but they may rapidly
awake in response to heat, as when warmed in the hand, so that
their lethargy is never very profound. At such times the body-
temperature rises rapidly, sometimes as much as 31° Fahren-
heit in fifteen minutes, the muscles shudder, the animal pants
and throbs, becomes lively, squeaks, and is fully awake. Before
commencing hibernation, bats are, as a rule, very fat, their
weight being appreciably greater than in early summer, at
which time some species are thin. Mr Hahn believes that the
degree of lethargy bears a close relation to the supply of super-
ficial fat, and is not connected with temperature or season. The
American bats examined by him became more active as the
winter advanced, but this is contrary to British observations.
At all times, as in Britain, there was much irregularity of
movement amongst individual bats.

The more general belief is, however, that the duration and
depth of the hibernatory sleep depend largely, although not
entirely, on temperature, and consequently on locality and
season. In warm climates bats, like hedgehogs, probably do
not hibernate at all, while in the temperate regions, such as
Britain, the process is by no means constant, but varies with
the species, for each of which there is no doubt a definite
temperature inducing torpidity. Individuals, however, may
constantly be found abroad at exceptional temperatures, but
the body never becomes so physiologically awake as to ovulate.

Leisler’s Bat is probably one of the earliest to discontinue
flying, and in Ireland, although it has been observed so late as
the 16th November, the majority as a rule disappear about the
end of September. Its congener, the Noctule, has been
observed abroad in every month except January, but its winter
appearances are the exceptions that prove the rule. The Long-
eared is more hardy and is frequently caught in winter: it
probably shifts its quarters at intervals throughout the winter.
The Barbastelle has been taken on New Year's Day. A
Daubenton’s Bat has been found on the wing at a temperature
of 42° Fahrenheit, and another contained recently digested
food in December. Natterer’s leaves its winter quarters in
February, while the Horseshoes, the Whiskered, and the

30 INTRODUCTION

Pipistrelle probably fly whenever the weather is mild throughout
the winter. An evening temperature of 40° to 43° Fahrenheit is
probably required to keep the latter at home, the corresponding
figure for the Long-eared being one or two degrees higher. On
the other hand, it is probably the temperature of midday that
influences Leisler’s Bat in its winter flights. Most authorities
are agreed that bats fly more frequently in the early than in the
later months of winter, by which time probably most insect life
has been killed by frost. In the equable temperature of
churches and caverns they are active although they may not
venture outside, and the delicate Horseshoes probably snatch
irregular meals on the cave-haunting moths and spiders which
share their winter quarters.

North American naturalists have proved that bats perform
regular migrations, but it is not clear that they do so to avoid
hibernation. Of six species found in Manitoba, according to
Mr Ernest Thompson Seton,’ ‘all are migratory and yet
hibernate,” and the seasonal journeys of at least three of them,
the Red, the Hoary, and the Silver-haired, are on a scale
comparable with those of birds. The second of these has
even been known to cross from the American continent to
the Bermudas, a distance of over six hundred miles. The
regular passage of these bats over the Atlantic was invested
with special interest in view of an old record of the occurrence
of one of them in the Orkneys. Unfortunately this turns out
to be an example of a Hawaiian species,’ which cannot possibly
have reached this side of the Atlantic unassisted.

Occasionally bats are reported from lighthouses as if
they were migrating,® but it is regrettable that for Europe
there is on this point not much definite information, although
it is more than a century since Spallanzani wrote that in
Italy most bats are migrants. More recently Bell accepted
Blasius’ belief in the migration of the Northern Bat,° based on

' Life Histories of Northern Animals, ii., 1909, 1161.

2 Nycterts (= Lasiurus) semota.

3 R. M. Barrington, Zhe Migration of Birds, 284: R. H. Porter, London, 1900,
gives records from Fastnet, Rockabill, Blackrock (Mayo) and Tearaght, on the west,
and from Lucifer Shoals and Arklow South on the east coast of Ireland.

* Rapports de Pair avec les étres organisés, ii., 179, édition Jean Senebier, Geneva,
1807. 5 Vespertilio nilssont.

CHIROPTERA a

the fact that no one had ever been able to find hibernating
individuals of this species in extreme northern countries.
Although this bat is believed to breed only in regions lying
approximately between 54° and 58° north latitude, it is observed
with its young, after the breeding season is over, as far north
as 68 or 70. Blasius, therefore, assumed a change of habitat
of at least ten degrees of latitude. It wanders north, however,
only at an advanced season of the year, so that, since rough
weather sets in at the beginning of October, Blasius supposed
that the bats could not remain for more than six weeks before
returning to their winter quarters in the south. Corroboration
of these facts is highly desirable, especially since the supposed
summer migration of the Eastern Pipistrelle* to central Europe
and Sweden turns out to rest on an error of identification, this
bat having been confused with another non-migratory species.”
Similar instances, although in different countries, are on
record. The Euryale Horseshoe® is plentiful in summer in
central France, but Messrs Rollinat and Trouessart in the
course of many years’ bat-hunting have never once detected it
between the months of October and June. If it does not
migrate south to pass the winter, it must retire to the depths of
caverns which are inaccessible to man—a somewhat unlikely
alternative when it is remembered how intermittent is the winter
sleep of bats. Farther south Macpherson * believed that in April
1891 he saw a small flock of large bats migrating through one of
the passes of the Pyrenees, at which point the available evidence
ends. Mention must, however, be made of Mr A. H. Howell’s
suggestion that when migrating bats fly high and by day.’
The breeding habits of bats have been but little studied
by British naturalists, who have recorded merely isolated
observations, from which it would be impossible to piece
together a connected history. Continental zoologists* have,

1 Pipistrellus abramus. 2 P. nathusiz. 3 Rhinolophus euryale.

4 Lakeland, 1892, 1. 5 Infra, p. 223.

6 B. Benecke, Zoologischer Anzeiger, i1., 1879, 304-395 (pipistrellus and auritus) :
Eimer, of. cit., 425-426 (noctula and pipistrellus): E. van Beneden and C. Julin,
Archives de Biologie, i., 1880, 551-571, pls. xxii. and xxiii. (daubentont, dasycneme,
myosotis, mystacinus, nattereri, emarginatus): O. Van Stricht, A natomischer Anzeiger,
xix. Suppl., 1901, 208-210 (noctula, pipistrellus, myosotis): Fries, Ueber die Fort-
pflanzung der enheimischen Chiropteren, Nachrichten, X. Gesellsch. der Wissensch,
Univ. Gottingen, 1879, 295-298, (noctula, pipistrellus, nathusit, abramus, serolinus.

32 INTRODUCTION

fortunately, paid more attention to the subject and have proved
conclusively that in their country the normal breeding season is
autumn. At that season spermatozoa are found numerously
in the uterus of the adult female, and the organs of the male
are also functional. Ovulation is, however, postponed until the
termination of hibernation, during which period the ovaries are
quiescent, but the spermatozoa retain their activity in the
uterus until fertilisation takes place in April. On this point all
authorities are agreed, and the facts, subject to correction as to
details, may be taken as substantiated. An alternative view,
that ovulation and fertilisation may take place in the autumn and
winter, with subsequent postponement of the development of the
embryo, seems to be unsupported by facts.

A point still open to discussion is the occurrence of copula-
tion in spring. Messrs Rollinat and Trouessart, whose observa-
tions, as briefly summarised below, are the most complete in
existence, altogether deny that it takes place even in the case of
a female which has missed autumnal impregnation. They
admit, however, the evident ability of the male, and the fact
remains that acts of courtship have been reported in spring, as
in Wales by Mr T. W. Proger, for the Lesser Horseshoe; his
observations, however, are not definite as to consummation.
Again, in France, Monsieur M. Duval* has noticed such acts
amongst captive bats, but it is possible that they may have been
abortive, and observations on captive animals are rarely of
value on such points. The matter deserves further study, but
since the breeding season may vary with locality and climate,
conflicting reports from observers in different countries are
naturally to be expected.

The work of Messrs Rollinat and Trouessart® is based
partly upon a study of a non-British bat, the Mouse-eared,’ and
myosotis, mystacinus, natterert, bechsteini, auritus, barbastellus, hipposideros): Carl
Vogt, Association francaise pour l’'avancement des Sciences, sess. 10, Alger, 1881,
Compte rendu, 655-662 : thus including all British species except /eds/ert.

1 Etudes sur ?Embryologie des Chiroptéeres, 1, Paris, 1879. For a description of
copulation, which takes place while the animals are at rest, not in flight, see Hahn,
op. cit., 161; Victor Fatio, “ Vertébrés de la Suisse,” i, 23, 1869; Duval, “Sur
Vaccouplement des Chauves-Souris,” in Comptes rendus Soc. Biol., Paris, 23rd Feb.
1895, 135-136.

* “Sur la reproduction des Chauves-Souris,” A/ém. Soc. Zool. de France, ix., 214-
240, 1896, 3 Myotis myosotts.

—— ee

a ‘HISTORY OF ©
BRITISH MAMMALS

GERALD E. H. BARRETT- HAMILTON

B.A.(CANTAB,), M.R.I.A,, F.Z.S.

WITH TWENTY-SEVEN FULL-PAGE PLATES IN COLOUR, FIFTY-FOUR iN
_ BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND
FIFTY SMALLER IZLUSTRATIONS

_ DRAWN BY

“EDWARD A. WILSON

B.A., M,.B. (CANTAB,)

y 214 tol

GURNEY AND JACKSON
PATERNOSTER ROW, LONDON, E.C,
IQI1

In Preparation

A NEW AND REVISED EDITION OF
YARRELL, NEWTON, AND SAUNDERS’

HISTORY OF BRITISH BIRDS —

EDITED BY

WILLIAM EAGLE CLARKE, F.R.S.E., F.LS.

Membre Honoraire du Bureau Central Ornithologique Hongrois;

Member of the British Ornithologists’ Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES H

SPECIALLY EXECUTED BY

MISS LILIAN MEDLAND

HE publication of Yarrell’s * History of British Birds” was BE
commenced in 1837 and completed in 1843. Its outstanding —

merits were at once recognised, and a Second Edition was

called for in 1845, followed by a third in 1856.
From the issue of the Original Edition down to the present

day, Yarrell’s “History of British Birds” has generally and |
deservedly been regarded as the standard authority on British fe

ornithology.

In the year 1871 a Fourth Edition was begun, under the a
masterly editorship of Professor Newton—the greatest British _
ornithologist of all time. | Unfortunately Professor Newton’s ©
official engagements at the University of Cambridge only allowed |
him to complete the first two volumes; and in 1882 Mr Howard

Saunders was selected to edit the remaining volumes, a task

which he successfully accomplished to the entire satisfaction of i

ornithologists in 1885.

During the period alluded to, a considerable number of new
and interesting species have been added to our avifauna. The

[Continued on Page 3 of Cover

CONTENTS OF PART V.

PAGES

GENERAL INTRODUCTION TO BATS 3 : : - 33-40
(70 be completed in forty-eight pages.)

VESPERTILIONIDE—
Genus Pélecotus . ; ‘ ; : ; » | 206
Genus Barbastella : : , : : < poe
The Barbastelle : ; : : é . leon
The Hoary Bat , : ; : : an 222
RHINOLOPHIDA
Genus Rhznolophus F ; : : 4 b. 220%
The Greater Horseshoe Bat : : : , © 225

ILLUSTRATIONS

FULL-PAGE (Coloured and Black and White).

Squirrel. (Coloured.)

Wings of—(1) Plecotus aurttus ; (2) Barbastella barbastellus ; (3) Rhino-
lophus hipposzederos.

Heads of—(1, 2, and 3) Rhznolophus hipposideros,; (4) Rhinolophus
Serrum-equinum.

FIGURES IN TEXT,

Diagram of Arrangement of Teeth in Genus arbastella—(1) Upper
and (2) Lower Jaw.

Front View of Incisors and Canines of Larbastella barbastellus (enlarged
and diagrammatic).

Diagram of Nose-leaf of RAznolophus.

Front View of Incisors and Canines of RAznolophus (enlarged and
diagrammatic).

Diagram of Arrangement of Teeth in Rhznolophus ferrum-equinum—
(1) Upper and (2) Lower Jaw.

CHIROPTERA 33

partly upon the Horseshoes. It is both the most recent and
one of the most complete essays on the subject, and may there-
fore be summarised somewhat fully. More exact details will,
where known, be found under the heading of each particular
species. Meantime, British naturalists have a model life-
history with which they can compare their own observations.
The authors found both sexes of the Mouse-ear living in
company in September. The genital organs of the male are
then highly developed, but the female organs are small and not
remarkable, except that the uterus contains numerous sperma-
tozoa. At this time the female is very fat, and remains so till
after parturition, whereas the male is very thin. No change
was observed in the bats throughout the winter. Towards the
middle or end of March hibernation ends and the active life of
summer is resumed. Ovulation ensues, and, the ovum having
been fertilised, the superfluous spermatozoa are expelled, and
gestation commences—almost invariably in the right horn of the
uterus.. A few days usually intervene between resumption of
activity and fertilisation, so that gestation commences most fre-
quently in the first week of April. After fecundation the females
form separate parties, to which no male is admitted, but in which
non-breeding females are present. These ‘nursing colonies,”
as they may be called, do not break up until the young have
been reared, after which they disperse to their winter quarters.
The date of birth of the young varies a good deal. Thus a
female, captured while still hibernating on the 7th March and
kept warm, became a mother on the night of the 4th May (fifty-
eight days), whereas others taken at later dates brought forth
their young between 28th May and gth June (a minimum of
fifty-eight days if they emerged from hibernation on rst April).
Apparently, then, the commencement of gestation may be
hastened by warmth. Allowing a few days between the termi-
nation of hibernation and ovulation, it would appear to last at
least forty-nine days, or seven weeks. In many cases its com-
mencement is delayed, and females captured on 3rd and 17th
April had not yet ovulated, although containing spermatozoa.

1 Duval, “Etudes sur PEmbryologie des Cheiroptéres,” in Journ, de Vanat. et de
la Phystol., xxxi., 93-160, March to April 1895 ; but the authors once found a Serotine
with an embryo in the left horn.

~
FE
4

34 INTRODUCTION

The authors think that these must have spent the winter in some
cold or windy situation, thus rendering spring revival sluggish.
At all events, the female gatherings increase in size all April.

By 5th June many wild young had made their appearance,
and their growth was very rapid.

The authors ascertained that the female does not pair
until her second autumn, when about seventeen months old.
Sometimes even, although to all appearance sexually mature,
she remains virgin throughout the following summer, appar-
ently until her third autumn. There are thus in the autumn
assemblages three types of female. That of the first year,
being virgin, may be distinguished by the small and undeveloped
mammary and generative organs, the latter never containing
spermatozoa, and by the usual external indications of imma-
turity especially in colour, teeth, and wings. That of the
second year, now breeding for the first time, is more developed,
but the uterus is always smaller than in the adult which has
borne young. The latter has the mamme plainly visible, and
the right horn of the uterus distinctly larger than the left.

When the bats leave their winter quarters, the virgin female
of the first year accompanies the adult of the same sex, but leaves
soon after the birth of the young. That of the second year, on
the other hand, even when herself virgin, never leaves the
company of the breeding females. The existence of the class
of virgin females, both of the first and of the second years,
throughout the spring, although it led Carl Vogt’ to suppose
the necessity for spring copulation, convinced the authors that
no such act takes place at this season.

That there should be no spring copulation is difficult to
understand, since the adult male is in train to breed from
September until April. The state of the organs indicates a
season of rut, and the authors express astonishment that, in
spite of this, they are so long inactive. They suggest that
reabsorption of the spermatozoa may take place in accordance
with the Brown-Séquard theory of internal secretion. Although
the authors express no doubt as to the absence of spring copu-
lation, the conclusion, in view of the condition of the male, is so
surprising as to seem in need of corroboration.

1 Of. cit. supra, p. 32.

CHIROPTERA 35

As with the female, so with the male, there are three
classes, comprising that of the first autumn, which does not breed,
that of the second autumn, which probably breeds but rarely, and
that of the third and subsequent autumns, which is fully adult.

Parturition was observed in a single case. The mother
had previously arranged herself with her head uppermost, and
the young one was born feet first and received in a cradle
formed by the maternal interfemoral membrane. The mother
severed the cord with her teeth. The whole process occupied a
period of slightly over half an hour. Thirty minutes later the
mother resumed her usual position with the head downwards,
and the young one occupied a position under her left wing with
the left nipple in its mouth. There it remained for eleven days,
after which it passed its time partly hanging by itself, partly
under the maternal wing.

The young Mouse-ear at birth, despite the fact that the
wings are of late development in the embryo,’ has an expanse of
130 to 160 mm. It is almost naked, its colour blackish or
brownish above, but unpigmented beneath, the wings slightly
pigmented. The eyes are closed, the ears more or less reflexed.
The limbs are well developed, especially the claws, both of the
thumbs and feet, which are sharp and curved. The hooked and
trilobed teeth of the milk dentition arm the front of the jaws.

It is with the claws that a young bat clings to its mother’s
fur, holding on also by its teeth to the nipples, of which there
are a pair, specially long corrugated structures, placed pectorally.*
So firm is the grip that a very young bat taken in the hand will
attach itself so firmly to the fingers as to be separated only by
vigorous shaking. The mothers have at first no difficulty in
carrying their young, which cling.to them with the hind feet
buried in the fur of the abdomen, the thumbs clutching the
breast and neck, a nipple firmly fixed in the mouth. Some-
times during flight the young one bends downwards so that

1 For an account of some embryos of bats, see Harrison Allen’s paper in Con-
tributions from the Zoological Laboratory of the University of Pennsylvania, vol. i.,
No. 2, 1895.

2 Dobson (Catalogue of Chiroptera, 79 and 83) observed that in the males of some
fruit-eating bats the nipples are as large as in any female during lactation, and
suggested that, where two young are born at the same time, the male may relieve the
female of the charge of one.

30 INTRODUCTION

daylight may be seen between its body and that of its
mother. |

The young grow rapidly, soon acquire hair, and are always
strong and vigorous. Their eyes open between the fourth and
ninth day,’ most usually on the fifth or sixth. Between the
ninth and thirteenth days they begin to leave their mothers
and divide their time between hanging by themselves and
attachment to the nipple under the cover of the maternal wing.
Probably after the second week the wild female leaves her young
one at home when she flies abroad to feed, but takes it under
her wing on returning. In fact, Mr Whitaker found that a
young Pipistrelle soon became a burden which was carried in
flight only with difficulty, and seriously impeded the feeding of
its mother on a flat surface, especially when she attempted to
“pouch” her food.

On the twelfth day a young Mouse-ear had attained an
expanse of nearly 250 mm., and was well covered with hair.
The permanent teeth were already replacing those of the
milk dentition. A few days later it no longer sought its
mother’s wing, even when being suckled, but stretched itself
frequently and expanded its wings.

By the thirtieth to thirty-fifth day the milk dentition had
disappeared, but the young one was not finally weaned until it
was about two months old. In the wild parties the young were
nearly as big as their dams in July, and the nursing colonies
broke up before the end of August. Other species probably vary
in these respects, since wild Notched-ears” were found to have
finished lactation on 26th August, whereas young Euryale Horse-
shoes, nearly equalling the adults in size, but with stomachs full
of milk, were on the wing only on the 18th of the same month.

Some naturalists have stated their belief that the females
suckle the young promiscuously,’ but, except in the case of the
Lesser Horseshoe, Messrs Rollinat and Trouessart, after many
years of research and experiment, find no evidence of this. Mr
Whitaker’s report on two Pipistrelles is similar.

The above observations may probably be regarded as

' Whitaker gives the eighth day for the Pipistrelle and twelfth for the Noctule.
* Myotis emarginatus.
* E.g., Osburn, in the case of a captive Monophyllus, op. cit., 84.

)

1 natural size

(About

}
4
4
a]

CHIROPTERA 37

typical for British vespertilionid bats. The Horseshoes differ
from them in certain important respects, and these have been
noted under each species. The period of gestation is longer,
extending to ten or twelve weeks ; the young at birth have the
upper surface covered with down; and they hold on to their
dams by means of a special pair of false nipples situated in the
inguinal region.

Differences of habit will also no doubt be met with in other
vespertilionid species. For instance, the number of young at a
birth is always restricted to one in the Mouse-ear, as it appears
to be also in every British species of this genus, as well as
in the Serotine and the Long-eared.'. The members of the
genera WVyctalus and Pzpestrellus, however, present a curious
problem, since, although the occurrence of two young at a
birth is almost unknown in Britain, continental naturalists
frequently credit them with that number, although not at every
birth. On this point they are quite definite, and there is nothing
inherently improbable in the statement, since in many exotic
bats’ twins or more are usual, and these are sometimes carried
about by the mother even when they have reached about two-
thirds of her size.” Confirmation in the case of the genus
Pipistrellus comes from America, where Mr Vernon Bailey *®
finds that the Little Canyon Bat* may have either one or two
embryos. The facts being as above stated—and, although
further investigation is desirable, there can, I think, be no reason
to doubt them—we have here a very interesting case of varia-
tion in fertility according to locality. My friend Mr Moffat
points out® that the lower degree of fertility occurs on
the outer limits of the ranges of these bats. He compares it
with somewhat similar facts in British and Irish birds, which

' For Barbasiella alone | have no information.

* See W. H. Hudson for Molossus bonariensis in The Naturalist in La Plata, 1o1-
104: Chapman and Hall, Limited, London, 1892 The Red Bat of N. America has
four mamme, and one weighing 11 grammes has been taken alive nursing four
young, weighing collectively 12-7 grammes (M. W. Lyon, jun., Proc. U.S. Nat. Mus.,
XXVi., 425-426, 1903).

8 North American Fauna, No. xxv., 210, 1905.

4 P. hesperus (H. Allen); P. abvamus of Java has also been found with two
embryos (see G. M. Allen, Bul/. Mus. Comparative Zool., Harvard College, Lit 3
July 1908, 45); and P. subflavus of N. America with three (Hahn, of. ct, 162).

® “The Problems of an Island Fauna,” /r7sh Naturalist, 1907, 141.

38 INTRODUCTION

are said to lay fewer eggs than on the continent, and in
Ireland than in England. Certain butterflies, also, produce
a single brood in Ireland as against two in England, while
in others the number of males is excessive.’ All the
facts lead him to suggest that diminished fertility occurs as
an exterminating factor at the outskirts of an animal’s range.

As regards other details, there is evidence that seasonal
change of domicile and the formation of nursing colonies obtains
in every British species of which we possess records in any
detail. Especially is this the case with the crowded cave
parties which only gather in the gloom of subterranean retreats
when the chilly blasts of winter drive them from the slighter
shelters of summer.” In spring the caves are deserted and their
occupants betake themselves to holes in buildings and trees.
Our knowledge is most complete in the case of the Noctule,
which in spring forsakes the sheltering house-roof wherein it
has lain dormant throughout the winter, and forms, usually in
hollow trees, nursing colonies, which keep together until the
autumn. Mr Symington Grieve’s description of the Water
Bat at Loch Dochart suggests similar habits. In the case of
the Noctule, segregation of the sexes is a general but not
invariable rule. The Lesser Horseshoes are as regular, but the
females of this species tolerate the males and frequently allow
them, both young and old, to associate with them. Less
clearly understood, and evidently more complicated, are the
summer ‘“‘swarms” of Long-ears which for a few weeks—some-
times only days—of July or August, cluster in outhouses, where
they are unknown throughout the rest of the year. These
gatherings, as well as those of the Pipistrelle, include bats of
all ages and both sexes.

The date of birth is subject to much variation: of nine
instances covering three species given by Mr Whitaker, the
earliest birth took place on 22nd June and the latest would
probably have occurred in August. Again, in south Wales Mr
Proger finds that the young of the Lesser Horseshoe and of

1 If, as has been suggested, there is a preponderance of males in British Lesser
Horseshoes, and that in Germany, as stated by Kuhl, there may be two young, a
thing unknown in France or Britain, that would be another case to the point.

* For Wales, see Proger, Proc. Cardiff Nat. Soc., March 1905, reprint, 4.

CHIROPTERA 39

Natterer’s Bat are not born until about the last week of June.
On the other hand, Mr Coward had a young Water Bat with a
forearm of only 20 mm., and a total expanse of only 110 as against
220 in the adult, taken on the wing on 28th June, and on the
same day a female which appeared to have recently given birth,
while I have examined a young Pipistrelle and Whiskered, fully
grown but immature, on 9th August and 26th July respectively.

As stated above, the breeding habits of the various species
probably vary with the degree of torpor which they undergo
during hibernation and the climate of the locality in which they
exist. The most simple are those of bats like Leisler’s, which
hardly fly at all during the winter. Others which, like the
Pipistrelle, fly at intervals throughout the year, probably have
a lengthened breeding season and may copulate frequently.
But even if they do, Messrs Rollinat and Trouessart find that
there is no fecundation until the spring.

The facts as observed in Britain seem difficult to reconcile,
unless either copulation takes place in spring, or the commence-
ment of gestation is retarded by a damp climate, just as it is
hastened by warmth.

The birth of bats in captivity has been independently
observed in England by Daniell and Mr Whitaker for the
Noctule, and by the latter naturalist for the Pipistrelle. These
observers were only able to approximate the period of gesta-
tion at not less than thirty-eight days for the former and
about forty-nine days for the latter species, so that as far as
they go they do not disagree with Messrs Rollinat and
Trouessart. They find, however, that the young may be born
head first. In one case the mother hung in the normal position
head downwards '; in the other this position was reversed.

Many naturalists, from Spallanzani downwards, have referred
to a supposed preponderance in numbers of the female sex. On
this point no satisfactory evidence is forthcoming, but the
apparently superfluous size and capacity of the male generative
organs would be accounted for if it could be shown that bats
are polygamous.

1 Sir H. A. Blake (Sez. Proc. Roy. Dublin Soc., 1V., N.S., 449-450, 1885) observed
the birth of an Indian fruit-bat, in this case also feet first. Professor Robert Collett
writes me that he has observed parturition once each in Daubenton’s and Nilsson’s
Bats. The mother in each case hung head downwards.

40 INTRODUCTION

It is perhaps difficult to account for the prejudices which
have always existed against these harmless and interesting little
animals, which are not only objects of superstitious dread to the
ignorant, but have proved to the poet and the painter a fertile
source of images of gloom and terror. That the ancient Greek
and Roman poets, furnished with exaggerated accounts of the
animals infesting the remote regions with which their commerce
or their conquests had made them acquainted, should have
caught eagerly at those marvellous stories and descriptions, and
rendered them subservient to their fabulous but highly imagina-
tive mythology, is not wonderful. It is, indeed, more than
probable that some of the Indian species of bats, with their pre-
datory habits, their multitudinous numbers, their obscure and
mysterious retreats, and the strange combination of the char-
acter of beast and bird which they were believed to possess, gave
to Virgil the idea, which he has so poetically worked out, of the
harpies which fell upon the hastily-spread tables of his hero
and his companions, and polluted, whilst they devoured, the
feast from which they had driven the affrighted guests. But
that the little harmless bats of our own climate, whose habits
are at once so innocent and so amusing, and whose time of
appearance and activity is that when everything around would
lead the mind to tranquillity and peace, should be forced into
scenes of mystery and horror, as an almost essential feature in
the picture, is an anomaly which cannot be so easily explained.

They have, however, been connected with various evil deeds
as foreign to their natures as sky from earth, with blood-
sucking,’ with maliciously entangling themselves in women’s
hair, with thefts of meat and bacon, and with mysterious
entombments in impossible places.” The “wool of bat” was
included by the witches of ‘‘Macbeth” amongst the in-
gredients used in preparing a charm,*® and Tennyson’s “black
bat night” was surely not used in a complimentary sense.

Sense organs:—No one who sees a bat on the wing in
pursuit of its ordinary vocation can doubt that these animals

1 Article on Horseshoes, z7/ra.
2 Zoologist, 1897,46; T. P. Bartlett, Journ. cit, 1844, 613 ; A.C. Smith, Journ. czt.,

1854, 4245.
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In Preparation

A NEW AND REVISED EDITION OF
YARRELL, NEWTON, AND SAUNDERS’

HISTORY OF BRITISH BIRDS

EDITED BY

WILLIAM EAGLE CLARKE, F.R.S.E., F.LS.

Keeper of the Natural History Department, The Royal Scottish Museumj; Member of the
British Association Committee on the Migration of Birds as Observed on the British
and Irish Coasts; Corresponding Fellow of the American Ornithologists’ Union;
Correspondirender Mitglied des Ornithologischen Vereins in Wien;

Membre Honoraire du Bureau Central Ornithologique Hongrois;

Member of the British Ornithologists’ Union, etc.

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES
SPECIALLY EXECUTED BY

MISS LILIAN MEDLAND

CaS publication of Yarrell’s “History of British Birds” was
commenced in 1837 and completed in 1843. Its outstanding
merits were at once recognised, and a Second Edition was
called for in 1845, followed by a third in 1856. .
From the issue of the Original Edition down to the Bosca
day, Yarrell’s ‘‘History of British Birds” has generally and
deservedly been regarded — as the standard authority on British i
ornithology. Hi
In the year 1871 a Fourth Edition was begun, under the
masterly editorship of Professor Newton—the greatest British
ornithologist of all time. Unfortunately Professor Newton’s
official engagements at the University of Cambridge only allowed
him to complete the first two volumes; and in 1882 Mr Howard
Saunders was selected to edit the remaining volumes, a task —
which he successfully accomplished to the entire satisfaction of —
ornithologists in 1885. ‘
The many excellences of this last edition advanced the work
more than ever in the public and in scientific favour. To its
stimulating influence is to be mainly attributed the marvellous and
unprecedented activity which has resulted in those extraordinary
advances made in all branches of British ornithology during —
recent years—advances which have rendered it essential that a —
new work based upon this classical and Forint founda-
tion should be issued.
During the period alluded to, a considerable number of new. j
and interesting species have been added to our avifauna. The

[Continued on Page 3 of Cover

con eeNTs OF PART VI.

GENERAL INTRODUCTION TO BATS (completion) ; . 41-48

RHINOLOPHIDZ—

Genus Rkznolophus—
The Lesser Horseshoe Bat . ; ‘ . 250

NoTe.—The Section on Bats being now complete, will—witth a General
Introduction to the whole work, Titles, Indexes, etc., to be published
later—form Volume J.

VOLUME II.—Lanp Mammnats

ORDER INSECTIVORA
TALPIDZ " é : : : ‘ , ‘ 2

Genus 7alpa . : ‘ : ; : ,
The Common Mole, Moldwarp or Want . : ; 5

The English local names have been revised in part by Mr W. W.
Skeat, M.A. (assisted by Professor W. W. Skeat), and in part by
Mr C. M. Drennan, M.A. Lond., late Scholar Emm. Coll. Camb. ;
the Celtic and Gaelic names by Dr E. S. Quiggin, M.A., Ph.D.,
Fellow and Lecturer in Modern Languages and Celtic of Gonville
and Caius College, Cambridge; while a list of Scottish Gaelic
names have been supplied by Mr C. H. Alston. Valuable
asststance has been rendered by Mr M. C. A. Hinton regarding
extinct Mammals.

ILLUSTRATIONS

FULL-PAGE (Slack and White).

Barbastelle Bats (on coated paper).
Greater Horseshoe Bats (do.).
Lesser Horseshoe Bats

FIGURES IN TEXT.

Diagram of Teeth of Rhznolophus hipposideros—(1) Upper and (2) Lower
Jaw.

Side View of Teeth of Zalpa Europea.
Encampment of Mole.
Mole Excavations (18 Diagrams).

“ansorian Inst;
a ene? an fac
MAY 27 1911

Mefionat ee :

CHIROPTERA 4l

possess a marvellous faculty for finding their way about under
conditions which would render human beings helpless. A
good illustration of this power is described by Professor C.
Lloyd Morgan," quoting Miss Caroline Bolton, who was present
during an experiment in which threads were fastened crossing
each other in all directions at intervals of about sixteen inches
in a room measuring twenty feet by sixteen. To each
thread a bell was attached in such a way that the slightest
touch would make it ring. Into this room a large bat was
liberated in absolute darkness, but, although the observers could
hear the animal flying about for half an hour no bell was rung,

So keen are their perceptions that, unlike birds, bats usually
perceive windows,’ and if they fly against them, it is not with
the blind dash of a bird. It has been noticed of a Lesser
Horseshoe that it persistently flew at a large mirror, ‘and
though it never actually touched it, it hovered in front of it in
such a way as to indicate clearly that it was in some way
deceived by it.”

All authorities are agreed that bats are in no apparent way
incommoded by the partial destruction of their eyes; but, as
regards the loss of power consequent upon the loss of other
organs, the older writers differed somewhat, and it seems quite
possible that the mere shock or pain caused by the operations
of the experimenters may have in some cases interfered with
the victim’s movements. The element of disagreement is parti-
ally lessened and explained by more recent investigators,* who,
after an examination of several species, conclude that the power
which enables bats to move with certainty in the most complete
obscurity, is not absolutely located in any single organ, but
arises from a combination of senses resulting from several
organs acting in unison and mutually assisting each other.
The most important of these, in the order named, are: (1) hear-
ing, or rather, as Mr Hahn puts it, a sense or senses located
in the internal ear ; (2) touch, specially distributed in the naked

1 Animal Life and Intelligence, 1890-91, 247.

2 Not always—see G. H. Caton Haigh on Daubenton’s Bat, Zoologist, 1889, 434 ;
also Osburn; and Hahn, whose experiments led him to regard the ordinary esti-
mates of a bat’s skill in avoiding obstacles as exaggerated (of. cit., supra, vide p. 3).

3 Messrs Rollinat and Trouessart, Comptes rendus Soc. Biol., Paris, 23rd June
1904, I-4 (reprint).

42 INTRODUCTION

parts and membranes; (3) sight; (4) smell;* and (5) taste.
This explanation is so apparently reasonable and in accord-
ance with probability, that it is to be hoped that these unfortu-
nate creatures will escape the pain of further investigations.

Without resorting to the cruel artifices of continental?
zoologists, an English naturalist, Mr Whitaker, has been able
to corroborate their work, and it may now be regarded as
almost certain that a vast expansion of the most exquisite
sense of touch over the whole surface of the wing, plays a most
important part in making nocturnal flight possible. In
addition, therefore, to their more obvious uses, the flying
membranes,’ and, it should be added, the ear, with the hairs
which scantily cover these parts, almost certainly function as
organs of perception. It seems that Cuvier was not far wrong
when he wrote that ‘‘it is by means of the pulsations of the
wings on the air that the propinquity of solid bodies is perceived,
by the manner in which air reacts upon their surface.”
Whether or no the numerous striations which traverse the wings
and the interfemoral membrane have anything to do with this
power is a matter for future study.

The variable development of the ear and tragus in the
different genera and species is indicative of corresponding
variations in use. The broad, short ear of Myctalus and
Pipistrellus is accompanied by extensive sweat or oil glands *
on the sides of the muzzle, which are connected with the highly
developed nasal branches of the fifth nerve. In Vesfertzfo and
Myotis the ear and tragus are larger, but the glands are less
extensively developed. ardastella and Plecotus, on the other
hand, present a very remarkable extreme, both of ear and
glandular development. Lastly, in Rhznxolophus we see a simple
ear without a tragus, but with well-developed antitragus, large

1 In the families Vespertiliontde and Rhinolophide the ethmoid bones depart so
widely from the plan usual in mammals that a strong development of the sense of
smell is suggested ; see Harrison Allen, Bud/. Mus. Comparative Zoology (Harvard,
U.S.A.), x., 3, 160, 1882-83.

* Spallanzani, De Jurine, Spadone, etc. See Edward Newman, Fe/d, 20th Dec.
1873, 628; J. McIntosh, Zoologist, 1850, 2814.

* The wing is richly supplied with blood, the circulation of which is assisted by
rhythmically contractile, valvular veins, as described by Jones; and Leydig; see
also for its structure, J. Schébl, Archiv fiir Mtkrosk. Anat., Bd. vii., 1, pl. i.-iv., 1870.

* Hahn suggests that these “fatty pads” may have a protective value in prevent-
ing injury to the animal’s head when it strikes an obstacle (of. ct, 176).

CHIROPTERA 43

auditory bulla, and a special nasal organ of sense, the “ horse-
shoe.” In all species the whiskers or vibrissz are prominent.

Although it is difficult to explain the movements of bats by
means of the five human senses, there is no proof of the exist-
ence of a sixth. Such a sense, if existing, might be one of
direction, and might be located in the inner ear.’ It must be
remembered also that different senses may predominate in the
various species, for it may be naturally supposed that the acute-
ness of each particular sense may vary in relation to the size, food,
and general habits of each. In those bats, such as the Noctule
and Leisler’s, which fly in the subdued daylight of morning and
evening rather than in the dark, the power of sight is probably con-
siderable, as it may be also in exotic day-flying species. Indeed,
the same authority, Mr Whitaker, who showed that in Natterer’s
Bat the eyes are not necessary for the performance of all normal
movements, believes that the Noctule habitually hunts by eye-
sight; and that the Long-eared Bat relies largely on hearing
and sight, especially the latter. On the other hand, Mr Bruce
F. Cummings finds the sense of sight weak, but that of hearing
very acute in the Greater Horseshoe.

As regards hearing, it is probable that this sense is
frequently useful as a means of communication. Its importance
to bats is indicated not only by the variable and remarkable
development of the ear, but by the great mobility and independ-
ence of this organ. It is indeed inconceivable that creatures
which, like some bats, are rarely silent should have no percep-
tion of the voices of their fellows ; and indeed a young bat calls
for its mother as persistently as any other young animal.
Most experiments on the hearing of bats are subjected to the
difficulty that, the cries of these animals being pitched on a
very high scale, their auditory organs are almost certainly
attuned accordingly. It is a well-known fact that their ordinary
cries, although very perceptible to some human ears, are to
others quite inaudible, especially to those of people of mature
age. The gamut is, in fact, fixed above that to which human
ears are attuned.

A captive Noctule, when it had become accustomed to its
novel surroundings, took no notice of such (to human ears) loud

1 Hahn, of. czz., 191.

a4 INTRODUCTION

noises as thunder, but always started at the tearing of paper,
and never ignored any sound approaching a chirrup or a click.
At the latter sounds it would invariably awake from sleep, and
they were always used to attract it to its food. High musical
notes invariably attracted it, whereas low ones, however loud,
had no perceptible effect.

According to Herr Herbert Elias,’ the shrillness of the cries
of bats has an intimate connection with the structure of the
larynx. The musculature is very powerful and the glottis short,
the latter being the main cause of the shrill cries.

Bats of many kinds have frequently been kept in captivity,
and with varying success. Spallanzani, for instance, reared
young ones on goat’s milk. Few species show any naturally
exaggerated fear of man when captured, and the majority
rapidly accustom themselves to their new conditions ; so much
so indeed, that when well fed they become lazy and slow to take
wing. The Horseshoes, however, were regarded as intractable,
and had never been kept alive for any length of time, until Mr
Coward, after three weeks’ coaxing, induced them to feed satis-
factorily. Others, as the Long-eared, the Noctule, Leisler’s Bat,
and the Pipistrelle—one of which Mrs S. C. Hall? is said to
have preserved in health for over two years—and, as shown
above, the Mouse-ear of continental Europe, are easily kept in
health, have in some cases given birth to their young, and
prove pets of considerable interest. In warm weather their
appetite is prodigious, and an adult Mouse-ear, according to
Messrs Rollinat and Trouessart,’ is capable of devouring a
thousand house-flies in a night and fifteen hundred in the night
following : sixty-seven large grasshoppers were eaten at a meal,
and eighty more followed during the ensuing night. The diffi-
culty of supplying a number of captives with food was overcome
by these authors by the help of the common cockroach, with
the result that the bats brought forth and reared their young
in perfect health, and after affording materials for the interest-
ing observations already summarised, were released and took

1 Gegenbaur’s Morphologisches Jahrbuch, xxxvii., i., 70-119, 1907, with plates.

* Vide Newman, Field, 27th Dec. 1873, 656.

3 These authors ingeniously moderated the appetite of their captives in winter by
keeping them out of doors in a temperature so low as to induce torpidity.

CHIROPTERA 45

wing—both young and old—none the worse for their captivity.
Other species appear to be no less voracious; thus Dobson!
mentions a fruit-bat which consumed food of twice its own
weight in three hours.

The behaviour of bats when in captivity is in many ways of
much interest, their actions being much influenced by their
habits while in a state of nature. A bat rarely thinks of look-
ing for a mealworm, even if it be struggling close under it on a
table, thus suggesting the inference that an insect missed is an
insect lost; but it will eat one if it accidentally touches its
mouth or head. When first offered moths or mealworms in
captivity, the various species behave very differently. Some,
such as the Noctule, Leisler’s, or the Pipistrelle, have no hesi-
tation in feeding, but others, evidently not expecting to eat
while resting, seem at first incapable of grasping the situation
and appreciating the meaning of the objects offered to them.”
The Whiskered Bat is at such times a particularly shy feeder,
and appears to gain no information whatever from the sight or
smell of a mealworm, although Mr Oldham found that the attach-
ment of a pair of moth’s wings to the worm has driven the lesson
home. Even the Pipistrelle, which readily accustoms itself to
captivity, has been known to insist on the unnecessary incon-
venience of pouching a small insect eaten while resting on a flat
surface.

There is nothing to guide us as to the possible age attained
by bats, except the statement quoted above that a captive
Pipistrelle lived in the possession of Mrs Hall for two years.
From the known facts of their breeding habits it may, however,
be argued that in Britain bats must, if the numbers of their
species are to be kept up, reach a minimum of at least four
years. The production and rearing of three young ones by a
pair of bats (the first born when the parents are two or even
three years old) occupies, if the number at a birth be but one, a
period of about three years, and it is very doubtful if such a
number would be sufficient to compensate for the admittedly
large death-rate existing amongst wild animals. Where two at
a birth is the rule, the period in question should be sufficient to

! Catalogue of Chiroptera, 83. a ; ;
2 The Long-eared is particularly easy to keep in captivity, since it readily searches
for and seizes its food.

46 INTRODUCTION

maintain the numbers, and these facts form a valuable com-
mentary on Mr Moffat’s views on the extermination by lack of
fertility at the outskirts of the geographical range of a species.

The following key to the characters of British bats will, it is
hoped, prove useful :—

AY) KEY VO. BRITISH * BA Ls;

I EXTERNAL CHARACTERS:

I. WirHout Noss-Lear :—
1. Lars separated—
(A) External ear-border terminating under the angle of the mouth—
(A) Post-calcarial lobe prominent—
(a’) Tragus broadest atapex. MYCTALUS pitaee

(a) Size larger, forearm | NN. noctula . + ‘perese
rarely under 49 mm.. f

(2) Size smaller, forearm | LV. leisierz . p. 0s
rarely over 44mm. . f
( PIPISTRELLUS \

(b’) Tragus broadest at centre \ papistrellus ye 103
{ VESPERTILIO \

(B) Post-calcariallobeinsignificant \ eerie j? 130
(B) External ear-border terminating |
under the base of the tragus J MYOTIS Peta

(A) Posterior margin of inter-
femoral membrane not
fringed—
(a’) Ears moderate ; when laid
forward not extending
beyond nose-tip—
(a) Calcar extending three- |
quarters way from ; . daubentont pte
ankle to tail 14
(8) Calcar extending half- )
way from ankle to tail f
(b’) Ears long ; when laid for- |
ward extending far be- ; WW. bechsteinz . >. Pony
yond nose-tip : |
B) Posterior margin of inter- | , Bae
e femoral eorre fringed . \ I AGRI a
2. Ears united at base—
(A) Ears very long, much longerthan f PLECOTUS \ p

M. mystactnus . / pe ase

head (about 34-38 mm.) . a auritus Hees ot

(B) Ears of moderate ican (about ( BARBASTELLA 5

13-16 mm.) : Fit barbastellus } ore

a. thes NosE-LEAF . : : . RAINOLOPAUS jp. 225
. Size larger, forearm 50-55 mm. . . RR. ferrum-equinum p. 228

2. Size smaller, forearm 34-39 mm. . LR. hipposideros . p. 250

CHIROPTERA 47

i tee AND SKULES:

J. NASAL CARTILAGES WELL DEVELOPED :—

1. Hour functional teeth in tooth-row behind upper canine—

(A) No rudimentary cheek-teeth { VESPERTILIO \ Pp

1 serotinus J

(B) A minute tooth internal to tooth-

row between canine and _ first
grinder.

(A) First lower grinder minute, its }
tip very much lower than
summit of third; upper sur-
face of rostrum concave, and
facial profile descending ab-
ruptly to nasal region

(B) First lower grinder large, almost

| BARBASTELLA\ :
barbastellus J p-. 212

equal in height to third ; upper

surface of rostrum convex, and

cranial profile a gradual slope

(a’) Outer upper incisors al-
most double inners in
section ; greatest length
of skull about 19 mm.

(b’) Outer upper incisors aL
most equal to inners in Ses
section ; greatest length Drea
of skull ‘about I5 mm.

2. Five functional teeth in tooth-row behead upper canine—
(A) Five teeth in tooth-row behind ( P7PJSTRELLUS \

NYCTALUS ) purges

~LV. noctula. « pk ags

lower canine. ra poypistrellus J ae
(B) Six teeth in tooth-row peti {, PLECOTOS * :
lower canine. Bal auritus J Bs

3. Sex functional teeth in paeteram ee upper canine—
(A) Upper pm. II. smaller, its tip about
reaching cingulum of fm. I. :—
(a) Size larger, greatest length of |
skull about 15 mm. im |
(B) Size smaller, greatest length of
skull about 13 mm.
(B) Upper fm. II. larger, its oe
reaching well above cingulum

M. daubentont . proms

\ M. mystacinus <) Pearse

of pm. I. :—
(A) Beet. II. about equal- 1 M. nattereri . :) ps. 178
(8) ee: I. distinetly s smaller ‘ M1. bechsteinz . / prays

II. Nasa CarRTILAGES AND UPPER INCISORS 4 RHINOLOPHUS
RUDIMENTARY

I. Greatest length of ull ae 24) mm. a Jerrum-equinum p. 228
2. Greatest length of skullabout 16mm.. &. Aipposideros . p. 250

-
ts
to
vat

VESPERTILIONID-.
TYPICAL INSECT-EATING BATS.

THESE are bats without a nose-leaf; with a distinct tragus ;
with a long tail and large interfemoral membrane; without
palatal processes to the premaxille, so that the bony palate is
defective anteriorly and the upper incisors are divided in their
midst by a wide space; without expansion of the nasal
bones.

The arrangement and classification of the Vespertilionidze
followed in the present volume differs widely from that of Bell.
The changes may be indicated by placing the two systems side
by side :—

Bells system of 1874 :— System of present work :—
SCOTOPHILUS NOCTULA becomes NYCTALUS NOCTULA.
S. LEISLERI x, N. LEISLERI.
S. PIPISTRELLUS * PIPISTRELLUS PIPISTRELLUS.
S. SEROTINUS 3 VESPERTILIO SEROTINUS.

VESPERTILIO BECHSTEINII
V. NATTERERI

- MYOTIS BECHSTEINI.
5 M. NATTERERI.

V. DAUBENTONII i. M. DAUBENTONI.
V. MYSTACINUS x M. MYSTACINUS.
PLECOTUS AURITUS remains P. AURITUS.

BARBASTELLUS DAUBENTONII becomes B. BARBASTELLA.

From the above list two doubtfully British species have been
omitted : the specific names of the remainder will be discussed
under their own headings. As regards the generic names,
Scotophilus was based by Leach (Zvaus. Linnean Soc., London,
xili. 71-72, 1822) on S. &uhlz, a species quite different from
any of those mentioned above. It was, therefore, employed in
error by Bell. Vesferugo of Keyserling and Blasius, a substi-
tute for Scotophilus, has been shown by Mr G. S. Miller to be

inadmissible in any sense in zoology (see under genus Vyctalus).
49 G

50 VESPERTILIONID£

In default, then, of the present arrangement, it is entirely open
to question what generic term could be used, and in any case
the trend of modern opinion is strongly against the retention
in one genus of two such dissimilar animals as sevotzmus and
noctula. If the separation of xoctula and J/ezslert from true
pipristrellus be refused, the generic name Vyc¢ta/us must, by the
laws of priority, be applied to all three; but this question will
be treated below. The transference of the name Vespertzlio
from the group of bats with thirty-eight teeth, as used by Bell,
to that with thirty-two, is due, as is the rest of the present
system, to the researches of Mr Miller, who showed (Aux. and
Mag. Nat. Hist., October 1897, 379-385) that much of our
previous classification and nomenclature was based either upon
laxity or upon intentional disregard of the laws of priority.
Mr Miller remarks that Linnzeus’s genus Vespertztio (Systema
Nature, \., X., 31-32, 1758), included seven species—vampyrus,
spectrum, perspicillatus, spasma, leporinus, auritus, and murinus,
only two of which, aurztus and murinus, are European. As
a non-exotic species illustrating most closely the original
meaning of the author should be retained as the type of the
genus, one of these two must be selected. The species
auritus was removed to the genus //ecotus by Geoffroy in 1818
(Description des Mammiferes...en Egypte,112). Thus murinus
is left as the type of the genus Vespertitio. The true Vespertilio
murinus, however, is a totally different animal from the one
formerly known by that name. To understand the matter
fully it is necessary to refer to the two editions of the Fauna
Suecica, in the first of which Linnzus mentions only one bat,
the ‘‘ Laderlapp,” ‘‘ Fladermus,” or ‘‘ Nattblacka.” This he calls
“Vespertilio caudatus, naso oregue simplict” (No. 18, p. 7, 1746).
In the second edition two species are mentioned, No. 18 of
the first edition (here numbered 2) and No. 3, the Long-eared
Bat ‘Vespertilio aurttus naso oreque simplicr, aurrcults duplicates
capite majoribus’ (pp. 1-2, 1761). These had already received
binomial names, Vesfertzlio murimus and V. auritus respec-
tively, in the tenth edition of the Systema Nature, where the
following diagnosis of V. murvinus is given: ‘‘V. caudatus, naso
oreque simplicz, auriculis capite minoribus” (p. 32, 1758). In
the second edition of the Fauna Suecica the teeth of V.

VESPERTILIONID/E 51

murinus are thus described :'—‘‘ Dentes primores superiores
6, acuti, distantes. inferiores 4. acuti contigui. Laniarii
superiores 2. anteriore majore. inferiores 3. antico maximo.
Molares utrinque 3. tricuspidati.”

It thus appears that the Vesfertilio murinus of Linnzus,
the type of the genus Vespertzlzo, is a common Skandinavian
bat with ears shorter than the head, and with the dental
formula—

.2-2? I-I I-I 3-3

ge es. “eer

The only known Skandinavian bats which combine these
characters are the members of the group to which V. serotznus
belongs, and which are commonly known as Vesferus in Europe
and <Adelonycteris in America, but to which Mr Oldfield
Thomas applied Rafinesque’s name, Zpéeszcus (Proc. Zool. Soc.,
London, 1896, 791, 1st April 1897).

The identification of the species murinus amongst the
Skandinavian members of the genus Vesfertz/io, although a
matter of considerable difficulty, does not affect the use of the
generic name. Nilsson (Skandinavisk Fauna, Diggdjuren, 17-
20, Ed. I., 1847) decided that it must have been the bat to which
Natterer later gave the name azscolor. He therefore placed the
latter in the synonymy of V. murznus of Linnzeus, and reinstated
Bechstein’s name myosoézs (correcting it to myotzs), for the Vesper-
tilio murinus of Schreber. Nilsson did not recognise Vesperugo
as distinct from Vespertzlzo. Hence he said nothing in regard
to the tenability of the generic names. Ten years later, Blasius
(Fauna Deutschlands, Stiugethiere, 74, 1857), though admit-
ting that the Vespertzlio murinus of Linnzus could not
be the bat commonly known by that name, considered the
species undeterminable, and therefore reasoned that the
name first applied to it might afterwards be properly used
by Schreber in a different sense. Thus Blasius continued to
apply the name Vesfertz/io of Linnzeus to the genus to which
he had restricted it eighteen years before, notwithstanding the
fact that, according to his own statement, it could not be

1 In the first edition the dental formula is the same, except that the lower
incisors are said to be five in number, an error corrected in the second edition.
2 In Linnzus’s statement the figures 4 and 6 are evidently transposed.

52 VESPERTILIONIDAX—NYCTALUS

made to include any of the Linnzan species. Lilljeborg (Sveriges
och Norges Ryggradsdjur, i., 124-126, 144, 1874) followed
Blasius, insisting that it is impossible to determine whether
Linnzeus’s bat is the species afterwards called Vespertiho
discolor by Natterer, or that called Vesfertzho nilsson by
Keyserling and Blasius, but, contrary to the opinion of Nilsson,
he favoured the latter. Lilljeborg, although aware of Blasius’s
mistake in applying the generic name Vesfertzlio to a group
containing no species known to Linnzus, concluded that as the
error had become time-honoured, it were better uncorrected.

Mr Miller concludes that, ‘“‘ Notwithstanding the incon-
venience to which such a course leads, there can scarcely be
any valid reason for rejecting the identification of Linnzus’s
Vespertilio murinus made by Nilsson. The doubt admittedly
lies between two species, one of which he deliberately chose with
all the facts before him. As nothing in the original description
is in any way discrepant with this determination, it should be
adopted.”

GENUS NYCTALUS.

1825. NyCTALUS, T. E. Bowdich, Excursions in Madeira, etc., 36; based on J.
verrucosus of Bowdich, antedating Pzerygistes madetre of Barrett-Hamilton, Am.
ana Mag. Nat. Ffist., Jan. 1906, 99.

1829. PTERYGISTES, Jakob Kaup, System der Europaischen Thierwelt, 1., 99, 100;
based on Vespertilio “ proterus et leislert.”

1839. VESPERUGO, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s Archiv
fiir Naturgeschichte, i.. 312 (part); based on Vesfertilio serotinus of Schreber and
eleven other species (see below).

1842. NOCTULINIA, J. E. Gray, Ann. and Mag. Nat. Hist., Dec., 258; based on “JV.
proterus and NV. fulvus.”

1856. PANUGO, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung (Dresden),
Neue Folge ii., 131 and 172 ; based on Zeisler of Kuhl, and octula,i.e. La Noctule
of Daubenton.

1878. VESPERUGO, G. E. Dobson, Catalogue of the Chiroptera in the Collection of the
British Museum, 183 (part) ; included also PIPISTRELLUS, VESPERTILIO, and others.

1893. NOCTULINIA, Harrison Allen, Proc. U.S. National Museum, xvi., 30, footnote.

Classification and Synonymy:—Mr Miller has shown that |
the genus Vesferugo of Keyserling and Blasius, as originally
defined, is inadmissible. It included twelve species: sevotznus,
discolor, nilssoni, savit, leucippe, aristippe, noctula, Jlerslert,
kuhlit, albolimbatus, nathusit,.and pipistrellus. These were

DORMOUSE. (3 natural size.)

NYCTALUS 53

arranged in two subgenera— Vesferugo including the thirty-four-
toothed species, and Vesferus those with thirty-two teeth. The
subgenus Vesferus is exactly equivalent to the restricted genus
Vespertitio of Linnzus, to the genera Zf/eszcus of Rafinesque
and Cuepheus of Kaup, all of which antedate it. Apart from
this, however, it would be necessary to find the type of the
genus among the species referred by the authors to the typical
subgenus. These represent two modern groups—the first con-
sisting of xoctula and Jezslerz, the second of the remaining
thirty-four-toothed species. Each of these groups had been
named by Kaup ten years previously. Therefore each of the
constituent parts of the genus Vesperugo was provided with a
tenable name at the time when the composite genus was formed.

The segregation of the xoctula-lerslert group has long been
urged by technical naturalists, on the grounds of their
generally heavy build and long narrow wing, with reduced
fifth metacarpal. It has even been recently advocated by
a field naturalist, Mr C. B. Moffat, who contends that (Lrish
Naturalist, 1905, 104) the very wide difference shown to
exist between the feeding habits of these bats and those of
the Pipistrelle must be correlated with some important internal
differences, a strong argument for generic separation. This
was first attempted by Gray, whose genus Woctulinia was
merely characterised as ‘with the feet quite free, the wing
being only attached to the ankle; they are otherwise like
Scotophilus,” and consequently contained more species than
Nyctalus, as used here; it was adopted by Blyth and Jerdon
(see under synonymy of JV. zoctula). In 1893 the genus was
for the first time accurately diagnosed in its present scope by
Harrison Allen, but the name which he proposed for it is clearly
antedated by Kaup’s Pterygzstes and Bowdich’s Wyctalus (see
Dr Andersen, Ann. and Mag. Nat. Hist., May 1908, 434), both
recently unearthed from oblivion. The present arrangement
was at first rejected by Mr Thomas (Zoo/ogist, 1898, 100), then
reluctantly accepted (Aun. and Mag. Nat. Hrst., July 1901,
34), and now seems likely to meet with universal approval.

The genus Vyctalus includes two British species (LV. xoc-
tula and N. letsteri). These are remarkable for their powerful,

swift-like flight, and the fact that, unlike the Pipistrelle, which
G2

54 VESPERTILIONIDA—NYCTALUS

flies all night, they, in the words of Moffat, “cram themselves
to bursting-point, either once or twice in the twenty-four hours,
during a 70-minutes’ career of mad excitement among the
twilight-flying beetles and gnats.”

The genus is of wide distribution, having representatives allied
to V. noctula—viz., in Japan, WV. /aszopterus (Schreber), with the
forearm measuring 60 or more mm. ; central and south-eastern
Europe,’ VV. maxzmus (Fatio), a mysterious giant of undetermined
status (see Mr Miller, Proc. Bro. Soc., Washington, 13th June
1900, 156), with the forearm of 68 mm. and remarkable skull ;
Sumatra, WV. sumatrana (Cuvier *); China, VV. plancyz (Gerbe) ;
the Himalayas, WV. /adzata (Hodgson); and Mozambique,
N. macuanus (Peters); the last four very little known. Allied
to WV. letsleri are N. stenopterus (Dobson) of Borneo and
Malaysia, a very distinct dusky bat with the forearm of about
37 mm.; my JV. montanus, of the Himalayas, and WV. verrucosus
(Bowdich), antedating my 4. madera, of Madeira, of size
similar to J/ezsZerz, but with distinct crania; and JV. azoreum
(Thomas), of the Azores, a small form, with the forearm vary-
ing from 35-39 in males to from 39-41 mm. in females.

The most nearly allied genera (see Winge, /ordfundne og
nulevende Flagermaus (Chiroptera) fra Lagoa Santa, Minas
Geraes, Brasilien, in E. Museo Lundiz, ii., 1, Kopenhaven, 1893)
are Pipistrellus and Barbastellus, described below : Chalenobus,
of the Australian and Ethiopian regions, with fleshy lobule to
the lower lip; Scotophzlus, of the Ethiopian, Oriental, and
Australian regions, with only thirty teeth; Harpyzocephalus,
of the Oriental region, with tubular nostrils; the long-eared
Otonypteris, from N.E. Africa and the Himalayas ; Myctophalus,
with thirty teeth and small nose-leaf, from Australia ; Lasiurus,
of North and South America, with four mamme and hairy
interfemoral membrane; and <Axtrozous, with remarkable
muzzle, and only four lower incisors, of North America.

The generic characters are as follows :—The body is large
and heavily built.

1 Not found in Switzerland recently (see Mottaz, Budi. Soc. Zool., Geneva, 15th
Nov. 1908, 150.

2 Stated to be of Cuvier, but I have not been able to find the original
description.

NYCTALUS 55

The head is broad and flat; the mouth wide; the muzzle
short and obtuse, the width of the face being increased by
a number of prominent glandular swellings on each side
between eye and nostril. The side of the head as far back
as the ear and upwards to above the eye, together with the
terminal portions of the muzzle, is very thinly covered with
hair. The nostril is tumid at its upper and inner margins,
and slightly channelled on its outer side; the pair project out-
wards and downwards, with a concave space between them.
The lower lip has a thick triangular mental plate. The
upper lip has a thick rounded border, and beyond a deep
groove, interrupted only at each angle of the mouth by a
fold of skin connected with the ear. The eye is, for a bat,
moderately prominent, and situated between the gland and
fae ear (Plate IV., Fig. 1, p. 60).

The ears, widely separated, are thick, broadly rounded
above, when flattened out nearly as broad as long, and
extending very little beyond the eyes when laid forward: the
outer margin of each is convex and reflected backwards,
notched below the tragus with a thickened convex lobe in
front of the notch, and terminating anteriorly behind the angle
of the mouth, considerably in front of the base of the tragus ;
the inner margin is nearly straight above, convex, and turning
inwards below, so as to form a moderately rounded basal lobe.
The tragus is short, curved inwards, expanded above but con-
stricted at its centre, so that it is broadest near the top, which
forms a distinctly rounded or reniform head, broad, rather thick,
and covered with numerous papillae (Fig. 2, Nos. 1 and 2, p. 7).

The wing, attached to the middle of the sole, is large, long,
and narrow, the fifth metacarpal being considerably shorter
than the third. The post-calcarial lobe is well developed,
and the interfemoral membrane ends posteriorly in a salient
angle. The tail is much shorter than the head and body, and
projects very little beyond the margin of the interfemoral
membrane. The thumb is short, and has at its base a small
callosity; the foot is thick; the toes are strong, and well
developed (Plate VII., Fig. 1, p. 86: Plate IX., Fig. 1, p. 126).

There is a single pair of mamme, situated in the pectoral
region.

VESPERTILIONIDA—NYCTALUS

{eran

Fic. 4.—SIDE VIEW (diagrammatic and enlarged) OF SKULLS AND TEETH OF

1. Myctalus noctula. 2. Vespertilio serotinus.
3. Rhinolophus ferrum-equinum.

NYCPALUS 57

The skull is massive and flat; the brain-case angular; the
nasal region somewhat inflated; the premaxillary gap large,
somewhat pointed above, deeply rounded below; the zygomata
are moderately flattened ; the cranial crests not prominent ; the
auditory bulla moderately large (Fig. 4, No. 1, p. 56).

There are thirty-four teeth (Fig. 5), arranged as—

-2-2 I-I 2-2 2
z eG , pu—, m 3_3
bia vd 2-2 Siad,

The upper incisors are arranged in pairs, inclined inwards, and
separated by the wide interval of the premaxillary gap in

I 2

Fic. 5—DIAGRAM OF ARRANGEMENT OF TEETH IN Genus WVycialus.

(1) Upper, and (2) Lower Jaw.

their midst: the outer of each pair lie close and parallel to,
but are shorter than the inner. The anterior upper premolar
is minute, and lying in the internal angle between the canine
and the posterior premolar, which meet externally, is invisible
from without.

According to the best authorities, this genus represents
the most specialised form of the Chzroptera, amongst which it
is nowhere excelled in power of flight ; a remarkable change of
Opinion, since in 1878 the leading authority of the day—
Dobson (Catalogue of Chiroptera, 100)-—wrote that ‘“ From
whatever point of their structure they may be considered, the
Rhinolophide are evidently the most highly organised of
insectivorous Bats.”

58 VESPERTILIONIDAZ—NYCTALUS

THE NOCTULE, OR GREAT BAT
NYCTALUS NOCTULA (Schreber).

1760. LA NOCTULE, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s /istcire
Naturelle, viii., 128-129, 135-137, pl. xvill., fig. 1; also Wém. de l’Acad. Roy. des
Sct., 380, pl. ii. (15), fig. 1, 1759, published 1765 ; described from France.

1775. VESPERTILIO NOCTULA, J. C. D. von Schreber, Die Sdugthiere, i., pl. lil,
166-167, evidently naming Daubenton’s Za Woctule ; Bingley ; Pennant ; Donovan ;
Fleming ; Jenyns; Bell (ed. 1) ; Clermont ; Newman.

1776. VESPERTILIO LARDARIUS, P. L. S. Miller, Watursystems Supplements und
Register Band, 15, naming Schreber’s Sfeckuiaus, t.e. V. NOCTULA, as above.

1776. GREAT BAT, Thomas Pennant, Aritish Zoology (ed. 4), 1., 128, pl. xiil., No. 38.

1789. VESPERTILIO ALTIVOLANS, Gilbert White, Seddorne, letter xxxvi. to Thomas
Pennant, September 1771 ; described from Selborne, England.

1789. VESPERTILIO MAGNUS, John Berkenhout, Syzopsis of the Natural History of
Great Britain and Ireland, ii., naming Pennant’s Great Bat.

1806. VESPERTILIO SEROTINUS, Isidore Geoffroy, Ann. du Mus. d’ Hist. Nat., viii.,
194, pl. 46, fig. 1 ; transference of name in error.

[1812. VESPERTILIO AURICULATUS, John Walker, Essays on Natural History,
472; described from Edinburgh, Scotland, but only doubtfully referable to
XN. noctula.|

1816. VESPERTILIO MAJOR, W. E. Leach, Systematic Catalogue of the Specimens of the

Indigenous Mammalia and Birds that are Preserved in the British Museum, etc.
(London), 5; renaming V. zoc¢uda, but without description.

1819. VESPERTILIO PROTERUS, Heinrich Kuhl, Ann. der Wetterauische Gesellschaft
fiir die gesammte Naturkunde, iv., 41 ; renaming Daubenton’s La Noctule.

1826. VESPERTILIO LAICOPTERUS, J. E. Gray, Zoological Journal, ii., 109 ; included
as a synonym of Vespertilio noctula; perhaps a misprint for lasiopterus.

1827. VESPERTILIO FERRUGINEUS, C. L. Brehm, Orazs (Jena), iii, 26; described
from Jena, Germany.

1829. PTERYGISTES PROTERUS, Jakob Kaup, System der Europiischen Thierwelt,
1.5 100.

1838. SCOTOPHILUS NOCTULA, J. E. Gray, Mag. Zool. and Bot., 497 ; MacGillivray,
Bell (ed. 2).

1839. VESPERUGO NOCTULA, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s
Archiv fir Naturgeschichte, i., 317; Blasius ; Fatio; Dobson; Harting, Zoologis?,
1887, pl. ili, 167-171 ; Blanford ; Woodward and Sherborn; Lydekker; Millais,
pl. vii.

1842. NOCTULINIA PROTERUS, J. E. Gray, Ann. and Mag. Nat. Hist., Dec., 258.

1845. NOCTULINIA NOCTULA, Edward Blyth, Journ. Asiatic Soc. Bengal, xiv., 340;
referring to T. Hutton’s Mussooree specimens (see Proc. Zool. Soc., London, 1872,
707); T. C. Jerdon, Mammats of India, 36, 1867; Harrison Allen, Proc. U.S.
National Museum, xvi., 30, footnote, 1893 (diagnosis of genus).

1856. PANUGO NOCTULA, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung
(Dresden), Neue Folge, ii., 131, 172-174.

THE NOCTULE, OR GREAT BAT 59

1869. VESPERUGO NOCTULA, var. MINIMA, Victor Fatio, Vertébrés de la Suisse, i., 58 ;
described from Geneva, Switzerland: See Mottaz, Bull. Soc. Zool. (Geneva), 15th
Nov, 1908, 151.

1897. PTERYGISTES NOCTULA, G. S. Miller, junior, Worth American Fauna, No. 13,
87, footnote ; Ann. and Mag. Nat. Hist., October 1897, 383-384 ; Oldfield Thomas,
Ann. and Mag. Nat. Hist., July 1901, 34; Johnston ; Méhely ; Cabrera; Millais, 61.

1898. PIPISTRELLUS NOCTULA, Oldfield Thomas, Zoologist, 100; Millais, 61.

La Noctule of the French; de friihfliegende Fledermaus of the
Germans.
Noctule, from the French zoctule, diminutive from the Latin xox,

Me a MPA (5) Senile
genitive octzs, 2.¢., “ night.

Local Names:—Rat Bat of Durham (Roebuck, Naturalist, 1886,
113), Cheshire (Oldham), Leicester (Montagu Browne), Buckingham
and Berkshire (Cocks, Zoologist, 1878, 334), and probably of other
Saumtics kee rot, 2.2, rere or rear tat (see “rere mouse” under
PIPISTRELLE) of Gloucester (Newstead). Ystlum (pronounced “ slim ”’)
Fawr, z.e., Great Bat of Wales (Forrest).

Distribution :—The Noctule, or bats closely resembling it, inhabits
the wooded districts of boreal and transitional Europe and ‘Asia, from
sea-level to over 4000 feet (in the Alps), from south Scotland and Norway,
Denmark, north Germany, and middle Russia, to the Mediterranean,
Black Sea, Caspian, and Turkestan (Samarkand), with (?) Minorca
(Barcelo), and Sicily (Blasius); and from Great Britain probably to
eastern Siberia; but the exact limits of its range are very imperfectly
known, as are those of the allied forms mentioned above (p. 54), upon
which, or others, are probably based records of Noctules from Japan,
China, Nepal, Sikkim, Kandahar, Ceylon, Singapore, Sumatra, Java, and
Mozambique.

Throughout the south of England, from Norfolk to Cornwall (Cocks,
Naturalist, (1), 1851, 37; Rodd, Zoologzst, 1891, 347), but rare in Wight
(More, Journ. cit., 1894, 148; Wadham), JV. zoctula is a common species.
Its flight may even be observed in the heart of London, above the Serpen-
tine (Macpherson), or in many of the parks (Millais). It probably occurs
in every county of Wales, where Forrest reports it from Flint, Mont-
gomery, and Radnor ; Newstead along the Dee Estuary ; Coward and
Oldham in Denbigh, Carnarvon (Zoologist, 1901, 53), and Anglesey ;
and Caton Haigh, in Merioneth (Zoologist, 1887, 293): in the latter
county it loves the oak-covered hills, and in Carnarvon it is abundant
at Nevin, right down to the sea-cliffs.

In the north it is widely distributed, and abundant as far as south,
and probably central, Yorkshire, where it is found at an elevation of
700 feet, at Carperby in Wensleydale; but towards the north and west
of the county it becomes less numerous, and Northallerton was its
most northern known British locality (Bell) until Nelson and Roebuck

60 VESPERTILIONIDH—NYCTALUS

extended its range to Bishop Auckland and Newcastle-on-Tyne respec-
tively, the latter the extreme northernmost limit of Durham (Naturalist,
1884-5, 202; 1886, 113 and 173; Harting, Zoologzst, 1887, 260). In
Lakeland, Macpherson, although denying its general distribution, placed
its range as “at least as far north as our most southern limits,’ and
particularly at Carnforth in Lancashire, and probably at Bowness-on-
Solway in Cumberland. MHarting, on the authority of Lee, records
its occurrence at Kendal in Westmorland (Zoologzst, 1887, 170).

For Scotland there was until recently no certain record, although
Sir William Jardine stated that it had been seen about the river Annan
in Dumfries (Statestical Account of the Parishes of Applegarth and
Sibbaldbie, 1835, 175), and Fleming identified with it John Walker’s
Vespertilio auriculatus+ Of late years, Service (Aun. Scott. Nat. Hist.,
1896, 202) has on several occasions noticed darkly coloured bats flying
high near Dumfries, and his description is suggestive of a member
of the genus WVyctalus. The first actually known Scotch specimen,
shot at Deune, on the Tay, in Perth, by Charles Eversfield, on 13th
October 1904, was forwarded to Millais: the skeleton is in the Perth
Museum (Zoologist, 1904, 425). A second was sent to William Taylor
from Duffus, near Elgin, on Ist October 1909, and others are stated to
have been seen about Elgin and Llanbryde, where there is probably a
colony (Aun. Scott. Nat. Hzst., 1910, 52-53).

So far as is known, JV. zoctula is absent from all the Scotch islands,
as well as from Man and Ireland, although Dobson at one time
identified with it two bats, a male and female, procured from Tandragee,
Co. Armagh, by Barrington (Zoologist, 1874, 4071-4074; Harting,
Zoologist, 1887, 168). In 1878 the same authority cited them as
“V. letslert” (Catalogue, 215), but in 1889, writing to Barrington of
some further Irish specimens, he thought that, if certain characters

1 Unfortunately Walker’s description of V. auriculatus, except that it clearly does
not apply to a Rhznolophus, was not drawn up with sufficient exactness to be definitely
applicable to any known Scottish species. It does not include any dimensions, and
the thirty-two teeth and lanceolate tragus would refer rather to V. sevotinus, which is
not known to occur in Scotland, than to JV. zoctu/a. Alston evidently considered the
description inexact, or he would not have connected it with Myot’s daubentoni (Fauna
of Scotland, 8), a species having thirty-eight teeth. On the whole, it seems better to let
such laxly written descriptions remain unconnected with any particular species than
to attempt a definite decision upon insufficient grounds. It may be interesting to
transcribe the whole passage :—

“Auriculis duplicatis. Vespertilio auriculatus, caudatus, naso oreque simplici,
auriculis duplicatis capite minoribus. . . . Descriptio maris. Dentes XXXII. numera-
vimus. Primores superiores 4 acuti distantes ; inferiores 4 acuti contigui. Laniarii
superiores 6: anticis maximis acutis; inferiores 6: anticis majoribus. Molares utrinque
6. Aures duplicate, capite multo minores. Exterior major ovata obtusa. Interior
minor brevior lanceolata. Palmze palmato-alatee maxima, pollice unguiculato. Plante
pentadactyle fissz, digitis pilosis unguiculatis. Cauda geniculata, 6 articulis.
Membrana juxta caudam, margine ciliato. . . .”

Se eee ee ee

THE NOCTULE, OR GREAT BAT 61

which he found in them were present “in the perfectly adult animal,
then the Irish specimens represent either a new species, or a curious
case of hybridization” (szc). The matter had not been settled at the
time of Dobson’s death, but de Winton compared these Irish with English
individuals of JV. /ezslerz, and found no difference between them (Alcock,
Trish Naturalist, 1899, 170-1). After careful examination, I find myself
entirely in agreement with de Winton, and in any case it is clear that
there exists no evidence for the occurrence of the Noctule in Ireland.

It is possible that, as suggested to me by Moffat, this species may
have recently extended its range in Britain. The idea can only be
regarded as hypothetical, but the present distribution of the bat is
consistent with a comparatively recent arrival. Its range on the conti-
nent of Europe brings it into contact with greater cold than it could
experience in Scotland, and the absence of so widespread and power-
ful a flier from Ireland is intelligible on some such basis. Such
a supposition would be in concord with its reputed rarity at
Selborne in Gilbert White’s time, its present abundance there, and its
recent appearance in Scotland. Moffat ingeniously adds that the much
shorter flying season reported by White would be natural in a species
newly arrived, which as it became acclimatised might be supposed to
extend its period of activity on the wing.

Distribution in time :—Bones of this bat have been found in cave
deposits of pleistocene age at Banwell, Burrington Combe, and Hutton
(Woodward and Sherborn).

Period of gestation :—Certainly not less than thirty-eight days
(Daniell, Proc. Zool, Soc., London, 1834, 129-132), but probably at least
forty-nine ; see above, page 33, and Whitaker on Pipistrelle.

Number of young, and breeding season:— There is rarely more
than a single young one in Britain (see p. 80). It is born most usually
towards the end of June.

Description:—The general form and appearance of the Noctule is
typically that of its genus. The forearm is massive, the lower leg
thick but short, the foot conspicuously powerful.

The fur is soft and long. The face is thickly furred from behind
as far as the glandular prominences, and anteriorly the muzzle carries
a few long hairs. Upon the upper surface of the wing the fur extends
as far outwards from the body as a line drawn from the middle of the
humerus to the knee-joint, and on both surfaces of the interfemoral mem-
brane as far back as the middle point of the lower leg: on the under
surface the wing is rather thickly clothed as far as a line drawn from the
elbow to the knee-joint, and a band of fine short hair frequently, but
not always, passes behind the forearm to the carpus. The inner side
of the ear is covered with fine short hairs, which also appear upon the
tragus, and sparsely upon the reflected outer margin of the conch.

62 VESPERTILIONIDA.—NYCTALUS

The colour above and below is yellowish or golden, almost ochraceous,
brown, very little lighter on the under surface; the hairs are almost
unicoloured, the basal and concealed portions being only of a slightly
lighter shade. The wing and foot are dusky; the lips, ear, and
nose lighter. In the newly-born young, the wing and interfemoral
membrane, the ear, muzzle, foot, and tail are pigmented nearly as
deeply as in adults, but all those parts, including the forearm, which in
the adult are covered with hair, are of a dull flesh colour. The eyes do
not open until at least the twelfth day, and the hair, preceded by
pigmentation, begins to make its appearance when the bat is about a
fortnight old (Whitaker, in MSS.). After the hair has been acquired,
the young at first are darker than the adults (Forrest, in MSS.).

I find no trace of seasonal variation or moult, but it is probable
that, as with JV. /ezsler¢, the coat is pale and faded in early summer,
and deepest and richest
in early autumn just ee ee
before hibernation. Old
males are said to possess

the brightest golden fur. \ ey y
The skull is typical of ae ae
the genus. It is character- (a7 ae Ga

ised by a strong lambdoid,
but, as a rule, weak sagittal
crest ; there is much indi-
vidual variation in size and
proportions (Fig. 4, No. I,
p. 56).

The teeth are shown
in Fig.6. The inner upper
incisor is bicuspid in the
young, but adults lose the
small outer cusp. The
outer upper incisor is
much shorter than the

inner, dut much broader in dle
transverse section at the a

base: its crown is hol- a ee
lowed out for the recep- oe

tion of the tip of the Fic. 6.—FRONT VIEW OF INCISORS AND CANINES OF

ipmericanine. ..%vie varie Nyctalus noctula (enlarged and diagrammatic.)

incisors are crowded and overlap, so that their broad crowns are oblique
and parallel to the jaw.

Laver mentions an almost black specimen, and Norton others “ pied
with grey, also with black, and sometimes a light colour round the

THE’ NOCTULE, OR GREAT BAT 63

neck” (Midland Naturalist, 1883, 151). Otherwise, individual and
geographical variation appears chiefly in size, the larger forms showing
a correlated development of the cranial ridges. In a large female from

Devon the forearm measures 54, or fully 3 mm. above the average, but
it is still far below that of maxzmus.

DIMENSIONS IN MILLIMETRES :—

MALES.
mulled :
sle|2 2 a : 5
a 5 sr) 5 SS e
Syl el @ ests NE gales eater 3 ES ae
so) sl = | 4 Ee 3 a ee oy >) OD
ee Se ira Be has 2 5 8 Se
3 | © | & } 8° Ss = 80 8 8 ne
S| ©| x His 4 3 = 2 3
-| 3 = B 4 = $
20) = Sp ee 5 = =
<=) Sl eo) = oS
No. of Items | 7 By Gal ad 7 5 5 5 3
NE ee =
Maximum . | 80 18 7 | 53 | 8 ( 51 | ( 375
Average sai ay Wg 5 | 49°5 | +} 17-18 S ~ 50 ;7|7-8 87-96 46°5-50 | 36°5-39°5 a0
Minimum .|76 | 16| 4| 43 = | 49 | | 353
FEMALES.
No. of Items 8 6 4 8 oe oe 9 an is 7 | u 3
Maximum .| 82 its ae || pees) | Ae 54 | a3 Ss ( 3887
Average manera -. | 49 17-18 | 11-12 51°3 -| 7-8 91-94 48-52 38-42 ~
Minimum .|73 | 16] .. | 41 | 50 | | 362
|

* Norgate (Zoologist, 1906, 26) gives the average of many as 14} inches, and the extremes as varying
from 14} to 15 inches.

The female is, on the average, the larger sex.

Proportionate lengths :—Foot, without claws, about -64 to -66 of
lower leg; fifth metacarpal about -79 to -80 of third; lower leg about
-33 to -34 of forearm, and about -24 of head and body.

Skull :—Greatest length, 19; basal length in middle line, 14; palatal
length in middle line, 7; from posterior border of 7° to anterior border
of canine, 7 to 7:5; same in lower jaw, 7 to 7-5; greatest breadth at
zygoma, 12-75; posterior breadth, 12; breadth between orbits, 12-2;
breadth at constriction, 5.

Very few published details of the weights of Noctules are available,
but Aplin, who weighed two on 25th July, each of about 26 grammes,
and one on 13th October of 35-44 grammes, found a considerable increase
of weight in autumn (Zoologist, 1885, 344). The following (in each
case converted to grammes) are of interest:—Two males, each 30-12

64 VESPERTILIONIDAE—NYCTALUS

grammes (Gilbert White, letter xxxvi, to Pennant); 15-95 grammes
(Fleming); a captive female, June, 17-72 grammes (Daniell); one, 28th
April, 24-80 grammes (C. H. B. Grant); a female, 27th March, 38-88
grammes (Charbonnier).

Distinguishing characters :—The Noctule is an easy bat to identify.
On the wing its dashing, lofty, swift-like flight is very distinctive,
and is only approached by JV. J/ezslerz, which is, however, too small
to be confounded with a mature Noctule. In the hand no species
approaches its size, with the exception of the Greater Horseshoe,
to be at once recognised by its nose-leaf and the absence of a tragus;
and the Serotine, also a very different animal.

We are indebted to Daubenton for the first discrimination
of this fine bat. He described it and figured the head so
long ago as 1759, and Buffon subsequently gave it a place
in his great work. The first notice of its occurrence
as a British species is in Gilbert White’s Natural Hestory of
Selborne,' in which it is mentioned three times, and its general
appearance, early autumnal retirement, “very rancid and
offensive smell,” and lofty flight are subjected to criticism. It
was the latter peculiarity which suggested to White the
appropriate name of Vesfert:lo altwwolans.

Other British writers were not slow to follow White, and
we find the Great Bat or Noctule and its habits described
with more or less precision in the works of Pennant, Bingley,
Donovan, Fleming, and Jenyns. It appeared in both editions
of Bell, who, himself a resident of Selborne, observed it
for several successive seasons, probably on the very ground
where it first attracted the attention of its venerable dis-
coverer. Its habits are now fairly well known, thanks to the
labours of a number of naturalists, amongst whom the names
of George Daniell, George Dowker, John Wolley, and Messrs
T. A. Coward, Charles Oldham, and Arthur Whitaker stand
out most prominently.

The Noctule is a tree-loving species, existing in some
abundance in the southern and midland counties of Great
Britain, especially where there are old trees in sufficient numbers
to provide it with secure retreats. It dwells gregariously in
large companies in holes of trees or of buildings, and its

' Letters xxii., xxvi., and xxxvi., to Thomas Pennant, dated 2nd January 1769,
8th December 1769, and September 1771: original edition, 63, 75, 93-94, 1793:

THE NOCTULE, OR GREAT BAT 65

presence is often betrayed by the excrement, which, as
at Ragley, the seat of the Marquis of Hertford, may
lie so thick as to darken the ground under some ancient
resort.

From such holes, the entrance to which is often polished
smooth by the frequent passing in and out of the occupants,
the insertion of a flexible stick sometimes dislodges a surpris-
ing number of bats. For instance, Pennant states’ that the
Reverend Doctor Buckworth saw taken from under the eaves
of Queens’ College, Cambridge, one hundred and eighty-five in
one night; on the second night sixty-three were taken, and
on the third night two. From another colony, described
from King’s Lynn by Mr H. B. Booth,’ nearly three hundred
bats were seen to issue for their evening flight. It has been
objected that, since there is no reason to believe that the bats
were all subjected to the rigid examination necessary to deter-
mine the specific distinctions of these animals, it is probable
that other kinds were mingled with them in these great con-
gregations ; but repeated observation by many naturalists has
led to the belief that the present species rarely, if ever, occu-
pies its retreats as a tenant in common with others, and
moreover the occupants of many lesser yet thickly crowded
Noctule dens have since been counted by reliable observers.
Indeed, the species is so abundant that Frederick Bond* informed
Edward Newman that he had observed no less than three or
four hundred on the wing together.

Although preferring in summer to seek its retreat in the
hollow cavities of trees, the Noctule is, as the above anecdote
shows, by no means loth to creep under the eaves of
houses, but, so far as I know, it has never been met with in
caves, a statement,* that it breeds in the Mendip caverns of
Somersetshire, being so contrary to the general experience that
I hesitate to accept it without corroboration. In fact, it seems
to bé almost a rule with it to seek good winter quarters
amongst houses or ruins. From these it issues forth nightly
for a few weeks in the spring, but after a time forsakes them
for an entirely arboreal life, returning, however, for its autumnal

1 British Zoology, London ed. of 1812, I, 179. 2 Zoologist, 1905, 427-429.

3 Field, 7th March 1874, 218. 4 Stanley Lewis, Zoologist, 1906, 69.
H

fe VESPERTILIONIDAZ—NYCTALUS

hibernation. In like manner, its feeding grounds change with
the season, the choice of them and the presence or absence of
the bats being no doubt determined by the distribution of the
insects upon which they feed. At Dunham, in Cheshire,
write Messrs Coward and Oldham, ‘‘at first these bats fly very
high, squeaking and chasing one another around and above
the tree-tops. During summer they frequent the open glades,
generally flying high; but towards the middle of September
they resort in great numbers to the water-meadows by the
river Bollin, flying up and down alongside the park-wall, often
not more than ten or twelve feet from the ground.”

The general conformation of this bat is essentially adapted
to the capture and mastication of beetles. Its broad muzzle
and strong jaws are quite equal to the reduction of the larger
kinds, such as the dor-beetle or the cockchafer, of which,
according to the elder J. H. Gurney,’ one has been known to
consume as many as thirty in half an hour. The stubborn elytra
are invariably cut off at the base and rejected, and these, where
the bats feed thickly over water, have astonished an observer,
ignorant of their habits, as they fell in showers on the surface.’
In fact, so strong is this species that, as related below, it seems
likely that it has but seldom need to call in the assistance of
the interfemoral pouch, the mainstay of the smaller kinds. At
all events, although he has seen the pouch used in captivity,
thus showing that this species is not unacquainted with the
habit, Mr Oldham, who has spent hours watching wild
Noctules, has never once detected them in the act of pouching
an insect; and Frank Norton,* although believing that the tail
“is certainly brought very much into play,” never succeeded in
proving this, as in the case of the Whiskered Bat, which, he
states, is easily caught dy ¢he wzng when a beetle is suspended
near its beat on a hook. On the other hand, Mr Whitaker
informs me that on one occasion he deceived one into pouch-
ing a pebble thrown up to attract it. The bat carried the
pebble a distance of some yards before dropping it with a splash
into some water. This propensity of bats to mistake pebbles
or other objects for their prey is well known ; and an object

1 Zoologist, 1874, 4153. 2 Thomas Ford, Fze/d, 1oth September 1898, 470.
3 Midland Naturalist, 1883, 151-152.

THE NOCTULE, OR GREAT BAT 67

thrown towards one, even when flying high, will often attract
it within gunshot. It does not necessarily indicate, as Mr
Whitaker suggests, that the bats hunt chiefly by eyesight.
Mr Arthur Patterson’ finds that a good mimicry of its shrill
notes serves the same purpose, and he relates that, having
on one occasion a slightly wounded one shrieking in his
pocket, its companions came so near him that he could feel the
whisk of their wings.

Besides beetles, this bat eats moths and lesser insects also
when they come in its way, and it will certainly devour
them freely in captivity. It is difficult, however, to
prepare a precise list of the creatures upon which it
preys, since digestion is very rapid and the contents of the
stomach of shot specimens are usually in a highly disinte-
grated condition. Mr Coward noticed* a number flying
low over the fields at the edge of the cliffs at Nevin, Carnarvon-
shire, their food being apparently the winged males of a species
of black ant, and Mr J. Steele Elliott has observed them
hawking mayflies over the river Severn.®

The flight of the Great Bat is, typically, high, straight and
rapid, and it may often be seen careering with swifts or
swallows at great elevations; but it is by no means tied to
habit in this respect, and frequently manceuvres near the
ground or occasionally dips towards a stream to seize its
quarry. During the fine midsummer evenings, when the
abundant cockchafers are humming on every side, it is in its
glory. Then it flies high and straight, and its shrill,
clear voice is heard as it passes overhead, interrupting itself
only to dart at some insect, and then passing on. But an
observer will not watch its actions long without noticing a
movement which looks like the falling of a tumbler pigeon,
with a consequent drop of about from one to six or eight
feet. Sometimes this is repeated every few yards as long
as the animal is in sight, and its meaning has been the
subject of some discussion. The usual explanation is that
the fall takes place with closed wings, and must be occasioned
by the capture of some insect so large and intractable that
the anterior joint of the wing, with its well-armed thumb, is

1 Zoologist, 1898, 304. * bid. f9OK;.53; 3 [bid., 1901, 70.

68 VESPERTILIONIDAZ—NYCTALUS

required to assist in retaining it until masticated.* Proof
of this contention is, however, not forthcoming, and it is note-
worthy that Mr Oldham, who has spent hours watching Noctules -
under favourable conditions, both with the naked eye and with.
a strong glass, has never detected them using the thumb. to
rend asunder their prey. It seems clear, from the evidence of
Messrs Coward, H. E. Forrest, J. G. Millais, and Oldham,
that the plunge takes place obliquely downwards, both wings
being clearly extended—a posture very different from a vertical
fall, such as occurs when there is a loss of balance, as when
one wing is broken by shot. My own conclusions are
thoroughly in accord with the opinion’ that the plunge is
made after insects below and off the direct line of flight. The
fact that it may take place not only downwards, but in an
oblique direction, is indeed an almost proof positive of its
objective. It is a falcon-like swoop, the tremendous force
of which is illustrated by Mr Whitaker’s* experience of a bat
which, in pursuit of a pebble, struck a butterfly-net so violently
as to break a wing and nearly dash the net from his hand.

Sometimes, however, food is not so readily obtained. With
a cold east wind, or indeed a strong wind from any quarter,
a change of hunting ground is required, and the Noctule may
then be seen taking a humble and silent flight in some
sheltered corner, fluttering about with half-closed wings, and
appearing to be very little at home or, indeed, like itself; for
instances are on record where several have been shot under the
belief that they were of some unknown species.

Long ago Wolley graphically described * its flight as always
strong, but varying “remarkably at different times, no doubt
influenced, like that of the swallow, by the casual range of its
prey; at one time it may be seen flying away, straight and
swift, at a great height in the air, no more to appear that
evening ; at another it will be performing a great circle, return-
ing perhaps once in five or ten minutes; or it may be flying
low (and then I think silently) along the streets of a town:
again it is wheeling round tall elms, in company with others

1 See, for instance, Oxley Grabham, Zoologist, 1899, 131.
2 Trans. Caradoc and Severn Valley Field Club, 1900 (Feb. 1901), 243.
5 Ian. MSS. 4 Zoologist, 1845, 953

PLATE III,

ow a we

ae Ld
castles Riinitgnmsncerr®

NocTULE BATs.

(Approximate natural size.)

THE NOCTULE, OR GREAT BAT 69

of its own species, at the time of year when the small
hairy cockchaffer . . . is swarming about them. Then its
powers are seen to perfection, and the great advantage over
the feathered tribes that it derives from the mammalian
articulation of its wings is beautifully evident. . . . Its latest
are by no means its lowest flights; even in November I have
observed it at such a height, that I could hardly have seen
it, had not my eye been directed to it by its cry. This is the
cricket-like chirp which it always makes with incessant repeti-
tion when flying high: ... it calls my attention to the
animal when it is within a hundred yards or so, frequently
giving me the first intimation of its presence: it is so readily
distinguished by its peculiar cadence from the chirp of other
bats, that however dark the evening, it gives me certain indica-
tions of the Noctule.”

The habit of sometimes flying straight away and to a
distance from its diurnal retreat may frequently have the effect
of preventing the species from coming under attention in its
actual home. Dowker,’ for instance, although observing them
issue from his house in Kent, whence they immediately took
flight across the marshes, searched for them in vain on the
wing in the immediate neighbourhood, to which they only
returned at the conclusion of their flight.

It was long thought that the Noctule remains in
activity for a shorter period than any other bat, coming
out later and retiring earlier: White’? never saw it abroad
till the end of April, nor later than July; but this excep-
tionally short season at Selborne must have been due to
causes other than hibernation, since subsequent observers
have lengthened the flighting period until it is now known to
include every month of the year, excepting only the latter part
of December and January. No doubt its appearance in the
spring and its retirement in the autumn depend to some extent
on the mildness of the season, but Mr L. Buttress * has noticed
it abroad in Nottinghamshire on 14th March, the thermometer

1 Zoologist, 1889, 258.

2 Letters xxii., xxvi., and xxxvi., to Thomas Pennant, dated 2nd January 1769,
8th December 1769, and September 1771.

3 Field, 23rd April 1892, 585.

70 VESPERTILIONIDAE—NYCTALUS

then standing at 40° Fahrenheit, and according to Mr Steele
Elliott,’ it flies commonly in Worcestershire towards the end
of that month, the earliest which he ever saw on the wing having
been active on 22nd February. Mr Oldham has detected
it in flight on 18th, 20th, 8th, and 18th March in the
years 1902 to 1905 respectively. In autumn the flying season
seems to terminate ordinarily towards the end of October,
during which month Mr O. V. Aplin has seen? it out in
Oxfordshire in very cold weather, both foggy and _ frosty.
More than one writer has reported * the flying of Noctules so
late as the second week of November, but probably Mr G. T.
Rope’s observation * of the 28th of that month, and Mr Aplin’s,
as quoted by Mr Millais, of 3rd December, are the latest
recorded.

So far as I am aware, Dowker’ was the first Englishman
to point out that the evening flight lasts only for about
an hour. He watched the bats emerging from their winter
quarters under the gable of his house early in May and
June. The weather seemed to have little influence on their
activity once they were in full flight, but at first only a
portion of the colony appeared to be active enough to
take wing. He counted only fourteen on 1st May, but
the number increased to sixty-seven on 4th June and
to one hundred and twenty on the 12th, after which they
appeared in decreasing numbers, until in the middle of the
month only one was visible. The time of their appearance
varied from 7.50 to 8.15 p.M., the return of the first taking
place at about 9, when, if many happened to be out, they
came flying round their home like a cloud of bees, awaiting
opportunity to enter. With unimportant variations, Dowker’s
statements are closely corroborated by Mr Booth,° who describes
the appearance of the members of a large colony in small
parties in the late April and early May evenings; their steady
increase, until in early June the numbers leaving nightly became
almost incredible, and later their rapid decrease, until in July

1 Zoologist, 1904, 455. 2 Tbid., 1885, 344.

3 E.g., Forrest, Steele Elliott, Henry Laver (specimen obtained on the 1oth),
Millais.

4 Zoologist, 1891, 167. 5 Ibid., 1890, 217; 1891, 305. 8 Tbid., 1905, 427-429.

LHe NOCTULE, OR'GREAT BAT 71

not a single bat would leave the dormitory, although crowds
might be seen hawking in the air outside each evening. It
appears, therefore, probable that a summer “camping out”
season is a regular feature of the animal’s annual routine.
The order of leaving the den was very ‘curious. A loud
squeaking would be heard for a minute or two near the inside
of the hole; then a Noctule would appear, and launch itself
into the air, followed in quick succession by four or five more.
Then silence ensued for a minute or so, followed by the squeak-
ing noises again, and another party of five or six would follow
each other out. This mode of procedure would be carried on
continuously. Silence, squeakings, and then bats in constant
succession. Most of the ‘parties’ consisted of from five to
eight individuals, though on occasions (particularly on evenings
when the greatest number were leaving) I have seen as many
as a dozen or more follow each other without a break, but this
was exceptional. These small parties were called families by
my friends, but they scattered off individually and in separate
directions immediately they were clear from the hole.” The
bats were never detected returning to their den in autumn,
although always there in the spring.

Working mainly in Cheshire, Messrs Coward and Oldham
have added largely to our knowledge of the times of flight of
this bat, and no account of its natural history could be regarded
complete that did not include substantial quotations from their
essays on the subject. In Cheshire, “on fine evenings,” writes
Mr Oldham,’ ‘one’s attention is often attracted by the shrill
squeak of the Noctules which are flying in company with the
Swifts, at an altitude difficult to estimate accurately, but
certainly not less than from seventy to eighty feet. ... As
the light fades, the Bats descend to a lower level, and feed
at a height of from fifteen to thirty or forty feet above the
fields, pools, and open places in the woods. The crunching of
their jaws as they masticate their insect prey may then be
heard distinctly.

“The time at which the Noctule issues from its retreat does
not always bear the same relation to the hour of sunset, and
sometimes differs considerably on consecutive evenings. Wind,

1 Zoologist, 1901, 51-59.

72 VESPERTILIONIDA—NYCTALUS

temperature, and other atmospheric conditions, rather than
the actual hour of sunset, probably determine the time at
which the Bats emerge, and the duration of their flight. Rain,
if not heavy, does not incommode them whilst feeding, but
if the night be cold and windy few or none will be seen. It
is probable that individual Noctules do not always resort to
the same den throughout the summer, for the numbers which
emerge in the evening are not constant, and even on con-
secutive evenings, when the atmospheric conditions appear to
be identical, the number sometimes varies considerably. On
the other hand, it is possible that on some evenings the whole
strength of the colony does not turn out, and that some of the
Bats remain in the den all night.”

Mr Oldham found that the time of first appearance varied
from twenty-eight minutes after sunset on 21st April to seven
minutes before sunset on 14th August. The hour of vespertinal
emergence thus becomes progressively more diurnal as autumn
approaches, until in the second week of September I have
observed them to appear at about 6.30 p.m., and the October
and November flights are, according to Mr Millais, all rather
diurnal than vespertinal. ‘The Bats,” writes Mr Oldham,
‘leave the den in rapid succession—on August 27th twenty-
two emerged within a minute—but their return is much less
regular. This is probably due to the varied success of
individuals in obtaining food. During August and September,
at any rate, on fine still evenings, the duration of the vesper-
tinal flight is sometimes less than an hour; on August 27th
a Bat entered the den fifty-five minutes, and on Sept. 4th fifty-
seven minutes, after the first had emerged. On each evening,
however, the flight in some cases lasted at least an hour and
a half; and on Sept. 5th some of the Bats were absent for more
than two hours. It is probable that on wet and windy evenings
the duration of the flight is even less than an hour, but I have
no data to prove this. It is easy to count the Bats as they
leave their den in the twilight, but a difficult matter to make
sure of the number that return. They do not often enter the
hole immediately on their arrival, but dash round and among
the trees, and in many cases pitch several times for an instant
on the tree-trunk near the hole. Their advent is proclaimed

ee es ee

THE NOCTULE, OR GREAT BAT 73

by the beating of their wings, but even on moonlight nights
all that one sees is a form silhouetted for an instant against
a patch of sky. When the Bat is flying against a background
of tree-trunks or foliage, one can see nothing. It is true that
a faint rustle may be heard when a Bat actually enters the
hole, but this resembles the noise made when it pitches for an
instant on the tree-trunk; and if two or more Bats arrive to-
gether, as often happens, the confusion is increased. A good
deal of intermittent squeaking may be heard in the den after
the arrival of the second Bat.”

In the paper above referred to, Mr Oldham, judging from
the habits and demeanour of captive specimens, expressed
the opinion that the period of activity is limited to the
short vespertinal flight, and that it does not again leave its den
before the following evening. Mr Steele Elliott, dissenting from
this view,’ mentioned two instances of matutinal flight which
happened to come under his notice; and later Mr C. B. Moffat,
from the analogy of the closely allied Leisler’s Bat, as well as
from his own slight experience of the present species, argued”
that there must be two flights each night. Mr Oldham sub-
sequently undertook an all-night watch*® outside a Noctule
den in Cheshire on 20th May, with the result that the
occurrence of a matutinal flight in this species may now
be regarded as beyond question, although it is possible that
fewer individuals are abroad at dawn than in the evening
twilight. Mr Oldham found that no bat entered or left the
hole after 9.36 p.m., but that there was intermittent squeaking
until ten o'clock, and ‘‘a slight squeaking in the den at long
intervals until 2.40 (eighty-five minutes before sunrise), when
the noise increased, and more than one bat emerged—in the
gloom I could not tell the exact number—and all was still
until 3.20 (forty-five minutes before sunrise), when three
returned. These dashed round among the branches, alighting
on the trunk at the mouth of the hole once or twice, and then
dashing away again before entering the den, as Noctules
generally do on returning from the vespertinal flight. There
was no squeaking after the Bats entered the den, and I heard
none until 4.2, when I left the tree.”

1 Zoologist, 1901, 153. ° Jrish Naturalist, 1905, 105-106. * Zoologtst, 1905, 307.

74 VESPERTILIONIDZ—NYCTALUS

Occasionally the Great Bat appears in broad daylight. Mr
J. W. Douglas observed’ one mobbed by swallows, the powers
of flight of which were, however, so clumsy compared with
those of the bat, that it soon left its pursuers behind. Again,
the Reverend A. Matthews told Mr Montagu Browne ‘that
one broiling hot day in July . . . at midday, when the
air was perfectly bright and clear, he observed swallows
circling at an immense altitude, and above them, at a much
higher elevation, four large bats, which he supposed to be of
this species.” On one occasion, writes the Reverend J. G.
Tuck,’ at about midday, a Great Bat suddenly dropped into a
pond about ten yards from the bank, and swam to shore with-
out the least difficulty. Whether the animal’s swim was under-
taken voluntarily or by mischance is open to question, but this
species, like others, loves to touch the surface of pools of water
over which it may happen to be flying.

When discovered in its diurnal retreat the Noctule will be
found cold and comatose, hanging head-downwards, but
tenaciously grasping the sides of the cavity with its feet.
Many may be huddled together, and frequently they will make
no attempt to escape, but when handled they rapidly become
warm and lively. They rise quite easily from a flat surface,®
exceptional failures being due to ill-health or overfeeding.

“The actions of the Noctules,’ wrote Gurney,* ‘ when
awaking from their diurnal sleep at the approach of evening,
are curious and grotesque. They frequently open and
shut their mouths for several consecutive seconds with an
exceedingly rapid motion of the lower jaw; this action is
succeeded by the tongue being protruded about the eighth of
an inch, and the lips being thus thoroughly licked. When
this is accomplished, a hearty yawn usually follows, the mouth
being opened in the process to its utmost width, and the
next employment undertaken is an attack on the small parasitic
insects which infest the fur of these bats. The sides of the
body are vigorously scratched by a rapid and continuous action of

1 Zoologist, 1843, 6; see also W. S. Lewis, Zoologist, 1848, 2252.

2 Zoologist, 1905, 231. 3 Gurney, Oldham, etc.

* Zoologist, 1874, 4194; see also Trans. Norfolk and Norwich Naturalists Soc.,
1869-70, 22 (published 1874).

THE NOCTULE, OR GREAT BAT 76

the hind claws, and the head is bent under the body, whilst the
mouth is employed in active investigation amongst the fur of the
under surface. These bats when fully awake usually begin to
crawl over one another, a process which generally evokes a
stridulous chirping cry from the individuals which compose the
lower strata of the cluster.”

Although the sexes fly together, the extent of their inter-
mingling in their places of sleep and hibernation has often
formed the subject of discussion. Gurney,’ working in Norfolk,
believed that when the females have young they are found in
separate colonies, which are larger than those of the males. This
opinion coincides with that formed by Leisler many years before
in Germany, as well as with the more recent work of Messrs
R. Rollinat and E. L. Trouessart.* It is also supported by the
recently recorded observations of Mr Booth, showing that both
sexes frequented the same den in about equal numbers until the
colony broke up for the summer in June and July, which is
about the time that the young are usually born. It is in agree-
ment with the known published records of the finding of both
sexes together, as by Mr Harting in February, by Daniell,
Mr Oldham, and Mr F. Norgate in May, and by Gurney and
Mr Whitaker in September.*® In the latter case, a small colony
was found on the 2nd of the month, and on the 5th contained
seven adult females and one male, which may be supposed to
have resumed companionship after the young had been reared.
Exceptional cases evidently occur, as when Mr Whitaker‘
met with fourteen individuals on 28th June, of which eleven
were males, one a pregnant female, and one a young male
aged about ten to fourteen days, and the mother of which was
presumably a bat which effected its escape.

There is no evidence on one important point from British
observations, but it may be presumed that, as shown to be the
case in continental Europe, the true breeding season is in
September, when the bats, having reared their young, are in
winter quarters awaiting the period of hibernation. The com-
mencement of gestation is, however, probably delayed until the
awakening of the animals in spring.°

1 Zoologist, 1887, 170. 2 Supra, p. 32. 3 Naturalist, 1905, 325-330.
4 In MSS. 5 Supra, p. 33-

76 VESPERTILIONIDAE—NYCTALUS

There is also some question as to a supposed preponderance
of females in this genus, a matter upon which it is

impossible to write definitely. Large numbers of females of .

the Noctule are certainly found not infrequently in their hiding-
places; but on the other hand we have the contradictory
experience of Mr Whitaker as related above, and that gentle-
man states that he and a friend had previously handled over
twenty-seven Noctules taken at different times and _ places,
without encountering a single female. In Ireland, again, Mr
Moffat points out that the males of Leisler’s Bat are more
frequently shot than the females, so that here once more the
available evidence agrees so little that more study is clearly
necessary. In this connection Mr Coward’s remark, as quoted
by Mr Millais, that in captivity the males are more active than
the females, is important, and may throw light on the relative
numbers in which the two sexes are encountered in nature.

Despite its offensive smell, the Great Bat is an object of
considerable beauty. Dr Henry Laver, indeed, contrasting
its rich brown fur, smoother and finer than velvet, with its
dusky wings, pronounces it to be the most beautiful of its order.
Although on the wing for so short a time, it is generally sleek
and plump, especially in autumn,’ when it accumulates much
fat both internally and as a thick layer under the skin, leading,
according to Mr Aplin, to a noticeable increase of weight as
compared with that of early summer.

All writers agree that,’ at least when new to captivity, the
Noctule, especially the male, displays much irritability, rest-
lessly biting the bars of its prison, its fellow-captives, and its
capturer, but confinement does not seem to interfere with its
health, since it may easily be kept alive for a period of some
weeks.

The first published account of this species when in
captivity, was that of George Daniell, who ascertained
many interesting particulars concerning its lactation and
parturition, and embodied them in a very pleasing essay.°

1 Gilbert White ; W. Borrer, Zoologist, 1874, 4125; Aplin, Zoologist, 1885, 344 ;
Dowker, Zoologist, 1891, 305.

2 F. R. Rodd, Zoologist, 1891, 347 ; Oldham, etc.

3 Proc. Zool. Soc. (London), 1834, 129-132.

———— a ee

PLATE IV.

2
be)

HEADS (natural size) OF

1. Nyctalus noctula. 2. Nyctalus lerslert,
3. Vespertilio serotinus.

“yt © Seana ee Sek

THE NOCTULE, OR GREAT BAT 77

Daniell’s observations show that he had observed, although
not with complete understanding, the use of the inter-
femoral membrane by bats as a pouch wherein to secure
their prey, and he particularly noticed that, unlike the
Pipistrelle, his captive Noctule made no use of this con-
trivance.

In feeding, he wrote, “the wings were not thrown
forward as in the Pipistrelle; and the food was seized
with an action similar to that of a dog: The water that
drained from the food was lapped, but the head was not
mused. in) drinking, as... . observed .. . in the Pipi-
strelle. The animal took considerable pains in cleaning
herself, using the posterior extremities as a comb, parting the
hair on either side from head to tail, and forming a straight
line along the middle of the back. The membrane of the wings
was cleaned by forcing the nose through the folds, and thereby
expanding them. Up to the 2oth of June the animal fed
freely, and at times voraciously, remaining during the day
suspended by the posterior extremities at the top of the
cage, and coming down in the evening to its food: the
quantity eaten sometimes exceeded half an ounce, although
the weight of the animal itself was no more than ten drachms,”
an observation which agrees very closely with those of Mr
Whitaker,’ who possessed a Noctule which on one occasion
actually devoured the enormous number of eight dozen
mealworms at a sitting, and which consumed an average
of about seven dozen, weighing in all about a quarter of an
ounce daily, or about a quarter of its own weight.

Of four females which came into Daniell’s hands on the 16th
of May, each was pregnant with a single foetus, but only one
survived to bring it to a natural birth. Of this one it
is related that on 23rd June, after exhibiting much rest-
lessness for upwards of an hour, she suddenly reversed her
usual attitude of suspension by the posterior extremities, and
attaching herself by her anterior limbs to a cross-wire of the
cage, gave birth to a young one. This, being born on its
back, passed into the interfemoral membrane, which was
expanded so as to form a perfect nest-like cavity for its

1 Naturalist, 1905, 325-330-

78 VESPERTILIONIDAE—NYCTALUS

reception. The young bat was at its birth perfectly destitute
of hair, and blind. As soon as it was born, the mother
“licked it clean, turning it over in its nest, and afterwards
resuming her usual position, and placing the young one in
the membrane of her wing, proceeded to gnaw off the umbilical
cord and eat the fAlacenta. She next cleaned herself, and
wrapped up the young so closely as to prevent any observation
of the process of suckling. The time occupied in the birth
was seventeen minutes. At the time of its birth the young was
larger than a new-born mouse, and its hind legs and claws
were remarkably strong and serviceable, enabling it not only
to cling to its dam, but also to the deal sides of the cage. On
the 24th the animal took her food in the morning, and appeared
very careful of her young, shifting it occasionally from side to
side to suckle it, and folding it in the membranes of the tail
and wings. On these occasions her usual position was reversed.
In the evening she was found dead; but the young was still
alive, and attached to the nipple, from which it was with some
difficulty removed. It took milk from a sponge, was kept
carefully wrapped up in flannel, and survived eight days, at the
end of which period its eyes were not opened and it had
acquired very little hair.”

Daniell’s remarks are in part borne out, in part opposed,
by those of Mr Whitaker,’ who has added not a little to our
knowledge of young Noctules. In Mr Whitaker's case a
female gave birth to her single young one on 3oth June.
Her attitude was the normal one, with the head downwards.
“As soon as ever the head of the baby Bat protruded it com-
menced to squeak lustily. The young was quite free in about
four minutes’ time, and worked its way under the shoulder of
its mother, and so round on to her back, where it clung quite
exposed, head downwards. . . . The note of the young
Noctule is a single chirrup even more highly pitched than
the note of the adult. It is so penetrating that when but a
few days old I could hear the young Bat calling after I had
gone to bed at night, though it was in a cage downstairs, and
in a room not directly under my bedroom.” For some time the
mother appeared disinclined to recognise her offspring, but at

1 Naturalist, 1905, 325-330.

THE NOCTULE, OR GREAT BAT 79

length on the third day, after she had been induced to eat
and was no longer hungry, it was accepted. ‘‘She first seized
it quite roughly in her mouth, and taking no notice of its
vigorous squeaking and struggling, proceeded to give it a good
wash and brush, much after the style of a Cat washing its
kitten. She then tucked it away under the skin between the
shoulder and thigh, pushing it so far round that it appeared
only as a protuberance on her back. This I found later was
the invariable method of carrying the young one, who was
packed away like this, head downwards, all along, and suckled
from this position by merely stretching its neck a little. For the
first week my Noctule always kept the young one tucked under
her right wing, as far as I saw, and I believe it suckled from
that side only. Later it put it under either wing indifferently.”

The young bat was quite capable of hanging by itself in
its cage, but sometimes the mother carried it with her, “in
which case the young one clung to her with its teeth
and was dragged along under her, trying to walk, with its
hind feet projecting from under its mother and close to
hers, so that she seemed to have four back feet, all working
out of time. The effect was decidedly comical. In spite
of this, the ‘baby’ did not seem to hamper its mother’s
movements very much when she was crawling. | When
disengaged from its mother the young one would cling with
extraordinary tenacity to the gauze sides of its cage, or toa
handkerchief on which we once or twice photographed it, and
great care was necessary to remove it without injury from any-
thing of which it had got a firm hold.”

When on one occasion allowed to fly with her young one,
the mother “hesitated a long while before making the attempt,
and when she did so only flew the length of the room, and
then dropped rather heavily upon the floor, the young one
remaining under her right wing all the time.”

After eleven days the young one, captivity having prob-
ably delayed its development, was still blind and naked, and
unfortunately on the night of that day, the mother having
escaped, it disappeared under circumstances suggesting that she
herself came back to the cage and stole it away.

According to British observations, the period of gestation

80 VESPERTILIONIDAH—NYCTALUS

exceeds thirty-eight days. To these should probably be added
another eleven,’ so as to complete the seven weeks laid down
by continental naturalists, but corroboration is still needed for
this country. The most usual time for the birth of the young
is evidently late in June. Thus MrH. J. Charbonnier found
within the body of a female killed in June a young one having
an expanse of wing of 33 inches, and Mr Oxley Grabham”’
writes me that he has received one, a dark, leathery-looking
object, just beginning to show indications of fur (and there-
fore at least fourteen days old), on 16th July. Gurney
informed Newman ® that he once disturbed a colony of females,
many of which had half-grown young ones with them, and
he observed that when the mothers flew away, the young ones
clung to their under sides, and seemed to be carried without the
slightest inconvenience to mother or young. On the other
hand, Mr Whitaker took a ten days’ old youngster from a hole
in a tree, where it remained after the departure of the adult
members of the colony, including its mother, so that it ig
evident that the young may either accompany their parents
when flying, or, when they have grown inconveniently heavy,
may remain suspended in the den (Plate III., p. 58). Whatever
else be the case, it is evident that the young must grow so
rapidly as to shift for themselves at an early date, and thus
relieve their mothers from burdens so tiresome. The increased
numbers which have been noticed in August, as by Mr Jeffrey
writing to Newman,* may probably have represented the influx
of the newly-fledged young, and Noctules exhibit marked dis-
agreement in size at this season.

Although there is only one published instance for Britain
of the production of more than a single young one at a birth,°
the statements of continental writers° are quite clear that fami-
lies of two have frequently come within their experience, a
remarkable instance of fertility, varying with locality, which is
discussed elsewhere (p. 37).

1 J.e., the true period from ovulation followed by fertilisation (not from copulation)
to parturition.

2 Naturalist, March 1899, 71. 3 Field, 7th March 1874, 218. 4 Jbid.

5 L. Jenyns, Observations on Natural History, etc.(London: Van Voorst), 1846, 56.

® As by Carl Vogt, who examined twelve females, in five of which were two, and
in six only one embryo (see supra, p. 31).

THE NOCTULE, OR GREAT BAT 81

The Great Bat has frequently been kept alive since Daniell’s
time. Mr Oldham’s description of its habits while in con-
finement is the most detailed, and may be quoted at length : * —
Like other bats in captivity, it ‘shows little inclination
for flight, especially in an artificially lighted room, and when
it does take wing, frequently collides with the walls and
furniture. A confined space is indeed unsuited to its bold
and dashing flight, although in a darkened room it will remain
on the wing for some time and avoid accidents. In walking—
a captive Bat’s usual mode of progression—the body is carried
clear of the ground, and supported on the feet and wrists only.
The tail is curved downwards and forwards, and the inter-
femoral membrane pressed against the belly. The fore limb is
spread considerably, but the phalanges with their connecting
wing-membranes are tightly closed and folded back along the
lower arm. In ascending a curtain or picture-frame, the claws
on the thumbs are brought into use, and the tail, instead of
being curved beneath the body, is then extended backwards,
with the tip pressed closely against the surface of the object up
which the Bat is climbing. For the time being, it is analogous
to the stiffened rectrices of a Woodpecker or Tree-Creeper.

«Any instinctive dread which Bats may have of man dis-
appears quickly in captivity, but the Noctule is exceptionally
fearless. Within a few minutes of their capture, I took two of
the Bats singly from among their struggling fellows in the bag,
and, holding them in one hand, offered mealworms with the
other. So cramped were they that they could not move their
limbs, but they seized and devoured the insects with the utmost
sang-froid. On the same evening others were climbing about my
arms and neck without any signs of fear; and the old female
which | had for several weeks used habitually to clamber up my
arm as it rested on the table, and snuggle against my neck.”

The captives drank freely. ‘‘Their food consisted of
mealworms ... , raw lean beef, and such moths, beetles and
other insects as I was able to procure. All food was thoroughly
masticated by an extremely rapid movement of the jaws before
it was swallowed. The wings of moths were generally con-
sumed, but the horny elytra of large beetles were bitten off and

1 Zoologist, 1901, 51-59. y
I

82 VESPERTILIONIDEZ—NYCTALUS

allowed to fall as the insect disappeared in the Bat’s mouth.
Mealworms and small moths, as well as...” beetles of lesser
size, were seized and eaten without any attempt to overcome
their struggles. On the other hand, large moths were some-
times, and powerful beetles always, thrust by the bat into the
pouch formed by the interfemoral membrane, in order to secure
them effectually before they were eaten. A cricket offered
to one of Mr Coward’s bats was treated in the same way, but
cockroaches were in some instances thrust into the pouch,
and at other times eaten without that preliminary. It
should be remarked, however, that cockroaches, despite their
size, submitted very tamely to their fate. On no occasion was
foot, carpus, or thumb used to secure or dismember prey.’

Mr Oldham’s bats seemed unable to see food if held but
little more than an inch in front of them, and this was the case
in natural twilight as well as in an artificially lighted room.
He, therefore, thinks it very doubtful whether in a free state
they would avoid a distasteful moth, because its nauseous
properties happened to be advertised by warning colour, but
Mr Forrest remarks’ that this does not prove that the Noctule
may not be “far-sighted, and see an insect in the air at a
distance of several feet better than close at hand on the table.”
It is, however, interesting to note that no amount of per-
suasion would induce the bats to eat notoriously unpalatable
insects.* These, although repeatedly offered, were invariably
rejected with disgust, as was an oil beetle in a similar experi-
ment tried by Mr Coward. Two other moths* also appeared
to be unpalatable, but in a lesser degree.

This bat is one of the largest of the species frequenting
Britain. Two others alone, the Serotine and the Greater
Horseshoe, are in this respect its equal, or occasionally
its superior: their flight is, however, distinct, and_ their

! The following, among other species of moths, were readily eaten by Mr Oldham’s
bats while in captivity :—MWamestra persicariea, Leucania pallens, Hepialis humuli,
1. sylvinus, 1. hectus, Rumia crategata, Urapteryx sambucata, Odontopera bidentata,
Fidonia atomaria, F. piniaria, Xylophasia polyodon, and Amphidasis betularia; the
list may be completed by the Poplar Hawk Moth, Smerinthus popult, with one of
which species Whitaker fed his captive Noctule (Waturalist, 1905, 325-330).

” Trans. Caradoc and Severn Valley Field Club, 1900 (Feb. 1901), 244.

3 As Euchelia jacobee or Abraxas grossulariata.
4 Spilosoma menthastri and S. lubricipeda.

LEISLER’S BAT 83

peculiarities will be pointed out in the proper place. With
these exceptions it could be confused only with the much smaller
Leisler’s Bat. Unlike some other species, the Noctule flies
with the tail directed straight backwards, or with only a very
slight downward curve, a fact which may be supposed to have
some connection with the less frequent use of the interfemoral
pouch than in smaller and less powerful species.

LEISLER’S BAT.
NYCTALUS LEISLERI (Kuhl).

1810, DIE RAUHFLUGLICHE FLEDERMAUS, T. P. Leisler, Magazin fir die Neuesten
Entdeckungen in der Gesammten Naturkunde (Berlin), 156; described from
Hanau, Germany.

1819. VESPERTILIO LEISLERI, Heinrich Kuhl, Avm. der Wetterauische Gesellschaft
fiir die gesammte Naturkunade, iv., 46, naming Leisler’s Rauhfliigliche fledermaus ;
Jenyns ; Bell (ed. 1); Clermont ; Newman.

1819. VESPERTILIO DASYKARPOS LEISL. (séc), Heinrich Kuhl, of. cé¢., 49, quoting
Leisler’s unpublished MSS..

1829. PTERYGISTES LEISLERI, Jakob Kaup, System der Europitschen Thierwelt, ic,
100; C. B. Moffat (doubtfully), Zrish Naturalist, 1905, 104; Miller; Thomas ;
Johnston ; Méhely ; Cabrera; Millais, 8 and 76.

1838. SCOTOPHILUS LEISLERI, J. E. Gray, Mag. Zool. and Bot., 497 ; MacGillivray ;
Bell (ed. 2).

[? 1839. VESPERTILIO PACHYGNATHUS MICHAHELLES (sic), J. A. Wagner’s ed. of
J. C. D. von Schreber’s Die Sadugthiere, Supplement i., pl. lv.6. There is no
description, but Fitzinger states that Michahelles found this bat in Dalmatia:
it appears to be a small WVyctalus.]

1839. VESPERUGO LEISLERI, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s
Archiv fiir Naturgeschichte, i., 318; Blasius ; Fatio; Dobson; Blanford ; Alcock ;
Moffat.

1856. PANUGO LEISLERI, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung
(Dresden), Neue Folge, ii., 131, 172.

1870. NOCTULINIA LEISLERI, L. J. Fitzinger, Sztzungsberichte der Kaiserlichen Akad.
der Wissenschaften (Vienna), Ixii. (1), 218; Harrison Allen, Proc. U.S. National
Museum, xvi., 30, footnote, 1893.

1898. PIPISTRELLUS LEISLERI, Oldfield Thomas, Zoo/ogzs?, 100.

Le Vespérien de Leisler of the French (Fatio), de vrauharmige
Fledermaus of the Germans (Blasius) ; but these are merely book-names,
as is the Hairy-armed Bat of Bell and others, there being no local
names for such a little-known species.

Distribution :—Leisler’s Bat, or species closely resembling it, is found
in the wooded districts of boreal and transitional Europe and Asia, from
sea-level to 4500 feet (Fatio) in the Alps, from middle Russia to Greece

84 VESPERTILIONIDA—NYCTALUS

(Winge), and probably the shores of the Mediterranean generally, and
from Ireland probably to China. As in the case of JV. zoctula, the exact
limits of its range are very imperfectly known, but allied forms (men-
tioned on page 54) are known from the Azores, Madeira, the Hima-
layas, and the Oriental Region.

The distribution of Leisler’s Bat is still very imperfectly known
for Great Britain, where it ranks as one of the rarest species. Its
admission to the British list rested for many years upon the slender
evidence of a single historical specimen in the British Museum, figured
by Bell in both his editions.

To this record Tomes, before 1874, added a specimen in the
Bond collection, taken near Cirencester. Tomes himself not unfre-
quently observed the species along the course of the river Avon,
in the counties of Warwick, Worcester, and Gloucester, and therefore
concluded that it must be less rare than had hitherto been supposed.

More recent study has added a few county records. Thus, for
Yorkshire, Bond informed Newman (/7ze/d, 7th March 1874, 218) that he
had seen specimens taken from a chimney-shaft near Leeds: Clarke
and Roebuck give the date of these as about 1840, and the exact
locality as Hunslet (see Grabham, WVaturalzst, 1899, 72); one of these
was in Bond's collection, and at his death passed into the possession of
Harting. Charbonnier obtained seven shot at Mexborough in 1890, one
of which is now in the British Museum (Zoologist, 1892, 329); lastly,
Armitage and Whitaker have found the species not uncommon near
Barnsley and Wakefield (Coward, Zoologist, 1905, 68; Armitage,
Naturalist, 1905, 37-38, and Whitaker, 1907, 384-388, and 416-418).

In Cheshire, Coward shot one in Dunham Park in 1899 (Zoologzst,
1899, 266). He has occasionally observed the species since, and believes
that it is more abundant than is generally supposed.

There is an old and doubtfui record for Norfolk: T. Paine, jun.
(Ann. and Mag. Nat. Hist., Nov. 1838, 181-183, published 1839), identi-
fied as referable to this species fourteen bats taken from a hollow
tree near Norwich. Jenyns, who examined one of them, declared
(Journ. cit., 481) that it was “not a specimen of V. /ezslerz,”? but was
undecided whether it was “the young of a Noctule or a distinct species.”
Apparently no skull was submitted to Jenyns, but the description and
dimensions given by Paine agree so well with those of JV. xoctula that
I have no hesitation in connecting his specimens with that species.
Possibly the band of hair on the arm, which occurs in JV. zoctula as well
as in JV. lezslerz, misled both Paine and Jenyns.

The bat is thus known to occur only in three isolated districts—
the West Riding of Yorkshire, Cheshire, and the valley of the Avon—
in Great Britain; judging, however, from its abundance and extended
distribution in Ireland, there seems to be no apparent reason why it

PLATE V,

‘quiny} pur 3003 jo AT[etoadsa ‘azis ut
QUdIAYIP PATV[A1 9Y} MoYs 0} ‘uUIOG A[jUad—aI *(Z) wyMj20u “Ay pur ‘(1) YyNpe ‘zcapszay SN]OJIAN

PLATE VII.

WINGS (4 natural size) OF

1. Myctalus leislert. 2. Pipistrellus pipistrellus.
3. Myotis daubentoni

EEISLER’S: BAT 85

should not occur in other parts of England, Wales, and even in Man
and Scotland (see Service’s observations under JV. xoctula).

This bat is well known in Ireland, where the first recorded was taken
in Belvoir Park, Co. Down, about 1848. A second was knocked down
by a mechanic with a fishing-rod in Blackstaff Lane, Belfast, Co. Antrim,
in July 1858. It passed into the hands of Darragh, and was exhibited
by J. R. Kinahan at a meeting of the Belfast Naturai History
and Philosophical Society on 25th April 18601 In 1868 the
species was found abundantly in Armagh by Barrington, who also
received a specimen from Kildare (Zoologist, 1874, 4071-74), and it
has since been discovered in Dublin (Ogilby, Journ. czt., 1874, 4236),
Wicklow (Barrington, Journ. cit., 1875, 4532), Fermanagh (Barrington,
Journ, cit., 1883, 116), Westmeath (Moffat, /rzsh Naturalist, 1897, 135),
Cavan, Louth (Jameson, /ourn. cit., 1897, 41-42), Londonderry (Alcock,
Journ. cit., 1899, 174), Carlow and Wexford (Barrett-Hamilton, Journ.
ctt., 1900, 134; Moffat, 162; Pack Beresford, 1906, 194). Its range,
although a few county records, eg., for Kilkenny, Meath, and
Monaghan, are needed for continuity, may be said to include the
whole east coast, together with a considerable portion of the north
and north-west. In many districts it is common, or even abundant,
and, although no western or south-western specimens are forthcoming,
R. E. Dillon (in lit., also quoted by Jameson) is well acquainted with
large, straight-flying bats, almost certainly of this species, at Clonbrock,
Co. Galway, and Laver informs me that he has seen many in the same
county ; one, labelled Tyrone, is in the Dublin Museum. It is probable,
therefore, that the bat has been overlooked in many parts of Ireland.

Distribution in time :—This bat is not known as a fossil.

Period of gestation :— Although no observations have been published,
there is no reason to suppose that this differs from that of the Noctule.

The number of young, although stated to be usually two in Germany,
is not known to exceed one in Britain (see Tomes, Zoologzst, 1854, 4365),
born early in summer, probably in June.

Description :—This bat resembles the Noctule, but is much smaller.
The body is relatively lighter, and the bones of the limbs less massive
and of somewhat different proportions.

Thus in the wing (again as compared with the Noctule) (Plate VIL.,
Fig. 1), while the greatest expanse, third metacarpal and longest digit
are relatively of smaller dimensions as compared with their forearm,
the lower leg is markedly, the tail and fifth metacarpal slightly, longer,
so that the wing and interfemoral membrane are slightly shorter and
distinctly broader.

1 The first published proceedings of the B. N. H. and P. Society date from 1871,
but Robert Patterson has most kindly looked up the present reference for me in the
old minute-books.

86 VESPERTILIONIDA—NYCTALUS

The foot is comparatively feeble, the post-calcarial lobe less con-
spicuous, and its outer margin not thickened (Plate V., p. 64).

There are two phases of colour. In the one represented by an Irish
female, dated 8th September, the fur is so dark—between “olive,”
“bistre,” and “sepia”—as almost to reach “clove brown,” the hair-tips
lighter ; in the other, represented most frequently by males in spring
and early summer, the hairs have a conspicuous rusty tip of almost
a third of their length, giving the bat a much more zoctuda-like appear-
ance: in this phase specimens vary between an intensified “wood
brown” and some shade of “mars brown.” The two phases are not
distinctly marked off, and may intergrade; in both, the under side is
lighter. The membranes, limbs, and ears are dusky, as in JV. xoctula.

Dr Alcock states that he sometimes, but not always, found Irish
males to be darker than their females, and the same remark is said to
apply to the young.

Nothing is known of the processes of moult, or of a seasonal colour
change, but a series of fourteen examples (seven of either sex) of
the allied WV. azoreum of the Azores suggests that the rusty phase
may represent the fading of the old winter coat in spring, and the
dark phase its replacement by a new pelage—a process which, as in
many other mammals, is characterised by much irregularity. The
Azorean specimens were obtained
by Ogilvie-Grant and N.C. Roths-
child between 8th March and 17th
May. ‘The females are in fully
tinted pelage, but the males, ex-
cept one, are much faded, as if in
very old coat; the exception,
killed on 17th May, appears to
be in process of acquiring the new
pelage, perhaps already donned by
the females. Similarly, a Rou-—
manian male of 22nd April is in
rusty faded pelage.

The skull is less than half the
size of that of JV. xoctula, its struc-
ture is weaker and more papery,
and the crests, especially the
sagittal, are absent or feebly de-

5 FIG. 7.—FRONT VIEW OF INCISORS AND
veloped (Fig. 16, No. 1, p. 174). CANINES OF Nyctalus /eisleri (enlarged and

The teeth are similar, but diagrammatic).
much smaller. The outer upper
incisor is comparatively small, its cross-section at the base being about
equal to, zot about doud/e, that of the inner incisor. The anterior upper

LEISLER’S BAT 87

premolar is relatively larger, and the /ower zucisors, being smaller, are less
crowded and do not usually overlap (Figs. 7, p. 86, and 16, No. 1, p. 174).

No material exists upon which to base any account of either the
individual or exceptional variation of this species, but Moffat informs
me that there are two melanic examples in the Dublin Museum.

Geographical variation, as in JV. xoctula, shows itself mainly in
regard to size and cranial characters, especially in the relative
development of the crests (for particulars, see above, p. 54). I have
carefully compared the Irish and English specimens in the British
Museum, but cannot distinguish them.

DIMENSIONS IN MILLIMETRES :—

4 » : S
we)
> op a 5 Es ee = é =
3 g ~ 8 » |S = 3 = = Z
» Pale 2 Eo | 3 io} 3 & Aan
2/8 |#e\21 8 |22) 2/3! 8 | |) ea) 88
3 3 Bales Bs °,.8 Fa © s 9 »s
3 2 | as 6 |/ee| eo 1% a & £ a
3 Sp 3 5 ey =| 5 we) > 2
o 2 ce) =] 2 3
= & es I HE A SI = $
ae 8 =] eS he
3 a0) o
Leach’s specimen in
British Museum (No.
d. 63a) . : 5 : a ae 48 Be is ais 43
Male, Mexboro’, Yorks.
(H. J. Charbonnier, No. |
90. 7. 6. 1. of British
Museum collection) “ sd ae Wy 46 ee “6 43
Male (in alcohol), Bray,
Ireland (R. M. Barring-
ton per J. E. Harting,
No. 90. 2. 14.1. of British
Museum collection) oi ai ws ie o6 || ite) 9 42 We Me 38 82°5
Male, Co. Clare, Ireland
(Dublin Museum) . fe Ar e 50) || GS [he 1 42 se ae 40°2 | 32
Male, Co. Dublin, Ireland
(Dublin Museum) . ae Se S 5m Il. 1s 8 42°5 40 31
Male, Co. Louth, Ireland
(Dublin Museum) . ae ae on Reo 8 40 37 30°5
Female (in alcohol), Ar-
magh, Ireland (R. M.
Barrington) F 57°5 | 14°5 | 6 we 18 7b | 42 6 71 36 30
Female, Co. ‘Armagh, Ire-
land (R. M. Barring-
ton per G. E. Dobson,
Wooes9., 11. 12. 5. of
British Museum collec-
tion; labelled “ var.
hibernicus ” in Dobson’s a
handwriting . 61 14 4 Ao ant) 9 44 “3 74 39 33
Female, Belfast, Ireland
(R. Patterson per G. BE.

H. Barrett - Hamilton,
Momo. 3: 16. 1. of
British Museum collec-

tion) 62°5 | 17 + A ee Sa 43 oe Se a es *295
Female, Co. Dublin, Ire- (approx.)

land (Dublin Museum) . 36 36 ae aay |e 8 44-5] .. oe 39 33 St
Female, Co. Dublin, Ire-

land (Dublin Museum) . Be a fe rata bes 8 43 aC “s 39°5 | 31

Male, immature, Co.
Cavan, Ireland (Dublin 9 5 9
Museum, in alcohol) 3c . oe | oes | 16° 89 |: os Sia

Male, immature, Co. (approx.)
Dublin, Ireland (Dublin x = 9
Museum, inalcohol) .| .- . oy [wate '|| 2 75 | 385] .. = os >

Average of adults g : ; ey
Papacy \ 59°5 | 15 45 seal lls AS 72°5 88°5 | 31°5

fo 4)
~
bo

* 118 inches in the flesh, fide R. Patterson, corroborated independently by Whitaker, who gives the
average wing-expanse of six Yorkshire females as 304 (Naturalist, 1907, 386).

88 VESPERTILIONIDAZ—NYCTALUS

For the following table of measurements I am indebted to Charbonnier.
They were printed on page 65 of Lloyd Morgan’s Animal Life and
Intelligence (London), 1890-1891 :—

Pig Digit III. Digit IV. Digit V.
Tiss
mk Ve Ved Vac ine re Pe rapes ete ¢.| ec
Bye ye |e 8 | ee toe) Bee eo eee
Svlves Waste) tea ths ceed Abaa | bts) (cet alates
Se A la be Se = a ican hae
41 |65| 88 | 40 | 36 | 19 | se | a4 | ¥ | 82 | 8)7 | te
41 6 38 40 16 19 39 155 Uf 33 8 |6°5| 16
Males . :
au 1@ |.89 (1740. | 6. a8: e9° |e” |! el es |e ciee tees
415|5 | 89 | 405|17 | 20 | 39 | 16 | 7 | 883 | 847 | 5
ao \6 | so | a7 | 166118 | 87 -| 445 | -7! | 82 Fos Vepieas
Females .4| 41 [5:5] 385] 39 | 165120 | 39 | 15 | 75] 33 | 8 |7-5] 17
Va 6 | 39 | 40 | 15:5] 205| 39 | 155] 7 | 38 | 8 |7 | 16
Averageof7 . . | 40°9/5-7| 386 | 39°5| 16 | 19:2| 385] 152| 7 | 32-7] 8 bi 15°8

Proportionate lengths :—Foot (without claws), about -45 of lower
leg; fifth metacarpal, about -82 of third; lower leg, about -41 of fore-
arm, and about -29 of head and body.

Skull :—Greatest length, 15 ; basal length in middle line, 11-25 to 12;
palatal length in middle line, 5-5 to 6; from posterior border of m?
to anterior border of canine, 6-5 to 6; ditto in lower jaw, 6 to 6-5;
greatest breadth at zygoma, Io to 10-5; posterior breadth, 9; breadth
between orbits, 8; breadth at constriction, 5 (nearly).

Weight :—14 to 20 grammes (Moffat, /rzsh Naturalist, 1900, 235).

Distinguishing characters :—Lack of familiarity with this bat has
led most naturalists to regard its identification as difficult, and even
to confuse it with JV. zoctula. The two, however, when closely compared
are remarkably different, and, apart from the distinctions based upon
dimensions, colour, and odour, the relative proportions of the wing,
lower leg, and foot, are infallibly diagnostic (Plate V., p. 64). The only
other British species which bears any resemblance to Leisler’s Bat is the
Pipistrelle, but here again there is no need to search farther than the
length of the forearm for points of distinction.

The bat on which Bell, following Leisler, bestowed the
not very satisfactory name of Hairy-armed, was first dis-
covered by the latter naturalist in Germany, and described by
Kuhl in 1819; it has ever since remained one of the least
known of English bats. It made its first appearance as a

LEISLER’S BAT 89

British mammal in the first edition of Bell, whose figure, taken
from a single specimen in the British Museum, was the first
representation of the species.

We are now familiar with the appearance and habits of this,
at least in Ireland, locally abundant species. In England,
however, its distribution is still very imperfectly known, but
its general reputation for rarity, judging from the late R. F.
Tomes’s observation of it at various localities in the valley of
the Avon, is probably exaggerated.

Inability to distinguish it from the Noctule—of which,
in all external characters, except colour and gdour, and
in many of its habits, it has long been considered a duplicate
in miniature— and the very short period of the day
during which it may be observed upon the wing, have
probably contributed to neglect of this bat by English
naturalists ; whereas by the Irish, who are not troubled with
the confusion created by the presence of the Noctule, it is more
easily identified and brought under observation. As a matter
of fact, the two are so distinct when in the hand, and the
shape and proportions of their wings are so different, that it is
likely that, now that these characters have been published, closer
study will reveal corresponding divergences in habits, and lead
to means of identification on the wing.

Until the year 1899, there existed no published account of
the habits of this species in English other than the few lines
inserted by Tomes in Bell’s second edition. ‘‘ Previously to
1849,” wrote Tomes, ‘‘a Bat had often been observed which,
from its smaller size and different mode of flight, appeared to
be quite distinct from the Noctule; but it was not until June
of that year that an opportunity occurred of examining a
specimen; the difficulty of obtaining specimens arising not
so much from its rarity as from its general habit and style of
flying. Whilst the Noctule may throughout the whole of the
summer be seen taking its regular evening flight, night after
night, near the same spot, the Leisler’s Bat, on the contrary,
will be seen once, perhaps for a few minutes only, and then
lost sight of. It appears to affect no particular altitude in its
flight any more than it preserves a regular or prescribed beat.

When the weather is fine, you may see this bat passing on in
K

90 VESPERTILIONIDAE—NYCTALUS

a kind of zigzag manner, apparently uncertain where to go,
generally, though not always, at a considerable elevation, and
in a few minutes it is gone.”

Such was the meagre sum total of our knowledge, until
in June 1868 Mr R. M. Barrington happened upon a large
colony in the beech woods at Tandragee, County Armagh,
Ireland." Thence he procured specimens in 1874 and again
in 1889, in spite of which the bat remained for many years
immersed in an obscurity which was at length only dispelled by
the labours of Mr H. Lyster Jameson’ on its distribution,
and by the excellent original observations on its habits by
Dr N, H. Alcock* and Mr C. B. Moflat* “The latteniayae
probably the first British naturalist to undertake a series of all-
night watches in the haunts of the bats, thus proving beyond
question the existence of two short flights, the one vespertinal,
the other matutinal, in this species, exactly as Mr Charles Oldham
has since shown to be also the case with the Noctule.

Like the Noctule, Leisler’s Bat is usually an inhabitant of
woods,’ where it frequents, often in large parties, the cavities of
hollow trees, communicating to them an odour which, how-
ever, is much less perceptible than that of its congener. Its
numbers evidently rival those of the Noctule, since Mr
Barrington estimated the strength of one band at from eighty
to a hundred, the combined squeaking being sufficient to cause
their discovery. In addition to trees, it retires also to the
roofs and recesses of buildings or walls. Mr Barrington®
found great numbers in the roof of a boat-house in County
Fermanagh; Mr P. W. Finn sent me one taken from a hole
in a barn in Carlow, while a party for some time occupied
a hole in the stable wall at Kilmanock, County Wexford.’
Extreme darkness would not appear to be a necessity for
its comfort, since the colony discovered at Tandragee were in
full view from without, and Mr Finn’s specimen was discovered

1 Zoologist, 1874, 4071-4074. 2 Trish Naturalis#, 1897, 41.

3 [bid., 1899, 169-174, and map. 4 Jbid., 1900, 235-240 ; 1905, 99-101.

5 Arthur Whitaker found a den in Yorkshire, in an oak at a height of about forty
feet from the ground, which elevation he states to be greater than any Noctule den
known to him (aturalist, 1907, 385). If it should prove that this bat has a preference
for inaccessible retreats, its reputed rarity might thus be to some extent explained.

5 Zoologist, 1883, 116. 7 Trish Naturalist, 1900, 134.

LEISLER’S BAT “Ol

while asleep in the sun at the edge of its abode. Mr Moffat
also informs me that he knew of one which slept in a hole ina
birch-tree, in a situation where the morning sun shone full on
its face.

Although fond of congregating in colonies, solitary indi-
viduals are frequently to be found. Mr Moffat has met with
them from the middle of May to the end of September, and he
suggests that our information concerning the number leading
a gregarious life may be quite misleading, since it is the
largest and noisiest assemblages that are the most easily
discovered.

This bat would appear to fly for a shorter period of
summer than any other known British species. In the
County Dublin it does not usually appear on the wing until
about 20th April, but by the end of that month it may
be seen in numbers, so that it is probable that the hardier
individuals may appear earlier in the month. It hibernates
early, retiring abruptly on or about 26th September, a date
arrived at independently by Dr Alcock at Dublin and by Mr
Moffat in Wexford. The abruptness of its disappearance is
remarkable; thus, in 1899, at Ballyhyland, County Wexford,
Mr Moffat made the following observations, as quoted from his
journal: ‘“September 25th, usual number flying; 26th, only
one; 27th, none”; the maximum temperature for the three
dates being respectively 574°, 53°, and 49° Fahrenheit.

Although hibernation commences early, it is liable, as in
the case of the Noctule, to interruptions dependent on the state
of the weather. Mr Finn’s specimen was taken on a2ist
October, and had been flying with regularity for the previous
fortnight, sallying forth ‘every evening to the minute at the
same time.” Mr Moffat has seen many on the wing in
Wicklow on 17th, and a single individual on 20th October,
and the late A. G. More identified one captured in a bedroom
on 16th November.’ Mr Moffat writes: ‘‘ The lowest tempera-
ture at which I have noted Hairy-armed Bats in the open is
46° (on October 8th, 1899), but on that evening they were out
in some numbers. I therefore think that this species is more
influenced by the heat of the day than by that of the actual

1 Moffat, Zr7sh Naturalist, 1897, 135.

g2 VESPERTILIONIDAX—NYCTALUS

time of flight ; for I have never seen it when the day’s maxtmum
was below 523°, and never xuerously with a lower maximum
than 56° Fahr.”

In 1899 Dr Alcock’ stated his belief that the activity of
Leisler’s Bat is probably restricted to a single hour's flight each
evening, commencing about fifteen minutes after sunset, ten
minutes earlier in spring, and ten minutes later in autumn.
The history of the discovery of the exact limits of this
vespertinal, together with the existence of a matutinal flight
as well, is so peculiarly Mr Moffat’s own, that I feel that I
should be doing him an injustice did I not quote his description
at full length.

‘For a mammal to enjoy so short a period of daily activity
as this one hour,” wrote Mr Moffat in 1900, ‘‘would be very
singular, and during a recent visit to Fassaroe [Co. Wicklow]

I obtained evidence that the flight is not so strictly
limited, for on July 22nd I saw several bats of this species flying
in the early morning :—the first and last being noted respec-
tively at 2.56 and 3.36 a.m. or 65 and 25 minutes before
sunrise. That they were Hairy-armed Bats I was certain;
but specimens could not then be procured.”

However, the discovery that Leisler’s Bat ‘is a morning
flier was a useful step towards tracing some of these animals
to their sleeping- place at Ballyhyland, the morning light
being infinitely preferable to that of evening for such a
purpose. On the morning of August t1oth I accordingly
watched from 2.45 a.m. in the pasture-field already referred to,
taking my stand near an old and hollow ash-tree which I had
reason to suspect the bats might inhabit. At 3.35 a.m. their
screams were audible in the open, where several were soon
afterwards seen, hawking and sporting above the level of the
tree-tops. At 4.2 one of them suddenly dipped from its eleva-
tion, and when near the ground darted towards the trunk of the
old ash, where it entered a hole about seven feet from the
ground. At 4.5 another dipped, and shot into the same hole.
At 4.11 a third followed suit; at 4.12 a fourth entered another
hole, some two feet higher up in the same tree. This was
the last bat seen that morning. The four individuals noted

1 Trish Naturalist, 1899, 169-174, and map.

i
tat

ats

SK: :

-<
Fp tb ta
ot o

PEATE I

PORTIONS OF FINGER SKELETONS OF BATS (diagrammatic), ENLARGED
TO SHOW STAGES IN OSSIFICATION OF THE JOINTS.

m, Metacarpal ; £41, phalanx 1; p42, phalanx 2; me, metacarpal epiphysis; p/1¢, first
phalangeal epiphysis ; £42, second phalangeal epiphysis.
A. Young specimen with epiphyses quite distinct from phalanges and metacarpal.

B. Older but still immature specimen, in which ossification, although advanced, is incom-
plete, and the outline of all the bones composing each joint is visible.

C. Fully mature specimen, in which all trace of the epiphyses has been lost.

LEISLER’S BAT 93

had homed respectively 33, 30, 24, and 23 minutes before
sunrise."

“The hole into which three of the four bats had vanished
was evidently the entrance to a cavity of some extent, but it
was too narrow to admit of any examination of the interior.
In the evening I saw the three come out again, one at 7.53 and
two at 8 P.M., or 16 and 23 minutes after sunset. On the
following evening five were seen, all issuing from the same
hole, between 7.47 and 7.57—their times of emergence being
respectively 12, 16, 17, 19, and 22 minutes after sunset. On
the 12th I again saw five come out: the first oze minute, the
others respectively 8, 11, 13, and 16 minutes after sunset.
When quitting their abode, though high fliers at other times,
these Bats skim very low over the grass.

“The question of the animal’s return to its sleeping-den
during the night was less easily settled. On two evenings I
watched the hole without any success, though the moon was
full and bright, and I hid in the shadow of the ash-boughs not
to disconcert the homing bats. The fact, however, that this
large and noisy species suddenly ceases to be either visible or
audible about an hour and twenty minutes after sunset weighed
strongly against the idea of its continuing on the wing all
night. Other considerations also pointed to the probability of
its retiring early, as the Noctule is known to do... . Dr Alcock
had drawn my attention to the fact that Hairy-armed Bats
shot by him an hour after sunset had their stomachs so
crammed with food that it seemed a physical impossibility
they could feed much longer.

‘“T therefore argued that since I could not detect the bats
going zz in the evening, I must endeavour to catch them as
they came owt in the morning. This would at least prove the
fact, though not the time, of their retirement after dusk. So,
on the night of August rath, I fixed a net at midnight over the
mouth of the hole. In the early morning hours of the 13th
I watched by this net . . . and at 3.15 a.m. heard a Bat gently
flop into it. I found that I had secured a fine female...

1 “Tn explanation of these figures, I should state that at Ballyhyland (lat. 52° 31’ N.,
long. 6 43’ W.) sunrise on August roth is 4 minutes later than at Dublin, and
sunset I minute earlier.”

94 VESPERTILIONIDA—NYCTALUS

caught emerging for her morning flight, one hour and twenty-
six minutes before sunrise. |

‘Having made sure of my specimen, I quickly removed the
net so as not to disturb the remaining inmates of the hole in
their egress. I failed to see any bats emerge—the light being
too dim—but at 4.20 had the satisfaction of seeing one re-enter.
As this individual cannot have left the hole while the net was
over it, the duration of its flight had not exceeded sixty-five
minutes.

“In the evening of the same day I saw four bats (the
survivors of the colony of five) emerge for their evening flight
in quick succession between 7.38 and 7.42, 6 and 10 minutes
after sunset; and at 8.53 by a fortunate chance, the moon's
rays falling full on the line of flight at the moment, I succeeded
in seeing one go in. This was one hour twenty-one minutes
after sunset. As the interval between earliest and latest
emergence had been so short, the duration of this example’s
flight had been told with some exactness. It cannot have been
on the wing for a Zonger period than 76, or a shorter one than
71 minutes.

‘““We may thus claim to have a set of data, limited in
number, but precise as far as they go, determining the flight-
time of this local and somewhat imperfectly studied species. It
has an evening flight and a morning flight, the two being of
about equal duration. The evening flight is usually commenced
a little earlier than 15 minutes after sunset, the average of
seventeen actual emergences noted being 13% minutes, the
earliest I minute, the latest 23 minutes, and the mean between
the extremes 12 minutes after sunset. The bat returns at night
to the same hole as serves it for a sleeping apartment by day ;
the precise time of its return, in the only instance noted, having
been 81 minutes after sunset. Its moment of leaving the hole
in the morning has also been noted only once, in the case of
the specimen caught on August 13th; but it should be
remembered that this individual must have been the fs? to
emerge on the date in question, and therefore the usual time
of emergence is probably a little later than 86 minutes before
sunrise. The time of going home in the morning, on an
average of five observations, is 26 minutes before sunrise, the

LEISLER’S BAT 95

-earliest and latest instances having been 33 and 21 minutes
before the sun. It has been shown that the duration of one
individual’s (evening) flight was at least 71 minutes, whilst that
of another’s (morning flight) was not more than 65 minutes.
The usual duration is, in all probability, not far from the mean
between these two figures. The animal, therefore, in summer,
spends one-tenth of its time on the wing, and the remaining
nine-tenths in its sleeping-hole. In the shortness of its flight-
time it is probably unique among Irish bats.”

In its hibernation this species appears to differ from the
Noctule in that, instead of collecting in numbers for the
winter sleep, the slight evidence which we possess seems to
indicate that it scatters and hibernates singly. Mr Moffat,
watching the daily flight of the bats in September, found that
early in the month they deserted their summer quarters and
bestowed themselves in separate crevices in the higher parts of
the same tree. They migrated singly on different dates. ‘On
Sept. 1oth three (instead of four) emerged from the common
den; on the 11th, two; on the 13th, none. This shows that
they changed their abode voluntarily; if they had been
disturbed or alarmed, they would all have left at once. They
still fly every evening from their new quarters. No two inhabit
the same hole,” a description borne out by Mr Finn’s solitary
Carlow individual, then probably in its winter quarters.

The flight of Leisler’s Bat resembles that of the Noctule,
but, as stated above, Tomes believed it to be a little less
regular in its haunts and less definite in its area of flight than
the larger species ; and in Wexford, where I have seen it flying
_ at a considerable elevation, I cannot be sure of finding it in any
particular spot. Further, the colony in the stable wall resented
disturbance, and has departed. Mr Moffat, however, informs
me that in localities where it is abundant, as at Enniskerry in
Wicklow, it may be observed evening after evening in fairly
numerous groups. _

At his own home at Ballyhyland, Wexford, the bats on
emerging from their den go off almost invariably in the same
direction. He found when he intercepted their line of flight that
they passed certain points more than a quarter of a mile away
with great regularity, still dashing along as if bound for a fixed

96 VESPERTILIONID4:—NYCTALUS

destination ; which shows that they range to some distance in
search of food. They are fond of frequenting cattle pastures ;
but here not only, like the Noctule, do they vary their hunting-
grounds with the season, but also with the hour, having
‘ apparently regular feeding-grounds both for the early and late
parts of the evening. Thus the pasture attracts them to a
certain extent from May to September ; but until midsummer,
when the cattle occupy it, the bats fly in it for only a few
minutes each evening, just before retiring for the night.
During the late summer months the half-hour before retire-
ment is spent in careering about the field.

Dr Alcock writes of this bat’ that ‘‘at first it commonly
flies at a considerable height, in open country taking long
sweeps and wide zigzags, often being seen but once in an
evening. Near woods and in favourable localities it will often
remain for some little time near one spot, flying at an altitude
of 30 to 40 feet, with a faster and less irregular flight than the
Pipistrelle, the tail being extended at a straight line with the
body. Later on, it flies near the ground, very commonly
shrieking loudly, and I have observed two Bats at this time
chasing one another . . . both flying very fast, and screaming.”

Mr Moffat has observed a curious peculiarity of the flight.
When a bat is hunting insects at a considerable elevation,
the observer who is high enough, say, on a hillside, to secure a
horizontal view of it, may notice that its motions consist of a
regular succession of very gradual ascents and abrupt descents.
Mr Moffat suggests that the descent is merely a means of
keeping the bat in the plane where its favourite food abounds ;
at all events the regularity of the process is very striking, and
leads almost to the inference that they may be additional to
those already described movements of similar regularity in the
Noctule which Tomes has otherwise interpreted.

According to Mr Moffat, the cry of Leisler’s Bat, when
flying in company, is a shrill strong screech, which he judges,
although without opportunity of actual comparison, to be
shriller and stronger than that of the Noctule. There is
besides ‘‘a peculiar ‘tinkling’ song emitted by a single bat
when flying alone; when a second appears it is greeted with

1 Trish Naturalist, 1899, 169-174, and map.

)
(4) and (5) Intermed

ize.

il
inter

natur

inter. (7
Summer ; (6) W

, IkISH HARE—Summer and W

, SCOTTISH BLUE Hart

and 2
to 6

s1ze

(+ natural

late.

(3)

4
ro]

LEISLER’S BAT 97

the shrill screech, and from that time one hears the tinkling
no more.” It is to the ‘“tinkle” and not to the screech,
Mr Moffat thinks, that Dr Alcock referred when he stated
that the cry of this species is moderately high in pitch,
corresponding to about 17,000 vibrations per second of
Galton’s whistle, and that a fair imitation of it may be
made by “striking a sixpenny piece against a halfpenny.”

A Leisler’s Bat was kept alive by William Darragh in 1858,
for ten days. Later, Dr Alcock, Mr Moffat, Mrs Oldfield
Thomas (in the latter case Mr Finn’s specimen), and Mr
Whitaker’ have kept various individuals in captivity for some
little time. Under such circumstances it is an interesting and
cleanly pet, devoid of fear, and easily trained to come for its
food on hearing a particular sound. It is more active in a
confined space than the Noctule, but soon becomes lazy, and,
although it will, if necessary, fly to and settle on the hand
that feeds it, at other times it takes the earliest opportunity
of alighting. But when once it has alighted it is active
enough, scurrying round tables, and even falling off on to the
floor.

Dr Alcock describes a live one, which he received on 13th
February. It remained in a typical state of hibernation until
11th March, when it awoke to be fed, returning to sleep on cold
weather again setting in, and thus being in alternate torpidity
and vigour until its untimely death on 6th April. Another,
kept under observation by Mr Whitaker from 27th September
to 16th April, passed through somewhat similar alternations.

In appetite this bat does not fall behind its congener. Indi-
viduals shot while on the wing, which could not have been flying
for more than an hour, are described by Mr Moffat as “ mon-
strously full—-so round and firm and hard (almost like cricket-
balls), with the quantity of insect food they had gorged in that
short interval.” Others submitted to Prof. G. H. Carpenter
by Dr Alcock were found to have been feeding largely on
flies. Professor Carpenter was also able to identify, amongst
other insects, the yellow-haired dung-fly,” a midge,’ besides
legs and wings of caddis-flies, and, as Mr Moffat informs me, a

1 Naturalist, 1907, 387, 415, etc. 2 Scatophaga stercoraria.
3 Probably a mycetophilid, also an acalypterate muscid.

98 VESPERTILIONIDZ—NYCTALUS

beetle. The latter naturalist has observed the capture of moths
of fair size by this bat while on the wing.

In captivity it will eat raw meat and drink milk, and Mr
Whitaker estimates its daily requirements in mealworms, which
were in no case pouched, at about five dozen to each bat. One
kept in confinement by Mr Moffat, which he fed exclusively
on insects for a week, refused various house-flies, as well as the
blue-bottle,* common crane-fly,? and horse-fly,* but enjoyed the
common volucella. She partook with relish of the honey-bee
and of five species of wasp. A humble-bee* and a red-tailed
bee® were eaten, but subsequent specimens of the former were
declined. A few small beetles were promptly devoured.
Above all, she evinced an extraordinary partiality for the
common cockroach, of which she was known to eat nearly a
third of her weight in one night.

“Next to the cockroach, I think her favourite food (among
the insects offered) was the honey-bee, when dead; but she
showed the greatest horror when I offered her a ueng bee,
though I held it securely in my hand, and had previously
extracted its sting. This was no mere accident, for the
experiment was repeated on successive days. In two instances
she managed, apparently by a sudden ‘flick’ of her tongue,
to throw the bee to a distance. At first I was surprised that
this nocturnal mammal should so well comprehend the offensive
capabilities of a day-flying insect ; but when we remember the
common propensity of both... to take up their abode in
hollow trees, it is at once seen to be quite natural that they
should know something of each other’s powers. The identical
ash-tree in which the Hairy-armed Bats now under notice
reside has often been occupied by bee-swarms.”

Leisler’s Bat has on more than one occasion been taken in
houses, for entering which, however, it seems, like the Noctule,
to have but little proclivity, its flight being usually high in the
air and away from buildings.° There remain many aspects
of its life-history in regard to which we possess no informa-
tion. In particular, as to its breeding habits, we have almost

1 Calliphora erythrocephala. 2 Tipula oleracea.
3 Hematopota pluvialis. 4 Bombus terrestris. 5 B. lapidarius.
6 Moffat, /r7sh Naturalist, 1897, 135 ; Jameson, Jrish Naturalist, 1896, 94.

LEISLER’S BAT 99

to fall back upon Blasius’ statement, that in Germany it is
pregnant in June, and that the young may be seen flying
with their parents before autumn. Tomes! examined two
females each containing one foetus, and another now in the pos-
session of Mr Barrington, and taken at Crum Castle, County
Fermanagh, Ireland, in June, had attached to her a single
young one; so that it is probable that, as with the Noctule,
whatever may be the case in Germany, the number of young
in Britain is usually one.

Again, as regards the relative numbers of the sexes and
their association or separation when in hibernation, there is
nothing to add to what has already been stated in the article
on the Noctule. A party of six females, found by Mr Whitaker °
in a hollow tree in Yorkshire on 22nd August, may be supposed
to have just finished rearing their young.

The habits of Leisler’s Bat, or of its representatives, would
appear to differ somewhat in the different parts of its range.
Thus, in Germany, according to Blasius, it may be observed
on the wing in the shade of thick woods, where it always flies
at mid-day ; a remark which is fully borne out by the series of
the Azorean form*® obtained by Messrs W. R. Ogilvie-Grant
and the Hon. N. C. Rothschild, and which were shot while
flying in or near pine-woods, between the hours of noon and
dusk, and frequently in bright sunshine. No similar observa-
tion has been recorded for Britain, except that of Mr P. E.
Freke,* who near Milltown Bridge, County Dublin, at half-past
ten o'clock on the morning of 22nd June 1881 watched
a bat (which from his description could not well have been of
any species other than the present) hawking for flies on the
river Dodder, in company with numbers of swallows and swifts.

British naturalists who take an interest in mammals should
search carefully for this mysterious bat, which even a practised
observer finds difficult to distinguish from the Noctule when
on the wing. Mr Coward, one of the few English naturalists
since Tomes who has had opportunities of comparing the
two, informs me that its flight is slower and more erratic
than that of the dashing and rapid Noctule, but even he is not

1 Zoologist, 1854, 4365. * Naturalist, 1907, 385-6.
3 N. azoreum. * Zoologist, 1882, 16.

100 VESPERTILIONIDA—PIPISTRELLUS

invariably able to recognise them. In the hand, the smaller
size, darker colour, the absence of the peculiar smell, and
especially the markedly smaller feet, cannot fail to form an
infallible criterion as between even the very youngest Noctules
and the present species.

GENUS PIPISTRELLUS.

1829. PIPISTRELLUS, Jakob Kaup, System der Europiischen Thierwelt, i., 98 ; based
on Vespertilio pipistrellus of Schreber.

1838. ROMICIA, J. E. Gray, Mag. Zool. and Bot., 495; based on R. calcarata of Gray
=P, kuhiit (Natterer).

1839. VESPERUGO, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s Azchiv
fiir Naturgeschichte, \., 312 (part); based on Vesfertilio serotinus of Schreber and
eleven other species (see under NYCTALUS).

1840. ROMICIUS, Edward Blyth, Cuvier’s Animal Kingdom, 75 ; misprint for ROMICIA.

1856. Hypsuco, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung (Dresden),
Neue Folge, ii., 131, 167-169; based on Vesferugo maurus of Blasius, and V.
kraschenintkowit of Eversmann.

1856. NANNUGO, F. A. Kolenati, of. czz., 131, 169-171 ; based on Vesperugo nathusit
of Keyserling and Blasius, Vespertilio pipistrellus of Daubenton, and V. kuhliz of
Natterer.

1878. VESPERUGO, G. E. Dobson, Catalogue of the Chiroptera in the Collection of
the British Museum, 183 (part); included also NYCTALUS, VESPERTILIO, and
others.

1893. VESPERUGO, Harrison Allen, Bull. U.S. National Museum, 43, 121.

Classification and synonymy :—The explanation of the use
of the genus MVyctalus applies also to Pzpzstrellus, which
presents no further difficulties.

In characters, the bats of this genus differ from those of the
preceding in their smaller size, lighter build, and shorter, broader
wing (Plate VII., Fig. 2, p. 86), attached to the base of the outer
toe, not to the middle of the sole. The third and fifth metacarpals
are about of equal length, and the tail is usually about as long
as the body without the head. The tragus lacks the prominent
rounded head, being broadest below its centre, and narrowed,
not expanded, above (Fig. 2, No. 3, p. 7). The foot is small.

The skull (Fig. 8, No. 1), as exemplified by that of
P. popistrellus, is small, weak, and rounded; the facial
region is saddle-shaped, namely, concave posteriorly, convex
anteriorly; the premaxillary gap is moderately marked; the
zygomata are weak; cranial crests are absent or hardly

PEAT Bie Wit

¢
i
€
i.

Rex!

LEISLER’s BATs. (Approximately natural size.)

PIPTISTRELEUS IOI

perceptible, and the auditory bulla are more prominent than
in AZyotzs, less so than in Barbastella.

3
CE Sa ei a a

Fic. 8.—SIDE VIEW (diagrammatic and enlarged) OF SKULLS AND TEETH OF

1. Pipistrellus pipistrellus. 4. Plecotus auritus.
2. Myotis daubentont. 5. Barbastella barbastellus.
3. MM. mystacinus. 6. Rhinolophus hipposideros.

The teeth are similar in number and arrangement to those
of Myctalus, but the inner upper incisor is both more bicuspid

102 VESPERTILIONIDA—PIPISTRELLUS

and more prominent than the outer, while the anterior upper
premolar is moderately large and usually takes its place in the
tooth-row (Figs. 9 and 10, p. 108).

a
«

Fic. 9.—DIAGRAM OF ARRANGEMENT OF TEETH OF /pistrellus pipistrellus.
(1) Upper and (2) Lower Jaw.

The genus includes a large number of species of wide
distribution, of which at least three occur extensively in
southern North America, from austral zones south to Vera
Cruz. In the Old World it is represented from Tasmania
—P. tasmaniensis (Gould)—to Ireland. The single British
representative is the well-known /P. pzfzstrellus, which in the
east and south is replaced by the closely-allied P. abramus
(Temminck). The latter has a larger forearm (34 to 35
mm.), more naked muzzle, and the anterior upper pre-
molar so small as to be scarcely visible from without. It is
believed to range from Australia and Japan through the Malay
Peninsula, Ceylon, Assam, Burma, and India, migrating in
summer to central Europe, and has been reported from Cadillac,
Gironde, France (Trouessart, Le Maturaliste, 1879, 16, 125),
and Sweden. But its reputed occurrences in Europe should
no doubt be credited to P. wathuszz (Keyserling and Blasius),
with light-coloured posterior wing-border and long lower
canine, described as of central Europe. Of other species,
P. kuhlit (Natterer), a southern form, also taken at Gironde,
and with a sub-species reaching Cape Colony (P. kuhlit fuscatus
of Thomas), has the posterior border of the wings white and
the outer upper incisor minute; P. savzz (Bonaparte), a form of
wide Palearctic range, with a representative—P. albolimbatus
(Kiister)—in Sardinia, and shown by Forsyth Major (AZé¢z della

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 103

Soc. Toscana dt Sct. Nat., 3, fasc. 1, 87, Pisa, 1877) to be synony-
mous with smaurus of Blasius and donapart: of Savi, has the
forearm reaching 34 mm., the tragus broadest at its middle, and
having two small lobes projecting from its hinder border, and
a broad callosity on the foot; P. sxbflavus (Cuvier) of North
America, has the thumb larger, and the forearm reaching
34 mm.

THE COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE.
PIPISTRELLUS PIPISTRELLUS (Schreber).

1667. FLITTER-MOUSE, or REAR-MOUSE, Christopher Merrett, Pzzax, 172.

1693. VESPERTILIO: The Bat or Flittermouse, John Ray, Synopsis Quadrupedum, 243. —

1760. LA PIPISTRELLE, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s
flistoire Naturelle, viii., 129-130, 135, 137, pl. xix., fig. 1; also, MWém. de l Acad.
Roy. des Scz., 381, pl. i. (14), fig. 3, 1759, published 1765 ; described from France.

1766. SHORT-EARED BAT, Thomas Pennant, B77tish Zoology, ed. 1, 55.

1775. VESPERTILIO PIPISTRELLUS, J. C. D. von Schreber, Die Sdadugthiere, i., pl. liv.,
167, evidently naming Daubenton’s La Pipistrelle; Jenyns, Trans. Linnean Soc.
(London), xvi., 163, read 3rd Feb. 1829, published 1833; Bell (ed. 1) ; Clermont ;
Flower and Lydekker.

1789. VESPERTILIO MURINUS, Gilbert White, Se/éorne, letter xi. to Thomas Pennant,
gth Sept. 1767, original edition, 32; Berkenhout ; Bingley; Donovan; Fleming ;
not V. MURINUS of Linnzeus, 1758 = (probably) V. DISCOLOR of Natterer; not
V. MURINUS of most continental writers = MYOTIS MYOSOTIS (Bechstein).

1811. VESPERTILIO PIPISTRELLA, P. S. Pallas, Zoographia Rosso-Asiatica, i., 123,
No. 48.

1825. VESPERTILIO PYGM&UuS, W. E. Leach, Zoological Journal, i., 560, pl. xxii.;
figured by Bell in both editions; described from a young specimen taken at
Spitchweek, Dartmoor, England, and now in the British Museum.

1829. PIPISTRELLUS PIPISTRELLUS, Jakob Kaup, System der Europaischen Thierwelt,
i, 98; Miller, Ann. and Mag. Nat. Hist., Oct. 1897, 384; Thomas, Zoologist, 1898,
100; Johnston; Méhely ; Cabrera; Millais, 79.

1833. VESPERTILIO BRACHYOTUS, L. A. F. Baillon, A7ém. de la Soc. Roy. d’Emula-
tion a’ Abbeville, 50 ; an unrecognisable species, probably referable to pipzstrellus ;
described from a specimen (collo super. nigro) from Abbeville, France.

1838. SCOTOPHILUS MURINUS, J. E. Gray, Mag. Zool. and Bot., 497 ; MacGillivray.

1839. VESPERUGO PIPISTRELLUS, A. Graf von Keyserling and J. H. Blasius, Wieg-
mann’s Archiv fiir Naturgeschichte, i., 321; Blasius ; Fatio; Dobson; Blanford ;
Lydekker.

1840. VESPERTILIO MINUTISSIMUS, Heinrich Schinz, Euvropdtsche Fauna, i., 9;
described from Zurich, Switzerland (type in Zurich Museum).

1840. ““V. MELANOPTERUS et STENOTUS, Brehm,” auct. e¢ op. cit.; described from
Renthendorf, Thuringia, Germany, and Switzerland.

1840. “V. PUSILLUS, Brehm,” aucé. et of. cit.

104 VESPERTILIONIDA—PIPISTRELLUS

1840. VESPERTILIO LACTEUS, C. J. Temminck, Monographies de Mammalogie, ii.,
245 (for date see Wiegmann’s A7chiv, 1841, 23) ; described from two nearly white,
immature bats of doubtful origin in the Leiden Museum (see Dobson, Ca/alogue,
225).

1841. VESPERTILIO VISPISTRELLUS, C. L., Prince Bonaparte, Fauna Italica, i., pl. xi.,
fig. 1; described from Italy.

1842. KERIVOULA GRISEUS, J. E. Gray, Aum. and Mag. Nat. Hist., December, 258 ;
described from a spirit specimen of unknown origin in the British Museum (Dobson,
Catalogue, 225).

1844. VESPERTILIO NIGRANS, J. Crespon, Faune Méridionale, i., 24, is thus placed
by E. L. Trouessart, Bull. de la Soc. d’ Etude des Sci. Nat. de Nimes, 7,1, 55, 1879;
and, on the authority of Paul Gervais, identified with P. Jzpistre//us though slightly
smaller; it was described from Nimes, France. But Gervais, Histoire Naturelle
des Mammiferes, \., 216, 1854, cites V. nigrans next to pipistrell/us, but as a distinct
species, under the name of 7. zigricans Gené: this is evidently not /. nigricans of
Maximilian, 1826, which antedates it.

1856. NANNUGO PIPISTRELLUS, F. A. Kolenati, Allgemeine deutsche Naturhist.
Zeitung (Dresden), Neue Folge, ii., 131, 170.

1858. NANNUGO MINUTISSIMUS, F. A. Kolenati, S7tzungsberichte der Kaiserlichen
Akad, der Wissenschaften (Vienna), xxviii. (3), 246, and xxix. (10), 334, fig. 7.

1862-1863. NANNUGO PIPISTRELLUS, a, var. TYPUS, p. 490; 8, var. FLAVESCENS,
Pp. 491; 7, var. NIGRICANS, p. 491; 6, var. LIMBATUS, p. 492; Carl Koch,
Jahrbiicher des Vereins fiir Naturkunde im Herzogthum (Nassau), xviii., 487-500 ;
described from Nassau, Germany.

1870. VESPERUGO MINUTISSIMUS, L. J. Fitzinger, S¢tzungsberichte der Katserlichen
Akad. der Wissenschaften (Vienna), Ixii. (1), 252.

1874. SCOTOPHILUS PIPISTRELLUS, Thomas Bell, “story of British Quadrupeds,
ed. 2, 34; E. T. Newton, Quart. Journ. Geol. Soc. (London), 1894, 193.

La Chauve-Souris, or la Pipistrelle of the French; de Zwergfleder-
maus of the Germans.

Synonymy :—This presents no difficulty. Those who object to
the Scomber scomber principle will probably find the combination
Pipistrellus pygmaeus (Leach) correct.

Local Names (non-Celtic):—Athern-bird, of Somerset (Forbes) ;
back-bearaway! (black-bear-away, Forbes), of Yorkshire (Atkinson,
Zoologist, 1878, 330; and Forty Years in a Moorland Parish, 1892, 137),
from back,? backie, bak, bakke, bauckie, baukie, or bawkie, an old form
of bat, having variants in Denmark, Iceland (compare /edhrblaka, i.e.,
leather-flapper), Norway (compare Old Norse d/aka, 1.e., to flap), and
Sweden (compare zattblacka, i.e., a bat), and still used in Scotland, often
in combination as backie-bird, etc. (Alston); barnmouse, bastat, bathy-

1 A word usually applied to ships, and probably meaning sailing or floating away
(Wright).
2 Compare—
“The laverock and the lark, The baukie and the bat,
The heather-bleet, the miresnipe, How mony birds be that ?”
— CHAMBERS, Popular Rhymes of Scotland (ed. 1870), 198.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 103

mouse (Forbes); bat-mouse, of Stafford (Adams, MS.); batty mouse,
billy-bat (Plecotus, Wright) ; bit-bat (Forbes), blin’bat, z.e., blind-bat, of
Kirkcudbright (Service, Zoologist, 1878, 427); blind-bat, of Dublin
(Moffat, MS.); brere-mus (Forbes), (?) = reremouse; chipper, ze,,
chirruper, from its cry of “chip, chip” (Forbes); flickermouse (Ben

Jonson), as in New Jna, iii., 1—

“Once a bat, and ever a bat! a rere mouse,
And a bird o’ twilight; ...
Come, I will see the flickermouse” ;

flinder (Forbes), compare Dutch v/znder, i.e.,a butterfly ; flinder-, flinter-,
z.é.. butterfly-mouse (compare in Caxton’s Reynard the Fox (1481), 112,
ed. Arber—

“Thenne cam . . . the flyndermows and the wezel” ;

fliner (Forbes); fletter-, flit-, flitter- (or vlitter-) mouse, flitty, flitty-
mouse, fluttermouse (compare German fledermaus), of many counties,
e.g., Yorkshire (Atkinson, of. cz.) and Essex (Laver), as in Ben Jonson’s

VAUCHEHLIST, V., 2—
“My fine flitter-mouse,
My bird o’ the night ;”

glaik of Lothian (Forbes), ze, a transient glance or gleam; haddabat
(Miller and Skertchley, Fen/and, 1878, 358); hatbat (Forbes); leathern
mouse, leathern wings (Forbes); leather-winged bat of Wexford (Moffat,
MS.) ; mouse bat, of Buckingham and Berkshire (Cocks, Zoologzst, 1878,
334); Oagar-triunse, of Shetland (Forbes), (?) = -hiunse, -hinnse or
-hunch, ze, a frightful creature (Wright); pipistrelle, mostly a book-
name, from French fcfrstrelle, from Italian pzpistrello, vipistrello, vespis-
trello, variants or diminutives of vespertz/lo or vespertilio, from Latin
vespertili9, i.e.,a bat; raamis, ramished, raamouse, raamse, raird, ramsh,
rare, rare-, rattle-, rye-, and ry-mouse, rawmil, rawmouse, rawmp, ray-
mouse (Gloucestershire), rearie, rearmouse, reelrall, reelymouse, reeraw,
reerd, reerie, being mostly words signifying riot or confusion, or con-
nected with reermouse, and reremouse, from middle English reremous,
and Anglo-Saxon hréremus, from /réran, i.e., to move, shake, stir, and
mus, 1.€..a Mouse, as in—
‘Some war with rere-mice for their leathern wings,

To make my small elves coats ”
—SHAKESPEARE, Midsummer Night's Dream, Act ii., Sc. iii. ;

rannermouse (Bingley), rannymouse, of south Hampshire (Corbin,
Zoologist, 1878, 429), see under SHREW.

(Celtic):—Most usually zaltag or zaltog, with variants dialtog, fialtag or
taltag, perhaps from za// = leather, sometimes combined with /eathazr,
ie., leather zaltag ; craicneach (?) = craicneog, i.e.. skinny, as in Man
(see Kermode); callah or cal-luch, and feascarluch, mioltog leathair,

106 VESPERTILIONIDA—PIPISTRELLUS

ie., leather fly; sczathan leathazr, i.e., leather wings ; yst/um,as in Wales;
see under NOCTULE (Alston; Forbes; O’Connell; Wilde, Proc. Roy.
Trish Acad,, vii., 189, 9th and 25th May 1859, published 1862.)

Distribution :—Pipistrelle-like bats occur throughout boreal and
transitional Europe and Asia, from sea-level to 6000 feet in the Alps
(Blasius), from about 60 N. lat. in Skandinavia, Russia, and Asia,
to the Mediterranean, with the Balearics (Barcelo, but not found
by Thomas and Pocock), Sicily, the Caucasus, Ural, and Altai
ranges, Kashmir valley and Gilgit (Blanford), and from Ireland
to an unknown point in Siberia or China. In austral and _ transi-
tional zones the direct representative in North America is P. subflavus
of the eastern United States.

From north to south of the British Isles this species is probably
numerous in every locality where bats can exist, and it was until recent
years a general assumption that it is everywhere the commonest species.
No doubt this is frequently true, especially in the south and east of Eng-
land, where no bat flies more frequently or persistently, or is more easily
observed and caught, so that it is almost certain that it is actually the
commonest to the eye. But in many localities close observers report that
its numbers are equalled or exceeded by those of A/Zyotzs mystacinus, from
which in flight it is with difficulty distinguishable, of Plecotus auritus
or Nyctalus noctula. In Lincoln it outnumbers all the other species
together (Caton Haigh), but in parts of Buckingham Cocks finds P. aurztus
more numerous. In the west and north its numbers decrease; thus,
although common on the Gloucester side of the Avon, it becomes rare on
the Somerset side, and Charbonnier and Lloyd Morgan believe that it may
be there replaced by JZ. mystacinus, as Jenyns found to be the case toa
creat extent at Bath (see Harting, Zoo/ogist, 1888, 164). In Shropshire
also its numbers are less than those of P. aurztus (Forrest), and in
Merioneth and North Wales generally it is less plentiful than either of
the other three species mentioned above. From the north of England
similar reports are forthcoming, it being in Cheshire locally less common
than JZ. mystacinus (Coward and Oldham), in Yorkshire probably about
as numerous as that species or P. aurvztus (Roebuck quoted by Harting,
loc. ctt., 165), and in Lakeland not so abundant as in the southern
counties (Macpherson).

In Scotland it is stated to be in most localities much the commonest
bat, and to range to the extreme north (Alston), becoming, however,
less numerous in Sutherland and rare in Caithness. It must be
remembered, however, that this is only a comparative statement, since
other bats, except P. auritus and MW. daubentoni are, as a rule, absent
from Scotland. For Ireland, Lyster Jameson gave a list of twenty-two
counties where it is known to occur, and it is certain that it is present
also in the remaining ten (see Patterson, /r7sh Naturalist, 1900, 233).

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE. to7

It has probably found its way to every island of any extent, and
has been identified from Orkney (Spence, Aun. Scott. Nat. Hist., 1909,
47), Islay, Mull (Alston; Gilmour, J/ag. czt., 1897, 191; Russell), the
Outer Hebrides (Harvie-Brown and Buckley), Man (where Kermode
considers it common, and whence Oldham has received a specimen),
Anglesey (Coward and Oldham), Wight (More; Wadham), and the
Scillys (Clark). It is probably the species which has been observed in
Shetland (hibernating in a peat stack, Tulloch, J/ag. cz¢., 1904, 125; also
Millais), Jura (by myself), the Blaskets (Kane, /7zsh Naturalist, 1897, 88),
and other Irish islands, and at such outlying lighthouses as the Fastnet,
Rockabill, and Blackrock, Co. Mayo (Barrington, MWzgration of Birds,
284). It is common in all the Channel Islands (Bunting). So diverse
are its haunts that, although reaching an altitude of at least 750 feet at
Carmichael, Scotland (Watt), it finds a sustenance equally well in the
heart of the largest towns, even of London (Rendall, Zoologzst, 1888, 24).

Distribution in time :—Pleistocene remains from Ightham Fissure,
Kent, have been provisionally referred to this species by E. T. Newton
(Quarterly Journ. Geol. Soc., ist May 1894, 193).

Period of gestation:—At least forty-four days (A. Whitaker,
Naturalist, 1907, 74, and in lit.), but see under NOCTULE.

Number of young and breeding season :—The (in Britain) single
young one is born sometime between late June and early August.
Continental naturalists write of two young ones at a birth as not unusual.

Description :—The form and general characteristics of P. pzpistrellus
are those of its genus. The ear when laid forward reaches to a point
about half-way between eye and nostril. The exterior margin is more
deeply notched than in Nyctalus (Fig. 2, No. 3, p. 7; for head, see
Plate XI., Fig. 4, p. 140).

The fur is long and thick, and almost hides the round, very small
eyes; above the anterior angle of each is a small wart, from which
grow a few black hairs; on the forehead a transverse tuft of long
upright hair alters the apparent contour of the face, making the fore-
head appear comparatively prominent, whereas it is in reality depressed
in contrast to the well-developed glandular prominences of the muzzle.
The furring of the wing resembles that of Vycta/ws, but there is no carpal
band. The interfemoral membrane is furred above nearly as far as a
line drawn between the ankles when the tail and interfemoral membrane
are extended ; beneath, the fur passes on to the membrane only at the
root of the tail and along the inner side of the thighs, about half the
remaining portion being covered with fine short hairs extending princi-
pally along the tail.

The colour of the upper side varies between “mars brown” and
“Prouts brown,” with tendencies towards “wood brown” and even
“clove brown”; the lower side is lighter. The hidden portions of the

108 VESPERTILIONIDA:7—PIPISTRELLUS

hairs are everywhere dusky, the dusky basal portion reaching a length
of 7 mm. out of a total of 10 mm. on the back. The wing, leg, foot,
ear, nose, and naked parts are dusky. The young are generally of a
more uniform, usually a darker, colour than the adults, owing to the
absence of the grizzled hair-tips, but there are exceptions, as in one
labelled 5th August, which is about the lightest individual which I have
seen (see Jenyns; Fatio; Couch, Zoologist, 1853, 3942).

Nothing is known of the seasonal changes of this species, but I
believe that they resemble those of the genus Vyctalus, since there seems
to be a tendency to deeper tints in March and to comparative pallor

in summer until August. There are cer-
a tainly no marked changes, and such as
—— : ‘

LAG occur are obscured by individual variation.
Two young ones (Plate II., Fig. 2,
— ee p. 16), born in the possession of Whitaker
Tee } had at birth a few straggling hairs on the
AQ UW? muzzle ; the body was dull flesh colour,
the wing and ear darker, but lighter than
\ in an adult. One was abnormal in that
it neither advanced in size nor apparent
development for a month. The other
began to grow darker after about five
days, and by the twenty-first day the
whole body had become dusky. The
eyes opened on the eighth day, and about
the same time hair began to appear on
the back and shoulders, later on the head
and chin, lastly on the breast: the belly,
however, was still naked on the thirty-

i a & ) third day, when the animal died.
pee oe agen Se: The skull and teeth are typical of
strellus (enlarged and diagrammatic). the genus (Figs. 8, No. I, p. 101; 9,

p. 102; and Io).

Individual variation is very frequent, and tends in three directions,
ze, towards melanism, erythrism, and albinism, intermediates of every
description being plentiful. One in the Grosvenor Museum, Chester,
killed in that city on 6th April 1896, is “coal black, with a rusty
tinge on the belly and flanks” (Oldham). Borrer mentions a rusty red
one (Zoologist, 1874, 4125), and there is an example of this type in
the Royal Scottish Museum at Edinburgh. Of all-white specimens, with
or without pink eyes, I know of about seven altogether (see Pelly, /ze/d,
14th Sept. 1889, 408, and J. M., Journ. czz., 2nd Oct. 1875, 368; Millais,
85; and one in Tring Museum). In the most remarkable the wings
and ears were white like tissue paper, the legs, arms, digits, nose and

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 1Io9

lips pinkish white, but the fur was only slightly paler than usual
(Charbonnier, Zoologist, 1901, 472).

Very little is known about the geographical variation of this species.
Eastern European examples are larger than those of the west, and
particularly of Britain. They have been distinguished as the sub-species
macropterus (Verhandlungen der kaiserlich-koniglichen szool-bot. Gesell-
schaft (Vienna), xii, 1862, 250), described by Zeittelles, from Kaschau,
Hungary. Cabrera’s sub-species medzterraneus, of the east of Spain, is
of a clearer red colour, and is said to present some structural differences.
Blanford states that Indian examples from dry, sandy regions are paler,
and sometimes have the underside almost white, and this is the case
also in the deserts of Turkestan, where the representative sub-species is
lacteus of Temminck (Thomas, Anzu. and Mag. Nat. Hist., March 1909,
258), and of a Quetta specimen in the British Museum.

DIMENSIONS IN MILLIMETRES? :—

MALEs.

3 4 ne

ys a op 3 » rm 2

2 EY & s 5 0 al eye labs 2

fe) <— ~~ ote) s = a aS) = = &

= & a By NS ie Sy vies 313 Sees
3 FA = r= = -= B BE | ss a Et % Sb
Salinas 2 | a bes .| &) |e he |S. he lesa
a 2 a fe} po £ fa 2 3 3 ge

3 0) oo = oO 3 [o) $ oO »

2 = eS} q a 4 = S

q K 60 = = r= ey =

o s q o

& 1
B

No. of Items . : 9 ae vs 9 4 12 12 3
Maximum. . | 44°5 | ( 34 11°75 | 6°75 | 80 | ( 212
Average. 5 . | 42 10°00 | 4°00 |< 31°85} 10°5 5°65 | 29°5 +! 3°5-5| 50 25 25 208
Minimum z . | 39 | | 29°2 | 10 4 PH fa) | S3 ‘a ae ate i 206

FEMALES.
No. of Items . al {toa lal ate ate 11 5 5 1l Se ae fe oe 6
Maximum . A Be | ss as pe 12 8 met | ati

Average . 4 . | 43°35 | + 10:00 | 4:00 30°5 | 11°36} 7°14 | 30°28 }/| 8°5-5| 50 25 25 213
Minimum 5 . | 345 | | 29°21|10°5 | 6°75 | 29 | i 203

1 For these I am partly indebted to Whitaker. The table suggests that, as in the Noctule,
females may be slightly larger than males.

Four young males in the Dublin Museum, taken at Bohoe Church,
Co. Fermanagh, Ireland, by Lyster Jameson, have the forearm measur-
ing 19-5, 24:5 (two), and 26-5 respectively. Couch (Zoologist, 1853,
3942) found the extent of wing varying between 175 and 210. The
following approximate dimensions of an embryo almost ready for
birth were sent me by Coward :—head and body 25, ear 3:5, tail 10,

110 VESPERTILIONIDA:—PIPISTRELLUS

hind-foot 3, forearm 12, thumb 2:5, digit iii. 13, digit iv. 10, digit v. 10,
expanse 70. :

Proportionate lengths :—Foot, without claws, about -55 to -62 of
lower leg; fifth metacarpal about equal to third; lower leg about -30
to -35 of forearm, and about -25 of head and body.

Skull :—Greatest length, 12-5; basal length in middle line, 9-5;
palatal length in middle line, 5 ; from posterior border of 7? to anterior
border of canine, 4:25; same in lower jaw, 5; greatest breadth at
zygoma, 8; posterior breadth, 6-5 ; breadth between orbits, 4; breadth
at constriction, 3-75.

Weight in grammes :—Couch (/oc. czt.) weighed 28, young and old,
in July, and found them varying from 2-6 to 5-4 (40 to 83 grains):
Moffat (/riskh Naturalist, 1900, 235) gives the weight as from 4-5 to
5-8 (70 to 90 grains): while Eagle Clarke sends me the following
details—two males, alive, Dec., 5-53 (86 grains) and 4-63 (72 grains) ;
one, 9th May, 4:5; one, 23rd April, 4.4; one, 9th May, 4-5: females,
one, 8th Sept., 6-3; one, roth April, 5-15; one, 9th May, 4-5; one
(young), 9th Sept., 4.

Distinguishing characteristics :—In the present state of our know-
ledge any small bat having an expanse less than 220 mm. may be
assumed to be an example of this species until proved to the contrary.
Its size at once distinguishes it from the British members of the genera
Nyctalus and Vespertilio, its ears from Plecotus, Barbastella, and
Rhinolophus, leaving only A7yotis, of which two species, dechstetnz and
natterert, are distinctly larger; dauwbentoni is less so, but has distinct
coloration and markedly larger foot ; szystacinus is of somewhat similar
size, but its thirty-eight teeth, longer oval ears with their outer margins
terminating anteriorly about under the base of the tragus instead of
near the angle of the mouth, attenuated tragus, and the total absence
of a post-calcarial lobe, are sufficiently distinctive. On the chance of
P. nathusit occurring in England it would be well to remember that this
species has the hinder border of the wing light coloured.

The Pipistrelle is the smallest British bat—Zin fact
the smallest British mammal—its weight about equal-
ling that of the Pigmy Shrew, which for long held
the reputation of being the tiniest. By the older English
naturalists it was confused with the Common Bat’ of conti-
nental writers, a very different animal, and one having no
claim to be regarded as British. This error appears to
have originated in the days of Pennant, perhaps with Gilbert
White himself, but was frequently repeated until Fleming’s

! Vespertilio murinus, not of Linnzeus = Myotis myosotis (Bechstein).

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 111!

time, to be at length dispelled by Leonard Jenyns, who
reviewed the whole question in 1829, and clearly demonstrated
that the Common Bat of Britain is the Pipistrelle of European
zoologists.

The widespread abundance of this bat, the lengthened
duration of its period of activity, and its love for the neigh-
bourhood of dwellings, make it to most people more familiar
than any other, if we except only the Long-eared, which
is in many localities as common. The only species likely
to be mistaken for it is the Whiskered, which it closely
resembles in size, in mode of flight, and in many of its
habits. So close is the resemblance that when flying it is
no easy matter to distinguish them. However, it is believed
generally that, while the Whiskered Bat often feeds amongst
trees, snatching its prey while at rest on the leaves or branches,
the Pipistrelle usually feeds near them, and takes its food
preferably on the wing. Even this distinction, however,
may not bear investigation, and the difficulties surrounding
the observation of small bats on the wing are so numerous,
that the differentiation of these two species in life must
be regarded as one of the most difficult tasks which confront
a field naturalist in Britain.

Sometimes the Pipistrelle flies alone in some sequestered
nook ; at other times, especially on windy evenings, a number
will gather together to tread an aerial dance, the passing by of
two at close range often leading to a spirited encounter and
chase. The small area over which it hunts, and the frequently
restricted extent of its beat are very noticeable, and Mr Charles
Oldham tells me that this is also true even when the animal
is flying over open country, where there is no natural boundary
such as a wall, house or fence. It is probably one of the species
which in winter time appears inside churches, distracting the
attention of the congregation from the evening service, and
causing scandalised rectors to set forth their grievances in the
public press in hopes that some zoologist may point the way
to relief. It was, perhaps, this trespassing in churches that
led to the inclusion of bats in the churchwardens’ accounts of
the parishes of St Paul's, Bedford, and Dean in Bedfordshire,

1 Field, 23rd June 1906, 1044.

112 VESPERTILIONIDA—PIPISTRELLUS

wherein, as Mr J. Steele Elliot has shown,’ they made their
appearance between the years 1797 and 1838 in the lists of
malefactors for whose head a reward was paid. In the latter
parish no fewer than eight hundred and fifty-two bats of one
kind or another were killed and paid for at the rate of sixpence
a dozen; and, indeed, their deaths cannot be altogether
regarded as judicial murders, since they often become a well-
nigh intolerable nuisance. At Claxby Rectory, Lincolnshire,
where, as Mr J. G. Millais relates, a wall had to be pulled
down before a large colony could be ejected, their squeaks
and scratchings and their odour were at one time unbearable.
Again, Mr Oxley Grabham states that the station-master’s
house at North Grimston, Yorkshire, was so overrun by the
parasites emerging from a colony of two or three hundred bats
as to lead to its destruction.

In its choice of nocturnal haunts the Pipistrelle is very
catholic. Wherever the Whiskered Bat is seen, this species
may be seen also. Its favourite resorts are sheltered corners
of an orchard, stack- or farm-yard, a lane, or indeed any other
quiet spot, most often near a homestead, gardens, or wood-
land clearing. Here it may be observed night after night,
just before dusk, and at about the same time as, but usually a
few minutes before, the Noctule. ‘Time after time it pursues
the same round, occasionally varying it by excursions to sport
in mid-air with some comrade on an adjacent beat, or to
snatch a brief visit to some neighbouring hunting-ground ;
at intervals it disappears altogether for a few minutes, but
generally returns to its accustomed circle, and thus continues
until darkness hides it from view. Backwards and forwards,
round and round, it flits with noiseless wing-beats, sometimes
at an altitude of fifteen or twenty feet above the ground, some-
times much lower. Frequently it darts with wonderful activity,
now here, now there, or pursues a minute and to human eyes
invisible prey into the corners of buildings, descending if need
be to within a few inches of the ground. It seems, like the
swallow, to seek its food in the neighbourhood of animals,
and will visit the cattle in their sheds, or flit about a man’s
head until a whip or butterfly-net cuts its flight short. So

1 Zoologist, 1906, 165 and 167.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 113

abundant is it that it is often easy to have several bats in view
at the same moment, and so familiar that, as far as the dusk
will permit, the shape of the wings—despite the extreme
rapidity of their motion—may be traced with the eye. The
interfemoral membrane appears to be extended nearly horizon-
tally, the ears are distinctly visible, and certain indefinable lumps
on the wings in front and behind suggest the thumbs and feet,
the latter, as far as I can make out, carried projecting backwards.

It must not be thought, however, that this bat is by
any means entirely confined to a lowly or monotonous flight.
Sometimes, as R. F. Tomes noticed, it accompanies Daubenton’s
Bat in its water-patrols, occasionally dipping its nose in
the liquid to slake its thirst. At other times, as Mr T. A.
Coward informs me, it flutters slow and moth-like around
the tops of trees, its wings appearing on such occasions
much broader than usual. Its appearance to Mr William
Evans, while out wild-fowl shooting amongst the sand-hills
of Aberlady Bay, Haddingtonshire, indicates that it may travel
to considerable distances in search of food, and may even find
its way out to sea, since Mr R. M. Barrington’ reports that
one was found dead on the Arklow South Lightship, some
miles off the south-eastern coast of Ireland, on 21st September
1898. Possibly this particular bat had been blown out to sea,
since we have no evidence that the species is migratory.

As the autumn evenings darken, the Pipistrelle issues forth
each night at an earlier hour,’ but its hibernation, at least in
the southern districts both of England and Ireland, is a very
uncertain and evidently a perfunctory affair. Although, in the
north, in all probability, many individuals enter upon their
winter sleep before the end of October, there is in the south
no night of the year on which, should the weather be pro-
pitious, this species or the Whiskered Bat may not be found
abroad. These winter flights have often been noticed.
Jonathan Couch® kept a day-to-day record of them in Corn-
wall in the years 1852-53, and Jenyns concluded that they are
more frequent before than after Christmas.

1 The Migration of Birds, 284 (R. H. Porter, London, 1900).

* For some notes of the time of appearance, relative to sunset, see Fauna of
Cheshire, 12.

° Zoologist, 1853, 3936-3943.

114 VESPERTILIONIDAE—PIPISTRELLUS

More recently, Mr C. B. Moffat’ has taken the trouble to
compare the movements of bats, hedgehogs, and frogs from
26th October rt901 to 21st February 1902, as observed
at Ballyhyland, County Wexford, Ireland. His facts are
carefully tabulated, and mark a distinct advance in our know-
ledge of the winter routine of three very different types of
animals. He finds that bats (and he thinks that practically
in every case they were Pipistrelles) ‘‘were observed on nine-
teen evenings in November, nine evenings in December, ten
evenings in January, and five evenings in February.” He
therefore concludes that at ‘‘all times during the winter, pro-
vided the temperature of the hour of dusk is above 43° Fahren-
heit, some Bats of the common species are pretty sure to be
found flying, if looked for in suitable localities. Below 43°
their emergence is not to be calculated on, but it sometimes
takes place at lower temperatures, down to 39.” It will be
noticed that a return of mild weather brings out the Bats
immediately, no matter how frosty the previous nights and
days may have been. The reason why so few Bats were seen
during February was that the whole of that month—until the
last week, when they reappeared—was continuously cold.”

Discussing the possibility that not all these observations
referred to the present species, Mr Moffat admits that he is
unable to distinguish it from the Whiskered Bat, which also
occurs at Ballyhyland. But he contends that, although a few
of the latter may have been noted in his table as Pipistrelles,
yet his general results would not be thereby affected, and I
think the same remark applies to the records of other observers.
Most of the bats which he saw at low temperatures cannot,
he believes, have been other than Pipistrelles; and in point
of fact he saw one captured on 28th December 1901, when the
temperature stood at 434° Fahrenheit.

Mr Moffat’s conclusions may, I think, be regarded as
sound so far as concerns the winter flights of the Common
Bat in the south-east of Ireland, and his limitations may
be taken as approximately correct; in fact I can myself
corroborate them for the same county, where at Kilmanock

1 Trish Naturalist, 1904, 81-87.
2 The corresponding figures as given by Jenyns are 4o° and 38° F.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 115

on windy winter evenings the Pipistrelle may be found hunting
within the shelter provided by a large partially open cattle-shed.
Bats have, however, been on occasions observed to fly at
temperatures below 38° Fahrenheit, as when Captain L. H.
Irby," on 2nd January 1887, observed one on the wing in
Surrey in bright sunshine at noon, the ground being covered
with snow and the temperature below the freezing-point—
an observation repeated by Mr G. H. Caton Haigh? at
Grainsby, Lincolnshire, on the 1oth of the same month,
in similar conditions as regards weather, but in bright moon-
light.

The partiality of small bats of some species for mid-day
flights is well known, and their frequent appearances
seem to show that they can have no inherently strong
objection to daylight. But the identification of such flights
with the Pipistrelle can only rest on conjecture, and at the
present stage of our knowledge it is quite beyond possibility
to say which species makes the most frequent appearances
by daylight. The evidence brought forward by good
observers is, however, on the whole, inclined to connect
them chiefly with the Whiskered Bat. Whatever be the
truth, bats which venture out at such unorthodox hours
must fain submit to the persecutions to which the original
in general are subjected, and have sometimes, as Couch?
has observed, been driven back to their dens by the angry
attacks of scandalised birds.

The fondness of this animal for different species of gnats
has been observed from the time when Pliny * wrote (although
probably of a different species), “Z¢ 2 czbatu culices gratissimt,”
and it is probable that these little flies constitute no small part
of its usual food. But, judging from its habits in captivity,
it doubtless consumes a variety of insects. According to
Mr Caton Haigh,’ it frequently captures comparatively large
insects, the well-known ‘“daddy-long-legs” or crane-fly being
one of its favourites, while Mr Millais states that it devours

1 Zoologist, 1887, 69. * Tbid., 1887, 143. 3 OD. cit., 1853, 3942.

* C. Plinit secundi Naturalis Historiae, ed. of Joannes Harduinus, lib. x., cap. Ixi.
(Ixxxi.), 454 (Paris), 1685 (“and for-food gnats [are] very pleasant [to it]”).

5 Zoologist, 1887, 293.

116 VESPERTILIONIDA—PIPISTRELLUS

large quantities of the smaller moths; and Mr Moffat has
noticed that when it secures anything of formidable propor-
tions, it invariably disappears for a few minutes, unlike the
larger Leisler’s Bat, which does not permit the struggles of
any insect to interrupt its flight.

Mr Moffat has taken considerable pains to show that,
unlike that of the Noctule and Leisler’s Bat, the flight of the
Pipistrelle is continued all night: and this in spite of the fact
that the majority of British naturalists, from Dobson’ to Mr
Millais, with the exception of William MacGillivray and a few
others, seem to have taken it for granted that this species,
like others which are without nasal appendages, must be more
properly crepuscular than nocturnal in its habits. Mr Moffat
writes :°—

‘‘Of course, in observing Bats, one must be very careful
that one knows what sort of Bat one is observing. The
difficulty of being quite certain on that point vitiates a good
many observations that might otherwise be useful. However,
I began my enquiries into the Pipistrelle’s habits by passing
a night in the open air in bright moonlight, in a spot where
large numbers of Bats generally fly. The result of this pre-
liminary mode of enquiry (on the night of August 21st-22nd,
1899) was that I found that there were lots of Bats visible on
the wing at all hours throughout the night, as well as in the
clear light of early morning. That was not conclusive, be-
cause, in the first place, these Bats might not all have been
Pipistrelles, and, even if they were, some might have gone
home early and others come out late, so that there was no
proof that any individual Bat, Pipistrelle or otherwise, had
been flying about the whole time. The next thing to do was,
therefore, to find out where some of these Bats went in the
morning. By watching on several mornings, in the summers
of 1899 and 1900, I ultimately got the retreats of half a dozen,
each living a perfectly solitary life in a little den of its own—
some in holes in walls, and some in the trunks of trees. That
made it possible to play the detective on these six individuals,
and I soon found that the hours of all six were very similar,

1 Catalogue of Chiroptera, xvii., footnote, 1878.
2 Trish Naturalist, 1905, 101-103.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 117

and, on the whole, very regular. Each of them left its retreat
every evening during the half-hour after sunset, and returned
to it every morning during the hour before sunrise. The time
of emergence would, indeed, vary, even for the same individual
Bat, from so early as ten minutes to so late as thirty minutes
after sunset, and the time of retreat similarly varied from so
early as forty to so late as eighteen minutes before sunrise ;
but in no instance did a Bat, whose sleeping-place was known,
on occasions when I watched for its emergence, fail to come
out during the evening twilight, or when I looked for its
return in the morning, disappoint my expectation of seeing it
go in. After ascertaining this much concerning their habits,
I caught three of these animals as they were coming out, and
they proved to be Pipistrelles. I have no doubt that the
remaining three were the same. That does not tell us all that
we want to know, but it tells us something. Not only is it
known that a good many Pipistrelles are on the wing during
the hour before sunrise, but it is also established that these
are the same individual Pipistrelles which left their retreats
early after sunset the previous evening, and not, as might be
imagined, mere belated individuals that had overslept them-
selves before coming out, and were making up for it by
breakfasting late.

‘But no amount of mere watching, moonlight or otherwise,
would tell whether these animals remained away from their
sleeping-places all night, or whether they follow the Hairy-
armed Bats’ rule of taking a midnight nap. So, on the night
of August 16th, 1900, I did what I had done four nights
previously in the case of the Hairy-armed Bat, and fixed a net
at midnight over a hole which a Pipistrelle had quitted the
previous evening twenty-eight minutes after sunset. The
result was the opposite to what had happened in the case of
the Hairy-armed Bat. At 3.45 in the morning no Bat had
come out of the hole, and as it now wanted only an hour to
sunrise, it was time to remove the net so as to let the Bat in.
Of course I kept watch to see that it did go in, and at twenty
minutes past four—some twenty-eight minutes before sunrise—
I had the gratification of seeing it make its usual return. Now
there could be no doubt, in the case of that animal, that it

118 VESPERTILIONIDA—PIPISTRELLUS

had been out all night. All that remained was to make sure
—a very important matter—that I was right concerning its
species; so the next evening I set the net again over the
same hole, caught the Bat as it came out, and found that it
was a male Pipistrelle.

“The above experiment was made on a fine bright night,
so I thought it safer to try it again on another Bat under less
comfortable conditions, choosing this time a dark and foggy
night, when nobody could suppose that Bats would be specially
tempted to fly late. Such a night occurred on August 30th,
1900, when I netted the residence of a second Pipistrelle. The
result, however, was just the same as in the former case.
No Bat came out after midnight, but, at the usual time before
sunrise, the occupant of the hole went in. Hence, it follows
that even during raw and foggy nights, when insects might be
presumed scarce, the Pipistrelle does not retire into its den, but
continues abroad till its usual hour for seeking sanctuary in
the twilight of the early morning. I even find that the same
thing happens in winter when the nights are warm enough for
the Pipistrelle to fly. I have several times seen it at midnight
in the long nights of December and January, and though |
have not stayed out at that season to see it going home at
seven or eight o'clock in the morning, I have trustworthy
information from one whose vocation brings him out at these
hours (Mr James Kelly, Ballyhyland), that it stays on the
wing till nearly daylight—in other words, flies through a sixteen
hours’ night.”

It should be noted that Mr Moffat’s results do not alto-
gether agree with those of J. R. Kinahan*' as quoted, more
particularly under Daubenton’s Bat. Kinahan’s observations
show that in a mixed colony of these two species some at least
of the bats returned to their den at ten o'clock in the evening in
June. In any case, Kinahan’s observations, interesting as they
are, are not to be compared for completeness with Mr Moffat’s,
and, as only a minority of the colony were Pipistrelles, are cer-
tainly not conclusive. Mr Moffat suggests that, in view of the
undoubted difficulty of distinguishing such small bats on a stormy
night at ten p.M., Kinahan may have been deceived in what

l Nat. Hist. Review (Dublin), 1854, 23-24.

COMMON BAT PIPISTRELLE*OR FPLITTER-MOUSE 119

he saw, and, perhaps, thought that bats passing close to the
wall were returning to their den. The probable truth of the
matter is that neither the Pipistrelle nor any other bat is
absolutely tied down to habit, but we may be quite sure that its
flight is continued off and on throughout the night. That inter-
ruptions may be frequent is averred by Mr Arthur Whitaker, who
assures me that he has more than once observed a short day-
break flight in this species, a fact which I can myself corro-
borate, having on one occasion observed a Pipistrelle active for
about five minutes just before sunrise. It is quite likely that in
inclement weather, or where there is a large colony, the bats
may come and go throughout the night, perhaps to visit and
feed their young or to devour more safely some large insect.

The Pipistrelle is by no means fastidious in its choice of a
place of concealment. No kind of crevice, crack, or aperture,
whether of tree or building, within or without, comes amiss to
it, and in such places it lives either singly or, more particularly
in summer, in large parties, the membership of which is not
always restricted to its own species; it has no objection to the
noisy companionship of man in the neighbourhood of its
sleeping-place. The inside of an old and disused wooden
pump has on occasion furnished a somewhat remarkable
resting-place, a bat having been seen to emerge from the
spout. It has been asserted that trees are much less frequented
by this species than by some others, but, apart from Mr
Moffat’s experience, instances are known of its having been
discovered hiding behind pieces of loose bark and amongst
ivy. The bats sometimes show a preference for some par-
ticular retreat; thus Mr Oldham informs me that on 16th
June 1888, he took an adult male from behind a window-
shutter at a farmhouse in Wisbech Fen, Cambridgeshire. On
the 18th he found an adult female on the wall in precisely the
same spot, and on the 19th and 21st two other Pipistrelles,
also adult females.

If there be any situation to which this bat apparently
objects it is to the interior of caves, resting within which, so
far as I am aware, it has been but rarely detected, although
Mr Coward has found it in secluded rock crevices just outside
a cave. It was not amongst the species found by Dr E. A.

120 VESPERTILIONIDA—PIPISTRELLUS

Wilson and Mr A. H. Cocks in Mr Heatley Noble’s cave at
Henley-on-Thames, nor does Monsieur Gadeau de Kerville?
include it amongst the seven species revealed by a three
hours’ search in the ‘‘ Grotte de la Briqueterie, Seine-Inférieure,”
France. In fact, he states expressly that he has never met
with it in such situations.

There is some evidence to show that, like the Noctule and
Leisler’s Bat, the Pipistrelle regularly varies its hiding-place
with the season, the sleeping-places of summer being frequently
distinct from the hibernacula of winter. One such instance
has been placed on record by Mr F. Boyes,’ who knew of a
retreat at Bishop Burton, near Beverley, from which no fewer
than one hundred and twenty-six Pipistrelles had been counted
as they emerged in summer, yet it was deserted in winter. In
its winter retreats, at all events, the animal is often solitary, but
immense troops gather together in summer. In these, unlike
the Noctule and Leisler’s Bat, both sexes participate, and a
Hampshire colony was estimated by Mr Whitaker to include
from three to four hundred individuals.’

The voice of the Pipistrelle is feeble when compared with
that of the larger bats, and can only be appreciated by sharp
ears, but Ovid was clearly guilty of poetic imagination when
he wrote of it that “ Wenzmam pro corpore vocem enattunt.” *
In truth, the little creature shrieks with might and main when
irritated, and it is hardly its fault, its gamut not being intended
for human ears, that its efforts are to most of us barely per-
ceptible. Mr Oldham remarks that individual bats differ widely
in the matter and manner of their utterings.

As with other bats, the breeding season is_ probably
autumnal, but hiemal activity is accompanied by desul-
tory pairing of the extent of which we are ignorant. Messrs
R. Rollinat and E. L. Trouessart,® whose interesting researches
on the breeding habits of bats have been already dis-
cussed at length, are quite firm in their assertion that, no
matter how frequently such winter breeding may take place,
as they admit it does, fecundation is retarded until spring

1 Bull. Soc. Amis Sct. Nat. (Rouen), 4th April 1gor, reprint, 2.
2 Field, 29th August 1903, 405. 3 Naturalist, 1907, 77.
4 Metamorphoses, \ib. iv., 10. 5 Supra, p. 32.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 121

precisely as in the case of other bats. On the other hand, Mr
Whitaker suggests that pairing may take place during the
latter half of May, at which time he has noticed bats
chasing each other, a fact not necessarily evidence of pairing.’
Unfortunately, no precise observations exist for Britain. It
is, however, certain that the date of birth varies a good deal,
at least from late June’ to early August, but it is never, so
far as is known, so early as to be inconsistent with Messrs
Rollinat and Trouessart’s results. Thus Mr H. Lyster
Jameson,® when visiting Bohoe Church, Ireland, on rith
July, found a number of young Pipistrelles, from a few
days old to half-grown individuals, crawling about the
floor of the church, having fallen through a hole in the
ceiling. As regards July births, four captive females, in
the possession of Mr Whitaker, produced their respective
young on the 2nd, roth, 18th, and roth; while Mr Coward
found an embryo nearly ready for birth in a female killed on
the 4th of the same month. The earliest of these would, no
doubt, have been born in June, and the possibility of early
births is further strengthened by the capture of a young one
on the wing on oth August at Exeter,* and of another at
Kilmanock, County Wexford, Ireland, on the 13th. Allowing
seven weeks each for gestation and rearing, the birth of these
two bats must have taken place some time between 2Ist and
25th June. Their southern habitats sufficiently account for
the early date, and it is interesting to note that the Wexford
bat was fully grown, although certainly immature.

In two of Mr Whitaker’s bats the known period of gestation
was not less than forty-one and forty-four days, and ina third it
probably reached forty-nine. The mother was captured between
27th and 31st May, had for companions two males of
her own species, and when she died on 14th July was found
to contain a small embryo, ‘probably not more than _half-

1 See his “ Notes on the Breeding Habits of Bats,” Maturalist, 1905, 325-330.

2 A statement by J. J. Briggs (Zoo/ogist, 1848, 2278) that the bats in Melbourne
Church, Derbyshire, “bring out their young about June 17th,” is unfortunately some-
what indefinite as regards the exact species.

3 Jrish Naturalist, 1896, 95.

* Now in the Exeter Museum, and kindly submitted to me for examination by
Edwin Hollis.

122 VESPERTILIONIDAZ7—PIPISTRELLUS

developed.” If the facts as reported by Mr Whitaker be
correct, and assuming that the true period of gestation could
not have exceeded some forty-nine days, we are left with
the alternative explanations that either the date of ovula-
tion was delayed by captivity, or that either fertilisation
or pairing had taken place after the date of capture. The
case is full of interest, but does not afford much ground
for speculation as to its meaning. It may, however, be
expected that the breeding habits of a bat having such
slender inclinations to hibernate, must present many features of
interest, and will probably be found to vary with the climatic
conditions of different parts of its range, as in fact we know to
be the case in regard to the number of young.

On the latter point, continental authorities from Pliny—
“ Geminos volitat amplexa tnfantes” '—downwards unite in con-
sidering two at a birth quite usual. This is certainly not the case
in Britain, where the observations of Tomes, George Daniell, Mr
Whitaker, and others show that, as with the Noctule, one is the
almost invariable rule. Of five female Pipistrelles received by
Daniell? in July 1833, each contained a single foetus ; and Tomes
also was convinced by the examination not merely of British
specimens, but of a great number of foreign ones, that the pro-
duction at a birth of more than one young one is exceptional.
The only (and that an extremely doubtful) British reference
to the birth of twins which I have been able to find is
the merely incidental statement of Couch, that a friend
‘observed of one to which the young ones were attached, that
they were separated from the teats with difficulty, and that
when separated they were not able to lay hold of them; and
the old one then seemed quite indifferent to her young, running
over them without care.” The act of parturition was observed
in one case only by Mr Whitaker. The mother clung head
downwards to the side of the cage and received her offspring
in her right wing, which she held partially extended for the
purpose.

The young ones born in Mr Whitaker’s cages were at
first stowed away under their mother’s wings, exactly like

1 Loc. cit. supra, p. 115, “[The mother] flies embracing twin young.”
2 Proc. Zool. Soc. (London), 11th Nov. 1834, 129-132.

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 123

young Noctules, and in that position were at first hardly
visible from without. After a few days the relation between
mothers and offspring became much less close, and they began
to hang themselves up alone at a distance of a few inches from
their parents. When they were a week or two old they usually
spent the day, often close together, at the side of the cage
opposite to that affected by their mothers, and would sleep
quietly by themselves for many hours. When wanting their
mothers or when touched, they would lift themselves well up
on the wrists, raise the head high and turn it anxiously
about from side to side, uttering a chirrup resembling the
soft smacking of human lips; this was very faint when the
creatures were young, but steadily grew in power as the days
went by. This sound was uttered with widely open mouth,
and after calling for a while the bat would set off in search
of its mother, whom it clearly distinguished from that of its
companion, and when found seized with its teeth by any
part available. Generally each managed to work its way
to a comfortable position under a wing, where, once arrived,
the mother would usually bend down her head to “tuck”
it in, at which times it could be heard uttering a soft, musical
twitter.

If the mother happened to be feeding at the time, she
would often take no notice of her young, but would drag it
carelessly about, clinging possibly to her interfemoral mem-
brane, or to the fur of her back. Whenever she happened
to pause, the youngster would try to improve its grip, and
the tenacity with which it clung to her was astonishing, so
that it very seldom dropped off, when once it had taken
hold. After feeding, the mothers always attended to the
young ones immediately. If the baby was under its mother’s
wing when she came out to feed, it did not hamper her
movements very much until it came to pouching a meal-
worm; the young one appeared to render this most difficult,
and seemed to be very much in the way. This was so
much the case that, when the young ones were three weeks
old, Mr Whitaker was compelled either to feed the mothers
by hand, or to remove the young ones from them whilst

they fed.

124 VESPERTILIONIDAZ—PIPISTRELLUS

Mr Whitaker more than. once allowed the mothers
to fly with their young ones, but they were always very
careful, before commencing to do so, to expand the
wings fully, and lift them up and down a few times, to
ascertain that ‘‘baby” was not clinging to the wing itself.
They seldom flew for more than five or six times the
length of the room with the youngster attached, but when
they did so, the youngster was plainly to be seen holding
to a nipple with its teeth, and to the fur with its feet,
the back being quite arched, so that the young one hung
well down from its parent, and was conspicuous enough
when the latter was flying in a room in the daylight.

The young Pipistrelles unfortunately did not survive to
eat solid food or fly, although the stronger of the two was
frequently observed to open and stretch its wings during the
last few days before its death on the thirty-second day.

The Pipistrelle has frequently been kept alive for long
periods in confinement, and possesses many lively and interesting
habits. It is far more active and ready to take wing than
the Noctule or Leisler’s Bat, and, although irascible to members
of its own species, recognises its patrons, circling easily round a
room, and soon learning to come to the hand for flies or other
dainties.

Bingley long ago showed that captive Pipistrelles are so
fearless as to take mealworms from the hand on the very day
of their capture. In this they resemble Noctules, and are as
a rule much less shy than the Whiskered or the Long-eared
Bats. When hungry they are not at all fastidious, and will
eat readily small pieces of raw meat, usually refusing such
as are not quite fresh and juicy. Their capacity for meal-
worms, according to Mr Whitaker, reaches thirty a day.
They like also many kinds of insects, sometimes eating, some-
times rejecting the wings; but Mr Oldham’s specimens could
only be induced to touch white butterflies or the yellow-
underwing moth after much persuasion, and Mr Whitaker's
refused the generally unpalatable magpie moth.

Mr Whitaker has experimented on the senses of this
bat in captivity, and writes me that sight and_ hearing
seem the most important used both by it and by bats

PEAT Ee It:

whe si Se

Fic. 1.—TyPIcCAL MOLAR TEETH OF AN INSECTIVOROUS BAT (after Miller).

A, Crown view of upper molar. 4. Crown view of lower molar.
C. Side view of upper molar,

ecd, entoconid mc, metacone mis, metastyle prc, protocone
hc, hypocone mcd, metaconid pe, paracone pred, protoconid
hcd, hypoconid ms, mesostyle ped, paraconid ps, parastyle

Fic. 2.—YOUNG PIPISTRELLE. (Natural size.)

From a specimen in the British Museum, to show the small wings, but
large feet and thumbs.

PEATE vildite

PIPISTRELLE BATS, (About #? natural size.)

COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 125

in general when feeding. A Pipistrelle which had become
accustomed to picking up mealworms from the floor of its
cage, was puzzled by finding a piece of white wool of
similar length, but seemed indifferent to bits of black or red
wool. It ate two blackened mealworms, but with hesitation,
after smelling them carefully, as it did also the white worm-
like pieces of wool. On the other hand, it almost sprang
upon ordinary mealworms and devoured them, without
attempting to smell them. But the peripheral nerves
generally must be extremely sensitive, since a temporarily
blinded Pipistrelle, liberated with a piece of paper firmly
gummed across the front of the tragus, not only flew round
a room briskly and without hesitation, but avoided contact
with obstacles, and smartly dodged strokes of a net.
Captive Pipistrelles frequently make use of the inter-
femoral membrane in the pouch-like manner (Plate VIII., p. 104)
which, although undoubtedly observed by older writers, and
mentioned by Dobson,’ appears to have been for the first time
thoroughly appreciated and intelligently interpreted by Mr
Oldham. The history of this habit is worth recounting at
some length. It came under the notice of White,” who wrote
to Pennant that he was much entertained at the manner
in which a tame bat, when fed, “brought its wings round
before the mouth, hovering and hiding its head in the
manner of birds of prey when they feed.” Bingley is
more detailed in his description, since he relates how a
bat, “raising itself somewhat higher than usual on its fore-
legs, bent its head with great dexterity under its belly,
and forced the insect into its mouth, by thrusting it,
from side to side, against that part of the membrane
which extended betwixt the two hind legs.” Again, in
1834, Daniell wrote of a bat falling over its prey, ‘with
all its membranes expanded, and cowering over the pros-
trate fly, with its head thrust under, in order to secure ”
it. Later, Joseph Clark* thus graphically described the
attack of a captive bat upon a fly: “If it missed its aim,
its next tactic was to use its long arm, to get it under its

1 Catalogue of Chiroptera, xxviii. 2 Letter xi., dated 9th Sept. 1767.
3 Zoologist, 1847, 1766-1767.

126 VESPERTILIONIDZ—PIPISTRELLUS

body ; it would turn and curl the membraneous part of its
tail inwards, forming a complete sack, into which, if it suc-
ceeded in getting its prey, it would thrust its head for the
purpose of capture.”

Clark’s remarks naturally lead up to the more detailed
observations of Mr Oldham, which will be found transcribed
at length in the account of the Whiskered Bat. It should
be remarked, however, that when making his observations
Mr Oldham was quite unaware of those which preceded his ;
in fact, he is in many respects in a position to correct them,
inasmuch as no previous observer had clearly stated that
what a bat actually does is to thrust its head, with
the insect actually tn rts mouth, into the interfemoral pouch,
its object being to prevent the escape of its prey, should it
by any chance break loose during its struggles. Mr Oldham
finds, however, that a bat only makes use of the pouch
when it has to overcome the struggles of a strong insect ;
at other times it is content to devour its prey with its head
entirely in the open.

The inference is that bats make a similar use of the inter-
femoral pouch while on the wing, but very few naturalists
have sufficiently keen eyesight to follow their evolutions while
thus engaged in the open. Amongst the few are the late
Frank Norton,’ Professor C. Lloyd Morgan,’ and Messrs E.
D. Cuming and Lionel E. Adams, of whom the first named
stated that he had often kept tame Pipistrelles in his house,
where they had full liberty of the rooms, and appeared to
be well content with the diet of flies thus afforded them.
When catching a fly, the bat, so far as he could judge,
struck its prey a blow with its wing, which disabled it, and
then seized it before it reached the ground, using its tail as a
basket until it had obtained a firm hold. Mr Cuming writes
somewhat similarly in his pleasant book, Zhe Arcadian
Calendar,? and in reply to my inquiry for further details,
he adds the information that he has “often watched the

' Midland Naturalist, 1883, 149-153, arranged and contributed by H. A. Mac-
pherson.

* Animal Life and Intelligence (London: Edward Arnold), 1890-1891, 65.

° George Newnes, Ltd., London, 1903, 35 (illustrated by J. A. Shepherd).

VESPER TIETO® 127

Pipistrelle swoop through a swarm of gnats, and have, on
favourable opportunities, seen the tail depressed to ‘bag’
the membrane, the animal’s flight at same moment—or in
the same movement—often turning slightly upward. It
appeared to me that this manceuvre could have no other
purpose than the capture of insects, and when I found that
another observer had recorded the same thing, I accepted it
as confirmation of my own opinion, and adopted it as fact.”
Lastly, Mr Adams informs me that he observed the manceuvre
in sunlight on 17th March 1906.

When feeding at large, Pipistrelles, like the Noctule,
execute falcon-like swoops on their invisible prey, with
the wings evidently spread to their widest extent, so that
way is kept on even at the fullest depth of the plunge. On
other occasions an intermediate line of action may evidently
be taken, since I have seen a remarkable headlong descent,
during which one wing was for an instant curiously curved
so as almost to enclose the body in front. No doubt, like
other animals, bats are not such rigidly red-tape disciplinarians
as to be forced to follow a fixed sequence of movements every
time they catch a fly. It is probable, therefore, that their
plans are varied, the smaller flies being snatched up without
trouble, while for the capture of the larger insects the mem-
branes may be to a lesser or greater extent employed, or the
struggling victim may even be carried off to be mastered at
leisure in some secure retreat.

GENUS VESPERTILIO.

1758. VESPERTILIO, Carolus Linnzus, Systema Nature, x., 31-32 ; based on Vespertilio
murinus of Linnzus (not V. murinus of Schreber, 1775, see above, page 51).

1820, EPTEsSICUS, C. S. Rafinesque, Aznals of Nature, 2; based on Eftesicus
melanops of Rafinesque = Vesfertilio fuscus of Beauvois.

1829. CNEPHZUS, Jakob Kaup, System der Europaischen Thierwelt, i., 103; based
on Vespertilio serotinus of Schreber.

1839. VESPERUGO, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s Archiv
Sir Naturgeschichte, i., 312 ; based on Vespertilio serotinus of Schreber, and eleven
other species.

1839. VESPERUS (sub-genus), auct. et of. cit., 313; based on the thirty-two toothed
species of the genus Vesferugo, e.g., serotinus of Schreber, discolor of N atterer,
nilssont of Keyserling and Blasius, savzz of Bonaparte, /eucippe of Bonaparte, and

128 VESPERTILIONIDAE—VESPERTILIO

aristippe of Bonaparte ; preoccupied by VESPERUS of Latreille, 1829, a genus of
Coleoptera.

1841. NOCTULA (sub-genus of P7fzstrellus), C. L., Prince Bonaparte, Fauna Italica,
i., xxl. (under Vesfertilio alcythoe) ; based on Vesfertilio serotinus of Schreber.

1856. CATEORUS (sub-genus), F. A. Kolenati, A//gemeine deutsche Naturhist. Zeitung
(Dresden), Neue Folge, ii., 131, 162-163; based on sevofinus, i.e., La Serotine of
Daubenton.

1856. METEORUS (sub-genus), F. A. Kolenati, of. cz¢, ii, 131, 163-167; included
nilssoni of Blasius, dzscolor of Kuhl, leucippe, aristipfpe, and savit, all of Bonaparte ;
preoccupied by METEORUS of Halliday, 1835, a genus of Hymenoptera.

1858. AMBLYOTUS, F. A. Kolenati, Szfzungsberichte der Kaiserlichen Akad. der
Wissenschaften (Vienna), xxix. (9), 252 ; based on A. atratus of Kolenati= Vesperugo
nilssoni of Keyserling and Blasius.

1863. ““ARISTIPPE, F. A. Kolenati, Bettraige zur Kenntniss der Phthiriomyiarien,
Petersburg, 1863” (thus in Koch, Das Wesentliche der Chiropteren, etc., 471, 473,
1863, under A/eteorus) ; included V. discolor of Natterer=V. murinus of Linnzeus
and Vesperugo nilssoni of Keyserling and Blasius.

1864. SCOTOPHILUS, Harrison Allen, Monograph of the Bats of North America, 28
(part).

1866. PACHYOMUS, J. E. Gray, Ann. and Mag. Nat. Hist., 90, February ; based on
Scotophilus pachyomus of Tomes.

1870. NYCTIPTENUS, L. J. Fitzinger, Szteungsberichte cit. supra, \xii. (1), 424; based
on Vespertilio smithiz of Wagner.

1872. MARSIPOLZMUS, W. Peters, Monatsberichte der Kiniglich Akad. der Wissen-
schaften (Berlin), 260, a sub-genus of Vesferugo based on Vesperus (M.) albigularis
of Peters= V. murinus of Linnzus.

1878. VESPERUGO, G. E. Dobson, Catalogue of the Chiroptera in the Collection of the
British Museum, 183 (part); included also NYCTALUS, PIPISTRELLUS, and others.

1892. ADELONYCTERIS, Harrison Allen, Proc. Acad. Nat. Science, 466, Philadelphia
(December 8, 1891) ; proposed as a substitute for Vesperus.

Classification and synonymy:—The use of the generic term
Vespertrlio has been fully explained on pages 50-51.

M. Méhely uses Rafinesque’s name /Aéeszcus for the serotinus-
fuscus group, restricting the name Vesfertzfo to the remainder of
the genus as here understood. Mr Thomas has pointed out
the objections to this proposal (Azn. and Mag. Nat. Fiist., July
I9OI, 31-32). A more recent arrangement, that of Mr Miller,
makes the division between V. murinus as sole representative of

Vespertitio, and the remaining species as Fféeszcus. This will
probably meet with general acceptance, since the short broad
tragus and large nares of typical Vesfertzlio are quite character-
istic, and not in accord with those of the other members of the
genus. The present work was written too early for its adoption.

As defined in the present work, this genus includes a

number of species of collectively wide distribution. The

VESPERTILIO 129

best known are, besides l”. serotznus, V. nzlssont of Keyserling
and Blasius (= V. dorealts auctorum) of northern Europe and
Siberia generally to the Arctic circle; V. murinus of Linnzeus
(= V. discolor auctorum) of the temperate portions of the Pale-
arctic region, mainly in moun-
tainous districts (for distinctions,
see page 140, under V. murznus) ;
and Cabrera’s V. ochromixtus
with forearm only 32 mm., in
central Spain and the Balearics.
There are representatives in
North and South America (see
under V. serotenus).

From those of the preced-
ing genera these bats differ in
their slightly smaller, narrower
ear (Fig. 2, No. 4, p. 7), distinctly
longer than broad. The tragus
is straight, short, moderately
pointed, broadest near the middle,
and with a slight forward direc-
fon. The wing» (Plate IX.,
Fig. 2, p. 126) is broader than in
Nyctalus, but spring's from the Fic. 11.—DIAGRAM OF ARRANGEMENT

: ay f OF TEETH OF Vesfertilio serotinus.
base of the toes, as in P2pr-
strellus. There are fewer teeth
(Fig. 11), the small anterior upper premolar being absent,
leaving only four teeth, not five, behind the upper canine, and
the formula is thus—

(1) Upper and (2) Lower Jaw.

eee: I-I 1-1 3-
7c c , pm ars
2-2) 3-3

3-3’ I-1
The skull (Fig. 4, No. 2, p. 56), as exemplified by sevotznus,
is heavily built, flattened, larger and altogether stronger than that
of Pipzstrellus, but, as compared with that of Vyc¢alus, longer,
less massive, and with more rounded brain-case ; the facial region
is angular and concave, the premaxillary gap much less marked,
the zygomata expanded, the cranial crests posteriorly prominent,
the auditory bullz smaller. In dorsal profile the skull presents
nearly a straight line, rising gradually from nose to occiput.
N

130 VESPERTILIONIDAZ—VESPERTILIO

Only one species, . serotznus, is a regular member of the
British Fauna, but /. survinus has twice been taken at or
near seaports.

THE SEROTINE.
VESPERTILIO SEROTINUS, Schreber,

1760. LA SEROTINE, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s
FHiistoire Naturelle, viii., 129, pl. xviil., fig. 2; also, Mém. de Acad. Roy. des Scz.,
377 and 380, pl. ii. (15), fig. 2, 1759, published 1765 ; described from France.

1774. VESPERTILIO SEROTINUS, J. C. D. von Schreber, Die Sdugthiere, 1, pl. liii.,
167, evidently naming Daubenton’s Za Serotine; Jenyns; Bell (ed. 1) ; Clermont ;
Dobson ; Flower and Lydekker ; Miller ; Thomas, Zoologist, 1898, 100 ; Johnston ;
Cabrera; Millais.

1806. VESPERTILIO NOCTULA, Isidore Geoffroy, Ann. du Mus. a’ Hist. Nat., viii., 193,
pls. 47 and 48 ; transference of name in error.

1811. VESPERTILIO MURINUS, P. S. Pallas, Zoographia Rosso-asiatica, i., 121, No.
46; preoccupied by V. murinus of Linnzus, 1758.

1811. VESPERTILIO SEROTINA, @uct. et op. cit., 123.

1827. VESPERTILIO WIEDII, C. L. Brehm, Ovazs (Jena), iii, No. 3, 24; described
from Jena, Germany.

1827. VESPERTILIO OKENI, auct. et op. cit., iii., No. 4, 25; described from Jena,
Germany.

1829. VESPERTILIO RUFESCENS, auct. cit., Oken’s /szs (Jena), 643; described from
Jena, Germany.

1838. SCOTOPHILUS SEROTINUS, J. E. Gray, Mag. Zool. and Bot., 497 ; MacGillivray;
Bell (ed. 2).

1839. VESPERUGO (VESPERUS) SEROTINUS, A. Graf von Keyserling and J. H.
Blasius, Wiegmann’s Archiv fiir Naturgeschichte, \., 313; Dobson; Blanford.

1844. VESPERTILIO INCISIVUS, J. Crespon, Faune Méridionale, 1., 26 ; described from
Nimes, France.

1844. VESPERTILIO PALUSTRIS, aucé. et of. cit., 22 ; described from Nimes, France.

1856. CATEORUS SEROTINUS, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung
(Dresden), Neue Folge, ii., 131, 162-163.

1857. VESPERUGO SEROTINUS, J. H. Blasius, Saugethiere Deutschlands, 76 ; Lydekker.

1862-1863. CATEORUS SEROTINUS, var. TYPUS, var RUFESCENS, and var. PALLIDUS ;
Carl Koch, Jahrbiicher des Vereins fiir Naturkunde im Herzogthum (Nassau), xVili.,
463-467, etc. ; described from middle Europe, south Europe, and the “die steppen
Sud-Russlands” respectively : the latter apparently = V. urcomanus of Eversmann.

1870. VESPERUS SEROTINUS, L. J. Fitzinger, Sttzungsberichte der Kaiserlichen Akad.
der Wissenschaften (Vienna), Ixii, (1), 88.

1900, EPTESICUS SEROTINUS, Méhely Lajos, Monographia Chiropterorum Flungariae,
209, 340, pl. xiv.; Miller ; Trouessart (1910).

La Sérotine of the French; ade spdtftiegende Fledermaus of the
Germans.

Serotine from the French sérotzn, from the Latin serotinus, z.2., late,
or backward.

“Rattle-mouse” of Wight (Guyon, Zoologzst, 1856, 5216).

— 3".

i a i a i i i i a a

THE SEROTINE 131

Distribution :—Bats of this type have a remarkably wide distribu-
tion, being found over the greater part of the known world. In
the Palearctic Region they range from north Germany, south
Russia, and corresponding latitudes in Siberia southwards to the
Barbary States, Asia Minor, Transcaucasia, Arabia, Persia, Kashmir,
China, and probably Japan, ascending in the Harz Mountains to 2000,
and in the southern Alps to 4000 feet. Representative or allied forms
of the old world are mentioned under variation.

In North America the closely allied V. fuscus of Beauvois ranges
throughout the austral, transitional, and the lower edges of the
boreal zones, and has sub-species in (1) Costa Rica, Guatemala, and
southern Mexico, (2) in Guatemala and Nicaragua, (3) the Bahamas,
(4) Cuba, and probably elsewhere.

In the British Isles the Serotine is entirely confined to the south
of England. The first British record, that of Gray for London in
1826 (Zoological Journal, ii., 109), was almost certainly made in error, but
it had the effect of introducing the species into the works of Jenyns,
Bell (ed. 1), and MacGillivray, to all of whom no other locality was
known. In 1846 it attracted the attention of H. N. Turner at Folke-
stone (Zoologist, 1847, 1635), and in or before 1851 of Bury and Borrer in
the Isle of Wight ( Journ. czt., 1874, 4126), to be followed by Martin’s,
More’s, and Guyon’s experience of it in the same island previously
to 1854 (/Journ. cit, 1854, 4179; see also 1856, 5216). Since then
numerous observers have detected it in the south-eastern counties,
between the valley of the Thames and the Channel, as in Sussex (Borrer,
Zoologist, 1874, 4126, and 1893, 223; Lilford, 1887, 65; Ellis, Fzedd,
14th Oct. 1893, 597, and Zoologist, 1893, 458; Jeffrey, 1894, 261;
Butterfield, 1897, 141); in Hampshire (Lilford, Journ. czt., 1887, 65 ;
Kelsall, 1891, 395); in Kent (Harting, Journ. cét., 1890, 380, and 1891,
203; Dowker, 1891, 305 and 424). It appears to be numerous in many
parts of Kent. It is rarer, although frequenting many localities, in
Sussex, and rarer still in Hampshire. In Wight, fifty years ago, More
found it a common species ; some later observers describe it as rare, but
Wadham finds it very common and widely distributed. Jersey has one
record (Bunting).

In the west it is almost unknown, or has escaped notice, but is
said to have been obtained near Newquay in 1902, and near Lostwithiel
in 1906, and to be not uncommon about Porth (Clark), while one from
Tintagel Castle, all in Cornwall, is in the British Museum (see Dobson).
D’Urban, in a letter to Hollis, connects with this species certain large
bats, of which he saw flights at Exmouth, Devon, on 24th Sept. 1892.

North of the Thames the Serotine must be regarded as very rare.
Harting denies its occurrence in Middlesex (Zoologist, 1891, 203),
and cites, on the authority of Bond, Dartford Heath, Kent, as its

132 VESPERTILIONIDA:—VESPERTILIO

nearest known locality to the metropolis. An Essex specimen killed
before 1863 at Coggeshall was detected by Miller Christy in 1883, and
another was taken by him in 1894 at Broomfield (Journ. czz., 1883, 173;
Proc. Essex Field Club, iv., iv., 1892, and Zoologist, 1894, 423-424). A
third Essex specimen, taken at Pitsea, near Tilbury, in August 1906,
was exhibited to the members of the Essex Field Club, by Rev. A. B.
Hutton, on 24th November. It was examined and identified by Laver.
The first-mentioned locality remains the most northerly authenticated
for Britain, although Coburn believes that he once had a Birmingham
specimen through his hands (Zoologzst, 1892, 403). A reputed example
in the Newcastle Museum has been examined by Southwell, and proves
to be a Noctule (/ourn. cet., 1887, 234), and Lilford’s belief that it
occurs in Northampton (/ouwrn. czt., 1887, 65) may have been founded
on a misapprehension.

The restricted distribution in Britain of a type of such wide range
in the world is very remarkable, and must be regarded as one of the
puzzles of British mammalogy, difficult or impossible to account for
unless on the supposition that the species is either newly arrived or
decreasing its range. The abundance of bats may depend on so
many circumstances of which we have little knowledge, that it may be
well to note More’s statement of this species that in the Isle of Wight
it was very common until the felling of the timber.

Distribution in time:—This species is not known in a fossil or
semi-fossil condition.

The period of gestation, breeding season, and number of young have
not been recorded for England; but in Germany, according to Blasius,
there is but one young one, born, according to Kuhl (Of. cet. supra,
p. 83, 1, 191), in the latter half of May: in other respects, this bat is
stated to be normal (see page 31, etc.).

Description :—The Serotine is of about the size of the Noctule,
but with the ear oval (Fig. 2, No 4, p. 7), somewhat triangular, and when
flattened out, with the broadly rounded tip reaching to a point about
midway between nostril and eye; the outer margin is concave for
the upper half, then convex, slightly emarginate opposite the base of
the tragus, and ending in a convex lobe behind the angle of the
mouth; the inner margin is slightly convex with a rounded basal
lobe. The tragus is elongate, broadest just above the base of the
inner margin, thence diminishing slowly in breadth to the bluntly
pointed tip; its inner margin is straight or slightly concave, the outer
convex with a small projecting rounded basal lobe. The tumid face is
sparsely haired in front, but with a fringe of stronger hairs on the upper
lip; the glands are not so prominent as in Vyctalus (Plate IV., Fig. 3, p. 60).

The wing (Plate IX., Fig. 2, p. 126) is broader than in JV. zoctula, the
lower leg, tail, forearm, and fifth metacarpal being all longer than in

THE SEROTINE 133

that species. The third metacarpal is slightly shorter than in xoctwla,
and is slightly longer than the fifth; the latter is relatively much longer
than its forearm. The foot (both actually and relatively) is smaller,
and the longest digit shorter. The tail projects markedly from the
interfemoral membrane, the last two vertebre being free. The thumb
is longer, and more slender. The total expanse is actually about the
same as in JV. zoctula, but is less relatively to the forearm length.

The fur is soft and longer than in (Vyctalus, but on the upper sur-
face extends on to the wing to a lesser degree; beneath, the wing is
sprinkled with fine hairs, and on the interfemoral membrane these are
restricted to an almost invisible scattering on the transverse dotted
lines. As in JV. noctula, a band of hair may extend from behind the
forearm to the carpus, but it is usually much less conspicuous.

The colour above is dark, varying
Between “clove” and “ Prouts,’ or
“mummy brown”; many of the hair
tips are of tawny shades. The under
side affects some tint of “broccoli”
brown, often inclining to tawny. The
line of demarcation is sometimes mode-
rately distinct, following the wing, and
thence running forward beneath the ear
to the mouth. The sparsely haired
portions of the face are deeply pig-
mented, as are the wing, interfemoral
membrane, limbs, and ear.

Bell states that the juvenile colours
are more obscure, and Ruskin Butterfield
writes me that the undersides of old
specimens are conspicuously darker,
but it is not known whether there is
any seasonal colour change or moult. vo
A very dark female in good condition,
shot by Ogilvie-Grant at Yalding, Kent,
on 23rd May, suggests that there may
be, as in so many bats, two phases, a bach alto aight ae
melanic and a brown, and seems to be AND CANINES OF Vespertilio serotinus
contrary to Bell’s statement that “the (enlarged and diagrammatic).
female is much brighter than the male.” !

In the teeth (Fig. 12) the inner upper incisor is long, strong, and
bifid, until worn down: its length is at least double that of the small

1 A similar sexual coloration is, however, present in Minéopterus dasythrix
(Temminck) and M. fraterculus of Thomas and Schwann (see Proc. Zool. Soc.
(London), 20th Feb. 1906, 161-162).

N 2

134 VESPERTILIONIDA:—VESPERTILIO

outer incisor. The anterior lower premolar attains to about half the
breadth and height of the posterior.

Variation :—This species has always been regarded as to a high
degree variable, but it is probable that many of its phases will eventually,
under modern criticism, prove to be of geographically specific or sub-
specific value. Little more than a note of the various described forms
can here be given. Daday distinguishes a large Transylvanian sub-
species as ¢ransylvanus, and Cabrera one with the forearm measuring
51 mm. from the Balearics as zzsularis ; the same author’s doscaz from
the Mediterranean coast of Spain was based on a young serotinus
(Miller). A number of cream-coloured Asiatic bats have been referred
to serotinus, but at least those bearing the names zsabellinus (Tem-
minck, 1827, forearm 41, from Tripoli), ¢urvcomanus (Eversmann,
1840, expanse about 200, from the Aralo-Caspian steppes), and
botte (Peters, 1869, forearm 40, from Botta, Arabia), are clearly not
conspecific. MJzrza (Philippi, 1865, forearm 53), and shzrazdensis
(Dobson, 1871, forearm over 50), both from Persia, may represent
desert forms. Blanford’s andersoni of Yunnan is said to have a smaller
foot, while Tomes’ pachyomus (forearm 52) of India is a distinct bat
(Miller, Proc. Biol. Soc., Washington, xiii, 13th June 1900, 155-156).
Chinese examples are of dark coloration.

DIMENSIONS IN MILLIMETRES :—

a 3 : 6
bo a ° = : 2
eal eB be a pM de SN al 1) eae
S| 4 2H | FPS Se EPS) | rot a S
a +> Zo a za = Le} ce} = Bi ma
no! m2 fo7_ Le HH -= 3 a ~ Sa See oO =
r=| SB at 3 3) 23 2 3 a BIEr aie ae
So eee Nos Ses Ail (ete ieeal imc 8 ete hae rc Wes he 2 E
os B = Sora & |g a |g S 2
a 8b s Hig 5 ole a $
4 = = a a} 4 2
3 =a
g ea) o
Male (in the flesh), Essex
(Miller Christy, Zoologist,
1894, 424) te ac 32 aie te 32 AC Sn leeds Be bc 355
Male do., Yalding, Kent, 28th
July 1906 3 79 | 20 8°5 | 51°5 | 20 11°56 | 50°5]8 85 47 47 348
Male, do., Surrey, ‘6th “Aug.
(W. R. Ogilvie-Grant) . - | 68 | 19 6 52 22 = 53 Bee Weiss 47 43°5
Male, Brede, Sussex . gall ee is - ie .. | 49 rs x sts
Female (in the flesh), Yalding,
Kent (W. R. Ogilvie- Grant),
7th Aug. 1897 . 72 | 15°5 47 52 20°5 9 53 7 89 49°5 | 46 354
Female, do., 8th Aug. 1897 A Wet Ss 50 55 21 10 54 i; 87 47 45 355
Female, do., 18th May 1898 .| 80] 19 te 52 21 36 51 7 84 45°5 | 41 3874
Female, do., 8rd July 1898 A033 |) 2283 6 54 no ae 53°5 |6°5| 85 48 44°5 | 374
Female, do., 2nd Oct. 1898 seca LG ore 48 20°5 | 10 53 7 90 47 44 365
Female, do., 2nd Oct. 1898 = | i) ao os 50 21 12 52°5 | 7 84 45 41°5 | 356
Female, Surrey, 6th Aug. 1902
(W. R. Ogilvie-Grant) . |) ee | PE 9 57
Average of females . |73 | 17°9 are 52°5 | 20°8 ae 52°8 |6°9| 86°5 | 47 43°5 | 368
Do. of both sexes . |74°2}18-25] .. 52°38 | 20°8 | 10°5 | 52:1 | 7 86°6 | 47 44 360

Dowker (Zoologist, 1891, 425) gives the wing expanse of two Kent specimens as reaching 14°5 and
15 inches =368°3 and 381 mm,

Proportionate lengths (both sexes) :—Foot without claws, about -50

THE SEROTINE 135

of lower leg; fifth metacarpal, about -93 of third; lower leg, about
-39 of forearm and about -28 of head and body.

Skull:—Greatest length, 21; basal length in middle line, 16;
palatal length in middle line, 8-5 ; from posterior border of third upper
molar to anterior border of canine, 7; same in lower jaw, 8-5; greatest
breadth at zygoma, 14; posterior breadth,9; breadth between orbits, 7 ;
breadth at constriction, 4:5.

Weight of a male from Essex, § 0z.=113 grammes (Miller Christy,
Zoologist, 1894, 424).

Distinguishing characters:—As compared with the Noctule, which
alone of preceding species is of similar size, this bat is deep brown,
with lighter under side, instead of wholly coloured reddish brown;
the colour is alone distinctive, but the cumulative evidence of broader
wing, longer ear and tragus, slight post-calcarial lobe, and greater
length of tail outside the interfemoral membrane, make error in identifi-
cation impossible.

The Serotine, notwithstanding the clear and intelligible
description of its discoverer Daubenton, was mistaken for
the Noctule by Isidore Geoffroy, but was well figured in the
eighth volume of Buffon’s great work. It is in the British
Islands only known from the south, and particularly the south-
east of England, where it is abundant, and perhaps the com-
monest bat, a very remarkable fact when it is remembered that
outside of Britain probably no bat has a wider distribution.

This fine bat has gained its name from the late hour at
which it is supposed to commence its evening flights. But
in this respect, and in other aspects of its natural history,
it has been frequently confused with the Noctule, and there
is no doubt that much that has been written of it is
incorrect. Thus, nearly every account of it—and the most
original are those of William Borrer,’ G. B. Buckton,’ Frederick
Bond,* and George Dowker *—mentions two quite distinct forms
of flight, difficult to reconcile as appertaining to the same
creature, and inconsistent with the descriptions of the best
foreign naturalists. In the first the bat is shown flying
with great strength at a high elevation in manner strongly
suggestive of the Noctule. In the second it appears as a

1 Zoologist, 1874, 4126. 2 Proc. Linn. Soc. (London), 6th Dec. 1853, 260 (1855).
3 Field, 14th March 1874, 263 (quoted by Edward Newman).
* Zoologist, 1891, 305-306 and 425.

136 VESPERTILIONIDA-—VESPERTILIO

somewhat delicate animal, not daring, as a rule, to venture
abroad except when the weather is fine, and seeking the shelter
of gardens and orchards, roads or lanes, where it appears in
the same haunts from year to year. Its flight is described as
low and heavy, and it often flutters with fully expanded wings
in manner resembling the members of the genus JZyodzs.

I am indebted to Captain Saville J. Reid for enabling
me to arrive at a partial understanding of this apparent con-
fusion. In the grounds of his house at Yalding, Kent, he has
long known the Serotine to haunt the lawns and open spaces
between the fruit-trees. Here, in the early summer months,
a party of ten or twenty of them may be seen circling and
twisting round the trees in pursuit of the buzzing cockchafers,
which they catch and devour on the wing. Captain Reid’s
experience early in the year thus bears out the descriptions
of those writers who find this bat a sociable, low-flying, glade-
haunting species, but he has not observed it actually snatching
the insects off the trees, as other writers assert that it does.’

On 28th July, when I joined Captain Reid in watching
the bats, there were no cockchafers on the wing, and the
Serotines, of which there were then only some half-dozen in
view, had perforce to look for other food. They now flew
higher, often at thirty or forty feet, but not, I think,
exceeding the height of tall elms or of gunshot, and
often descending near to the ground. Their flight was
not unlike that of the Pipistrelle, but their beat was wider
and their pace relatively less rapid. They could not be
described as weak flyers, nor was their pace slow, but they
clearly lacked the dash and finish of the Noctule, one or
two of which were present for comparison.

Like other bats, they indulged in frequent swoops and
somersaults, evidently in the act of seizing their prey, but
the only one heard to utter a sound was a large one, which
Mr H.C. Schwann shot to test our identification, and which
shrieked before it fell dead to the ground.

To the above notes may be added the information kindly
supplied by the Rev. E. N. Bloomfield, of Guestling Rectory,

' £.g. Biasius in Germany, Borrer and Bond in England; see Zoologist, 1891,
203.

0 EE

THE SEROTINE 137

Hastings, that a colony occupying a cavity in the roof of
his house, vary their flight with the weather. When a breeze
is blowing they seek the shelter of trees, and fly backwards
and forwards along the short drive at the Rectory. In
calm weather they venture farther afield.

As far then as these very limited observations go, the
Serotine usually feeds at moderate, not necessarily humble,
elevations. It looks fully as large as the Noctule, but may be
distinguished by the breadth of its wings, apart from the two
different styles of flight, which suggested to Dowker and to
Captain Reid a comparison between snipe and woodcock.

The food evidently varies with the season and oppor-
tunity. At Yalding cockchafers seem to be the most highly
esteemed, but, when these insects are not present, substi-
tutes must be found, and Buckton saw the bats pursuing
white moths,’ as well as beetles, and successfully angled three
of them by means of shreds of white paper attached to an
ordinary fishing rod and line. At Yalding, when the cock-
chafer flights are over, they seek some more congenial hunting-
grounds, and by autumn the garden is almost deserted.

Despite its name, the Serotine is an early flier, perhaps
the earliest of all British bats. Captain Reid has observed
it in numbers soon after eight in the evening in the third
week of July, the sun setting at about eight. On the 29th,
the sun then setting at a few minutes before eight, several
made their appearance at about sunset, the first at about
twenty minutes to the hour. Buckton believed that, like the
Noctule and Leisler’s Bat, this species retires after about an
hour’s exercise in the evening, and that was the impression
which I gained from my brief acquaintance with it; but further
study is required before this point can be regarded as settled.”
I watched for the Yalding bats from a quarter to two to four
o'clock on the morning of 29th July, the sun rising at about
twenty minutes past four, but without seeing a single one,
although a small bat was very busy on the wing for a few
minutes at about four.

1 Porthesia chrysorrhea, Linnzus.

2 Miller Christy’s (Broomfield, Essex) specimen entered a room at I A.M., on
25th August, and A. B. Hutton’s bat, from the same county, after midnight,-on
27th August (Henry Laver in lit.).

138 VESPERTILIONIDA—VESPERTILIO

In the daytime the Serotine retires to holes, which a num-
ber of individuals inhabit in company. They seem to have
a preference for cavities under the roofs of houses, and Captain
Reid states that they do not object to the presence of other
species. Borrer’ knew of acolony of at least eighteen individuals,
while Mr H. G. Jeffery,” who sent one to Mr J. E. Harting,
has described another of about twenty; but assemblages of
numbers rivalling those of the Noctule seem to be unknown.

Hibernation probably commences for the species as a whole
in the latter part of October, but several observers have noticed
a few on the wing in November,* as Mr Percy Wadham in the
Isle of Wight until the roth; he remarks, however, that these
late-flying specimens haunt unusual places, and _ restlessly
change their beat every few minutes. The dates of first spring
appearances have not been placed on record.

Nothing is known definitely about the breeding habits,
except statements by continental naturalists (Blasius and Kuhl)
that only one young one is brought forth at a time, its birth
taking place generally in the latter half of May, and that in
other respects it is not known to differ from other species.

A bat of this species is said to have been kept alive by
Mr George Guyon* from 3rd January to 7th March 1856. It
was taken while hibernating in an old chimney, and immediately
after its capture ate some raw meat left with it, and in four
days it would take meat from the fingers and allow itself to
be stroked. ‘That it knew me,” wrote Mr Guyon, ‘I would
hardly venture to say ; but certainly on one occasion it squeaked
in alarm when another person offered to touch it, which it
never did with me after the first few days.”

The Serotine is frequently mistaken for the Noctule, but,
except in size, the two have few points of resemblance. The
generally darker colour of the Serotine, the lighter under side,
broader wings, longer and more pointed ears, and lanceolate
tragus, very slightly developed post-calcarial lobe, and failure
of the interfemoral membrane to extend to the tail tip, are quite

1 Zoologist, 1893, 223-224. 2 Jbid., 1894, 261.
3 E.g., Rev. E. N. Bloomfield at Hastings (in lit.) ; also A. G. More, in Wight, on
3rd Nov. ; see Life and Letters of Alexander Goodman More, etc., 34, 1898, edited

by C. B. Moffat (Dublin: Hodges, Figgis, & Co., Ltd.).
4 Zoologist, 1856, 5216.

THE PARTI-COLOURED BAT 139

diagnostic. Both species have a band of hair on the arm, so
that this point is of no value for purposes of determination.

On the wing, the Serotine can only be confused with two
other British bats, the Noctule and the Greater Horseshoe.
Its flight has already been compared with that of the former,
which rarely descends to hunt insects around the lower branches
of trees, and both are. conspicuous enough to exhibit the
different proportions of their wings in flight.

[THE, PARTI-COLOURED., BAT.

VESPERTILIO MURINUS, Linnzus.1

The Parti-coloured Bat was included in the British list
by Bell, on the strength of a single specimen taken by
W. E. Leach at Plymouth, probably in the early thirties,
and now in the British Museum.’ A second example came
into the hands of John Hancock when ‘either alive or jus¢
dead.” It was taken (as he wrote to Mr Thomas Southwell),
“JT am almost sure, on board ship, undoubtedly off Yarmouth
Roads, in the year 1834.”

The coloration of this species is so remarkable that, were
it a true native of Britain, it could not for so long have escaped
the attention of zoologists. We may, therefore, conclude
that its visits to this country have been entirely accidental.
Posstbly they may have been effected by the assistance of
shipping, or it may be that the individuals which have reached
our shores have been blown across the Channel while engaged
in the migratory movements which both the Parti-coloured
and its ally, Nilsson’s Arctic Bat,* are believed to perform
annually. With the prevailing winds blowing from and not
to the British Isles, however, its frequent occurrence within our
boundaries would seem to be improbable.

1 Systema Nature, x., 32 (7), 1758: until recently cited as Vespertilio discolor of
Natterer.

a. No, 374.

* Southwell, Zvans. Norfolk and Norwich Nat. Soc., 1873-74, 80; (1 am indebted
to Southwell for further particulars by letter); E. Newman, /%e/d, 7th March 1874,
218, and Feld, 5th Sept. 1874, 246.

1 V. nilsson.

140 VESPERTILIONIDA:—MYOTIS

It ranges from southern Sweden to Dalmatia, and from
France to Asia, and is, perhaps, representative of the American

genus Laszonycterts.
The rich tints of this species, dark brown marbled with

pale yellowish brown, and the strongly contrasted whitish under-
parts, make it one of the most beautiful of European bats.
It may be at once distinguished from the Serotine by its colour
and smaller size, the forearm measuring only about 43-45
mm. From Nilsson’s Bat also it differs in colour, the latter
being deep brown grizzled with tawny above and yellowish
brown beneath, in its slightly larger size, the forearm of the
latter averaging 39-45 mm., and in having the tragus broadest
above, instead of below, the middle of the inner margin. |

GENUS MYOTIS.

1829. Myotis, Jakob Kaup, System der Europiischen Thierwelt, 1., 105-106, 188 ;
based on Vesfertilio murinus of Schreber (not V. murinus of Linnzeus) ; Gray,
Ann. and Mag. Nat. Hist., December, 1842, 258.

1829. NYSTACTES, Jakob Kaup, of. cz¢, 1, 106, 108-109; based on Vespertilio
bechsteini of Leisler ; name preoccupied by NYSTACTES of Gloger, 1827, a genus of
birds (see Palmer, /udex, 467, 1904).

1829. VESPERTILIO, Jakob Kaup, of. cz4, i, 110; based on Vesfertilio daubentont,
V. natterert, and V. mystacinus; A. Graf von Keyserling and J. H. Blasius,
Wiegmann’s Archiv fiir Naturgeschichte, i., 307, 1839; Harrison Allen, 1864 and
1893 ; Dobson, 1878 ; not VESPERTILIO of Linnzus, 1758.

1830. LEUCONGE, Friedrich Boie ; Oken’s /s¢s (Jena), 256-7 ; based on die Wasser-
fledermiause, type V. daubentonitz.

1841. SELYSIUS, C. L., Prince Bonaparte, Fauna Jtalica, i., introduction (3); based
on Vespertilio mystacinus of Leisler.

1841. CAPACCINIUS, C. L., Prince Bonaparte, Fauna Jtalica, 1., introduction (4) ; based
on Capaccinius megapodius of Bonaparte = Vespertilio capaccinit of Bonaparte.

1842. TRILATITUS, J. E. Gray, Ann. and Mag. Nat. FHist., 258, December: in-
cluded Vesperiilio hasseltit of Temminck, V. macellus of Temminck= V. adversus
of Horsfield and 7. dlepotis of Gray = Miniopierus sp.

1849. TRALATITUS, F. L. P. Gervais in Charles D’Orbigny’s Dictionnaire Universel
@ hist. Nat., xiii, 213; a misprint for 7rzlatitus.

1856. BRACHYOTUS (sub-genus), F. A. Kolenati, Allgemeine deutsche Naturhist.
Zeitung (Dresden), Neue Folge, i1., 131, 174-177; based on mystacinus of Kuhl,
daubentonit of Kuhl, and dasycnemus (sic) of Boie; name preoccupied by
BRACHYOTUS of Gould, 1837, a genus of birds.

1856. IsoTuS (sub-genus), F. A. Kolenati, of. cz¢., ii., 131, 177; based on mattereriz of
Kuhl, and cz/éatus (p. 131) or emarginatus of Geoffroy (pp. 177-179).

1856. Myotus, F. A. Kolenati, of. czz., ii., 131, 179-181 ; based on dechsteiniz of Kuhl
and murinus of Linnzeus.

a ee

MYOTIS I4I

1866. TRALATITIUS, J. E. Gray, Ann. and Mag. Nat. Hist., 90, February ; a mis-
print for 77zlatitus.

1867. PTERNOPTERUS, Wilhelm Peters, Monatsberichte der Koniglich Akad. der
Wissenschaften (Berlin), 706; a sub-genus of VESPERTILIO = Myotis, based on
P. lobipes of Peters=V. muricola of Hodgson.

1870. EXOCHURUS, L. J. Fitzinger, Sztzungsberichte der Katserlichen Akad. der
Wissenschaften (Vienna), Ixii., 1, 75-81; based on Vesfertilio macrodactylus of
Temminck, V. horsfieldii of Temminck, and l. macrotarsus of Waterhouse.

1870. AEORESTES, L. J. Fitzinger, of. cz¢., 427-436; included Vespertilio villostssimus
of Geoffroy, V. albescens of Geoffroy, V. néigricans of Maximilian, and V. levis of
Geoffroy.

1870. COMASTES, L. J. Fitzinger, of. cz4, 565-579; included Vesfertilio capaccini of
Bonaparte, V. megapodius of Temminck, V. dasycneme of Boie, and V. limnophilus
of Temminck.

Classification and synonymy :—As shown by Mr Miller, the
genus Vespertilio of Linnzeus contained none of the large group
of bats with thirty-eight teeth, to which the name was applied by
Bell and others. These have, moreover, clearly nothing to do
with the thirty-two toothed members of that genus as here
applied, and have accordingly been allocated by recent writers
to the genus A/yotzs of Kaup, the first based upon a member of
the group.

This large genus comprises a multitude of species, some
of which are found in nearly all tropical and temperate regions
of the globe, so that the area of their distribution is probably
unexcelled in extent by that of any other bats.

There are four British species —daubentont, mystacinus,
the rare dechstetnt, and natterert. A fifth, the common con-
tinental JZ. myosotzs (see page 190), has often been included in
works on British natural history, but is certainly not a regular
inhabitant of this country; a sixth, JZ. emarginatus, is stated
to have occurred once (see page 189). Other European
species are MW. capaccin? of Bonaparte, a bluish-grey bat, with
narrow, hairy, interfemoral membrane and elongated backward
pointing, falcate tragus; JZ. dasycneme of Boie (see page 157) ;
and the Mediterranean JZ. oxyguathus (Monticelli).

The generic characters are slender, delicate form, long tail,
hairy face, narrow ear, and tapering, straight, or recurved
tragus, combined with variable size.

The muzzle is long, the forehead not prominent, the face
hairy, the glandular prominences being much less developed
than in WVyctalus, Piprstrellus, or Vespertilio, and scarcely add-

142 VESPERTILIONID#—MYOTIS

ing to the breadth of the face. The nostril opens without
prolongation sub-laterally, the aperture being crescentic.

The ear (Fig. 2, Nos. 5-8, p. 7) is long and oval, distinctly
longer than broad ; the outer margin terminates anteriorly below
the base of the tragus, or very slightly in front of it, and does not
run forward towards the angle of the mouth; the internal basal lobe
is angular. A number of cross-folds or pleats running horizontally
across the ear, near the outer margin,
are characteristic of the various species,
but are often inconspicuous after death.
The tragus is long, narrow, and gene-
rally attenuated to a point.

The foot, lower leg, and calcar are
of variable size and development.

The wing is broad, or moderately
so, the fifth metacarpal being slightly
longer than or about equal to the third ;
the attachment may be from a little
above the ankle, to the base of the
toes. The post-calcarial lobe is poorly
developed. The tail is rarely so long
as the combined head and body.

The skull varies in strength accord-
ing to the size of the species: it has

2 usually a rounded brain-case, elevated
above the facial region, which is narrow,
depressed, and markedly saddle-shaped:
the premaxillary gap is developed to a
degree about intermediate between that
of Myctalus and Vespertiho , the zygo-

Fic. 13—D1AGRAM or ArraNcE- Mata are flattened, the auditory bulle
oe oF, TEETH IN'GENUS moderately developed.
The number of teeth (Figs. 8, p. 101,
13, and 16, “p. 174) 1s increased "te
thirty-eight by the addition of a third premolar in each jaw and
side, so that the formula is—

(1) Upper and (2) Lower Jaw.

The upper incisors are sub-equal, their points generally diver-

DAUBENTON’S, OR THE WATER BAT 143

gent, the outer slightly outwards, the inner inwards. The
two anterior upper premolars are small, especially the central,
which is often minute and crowded out of the tooth row
internally. The last upper molar is rather less in section
than half the second. The lower outer incisors are much
larger than the inner. The lower premolars follow the relative
proportions of the three upper, but the central, although the
smallest, is rarely so minute as the corresponding tooth in
the upper jaw.

Group Leuconde :—Myotis, as shown in extra- British species,
forms an unwieldy and heterogeneous assemblage, which is
certain to be subdivided as the relationships of the bats
included within it become better known.

The first subdivision to suggest itself is that of which JZ.
daubentonz is the sole British representative, and the members
of which are characterised by their large feet. This character
in daubentonz, sufficiently evident as it is, reaches an extreme of
development in vzcketz of Thomas. For these bats the name
Leuconée, first used by Boie in 1830, is conveniently available,
but, unfortunately, the sharp definition of the group is destroyed
by certain exotic intergrading forms, and thus cannot be upheld
on careful analysis of the dimensions even of British species
alone. The small ciliated interfemoral of A/. nattererz is a more
distinct character.

DAUBENTON’S, OR THE WATER BAT.
MYOTIS DAUBENTONI (Kuhl).

1819. VESPERTILIO DAUBENTONI, Heinrich Kuhl, Meuwe Ann. der Wetterauischen
Gesellschaft fiir die gesammte Naturkunde, i., ii., 195, pl. xxv., fig. 2 ; described from
Leisler’s MSS. from Hanau, Germany ; Bell (ed. 1) ; MacGillivray ; Blasius ; Cler-
mont ; Fatio; Bell (ed. 2); Dobson; Blanford ; Flower and Lydekker ; Lydekker.

1828. VESPERTILIO EMARGINATUS, John Fleming, British Animals, 6; Jenyns;
not V. emarginatus of Geoffroy.

1830. LEUCONGE DAUBENTONI, Friedrich Boie, Oken’s /szs (Jena), 256-257.

1839. VESPERTILIO £DILIS, Leonard Jenyns, Anm. Nat. Hist, ii1., 73, pl. iii., April ;
described from an albinic variety, from Auckland St Andrew, Durham, England.
1840? VESPERTILIO VOLGENSIS (? species), Eduardo Eversmann, Bull. Soc. Impériale
des Nat. de Moscou, i., 24; described from Des Kasanischen und Nisch nigoro-

dischen Gouvernment, und im Uralgebirge ; placed here by Dobson, 1878.

1844. VESPERTILIO PELLUCENS, J. Crespon, /aune Méridionale, 1, 16; vide
Trouessart, Bull. de la Soc. ad Etude des Sci. Nat. de Nimes, 7, i. 35-39, 1879;
described from Nimes, France.

144 VESPERTILIONIDAA—MYOTIS

1855. VESPERTILIO DAUBENTONH, var. EMARGINATUS, G. B. Buckton, Proc. Linnean
Soc. (London), 6th Dec. 1853, 260.

1856. BRACHYOTUS DAUBENTONII, F. A. Kolenati, Ad/gemeine deutsche Naturhist.
Zettung (Dresden), Neue Folge, ii., 131, 175-176.

1887. MyoTIs CILIATA,! J. Daday, Ertekezések a Természettudominyok Korébola
(Budapest), xvi., 7, 35, 1886; not Vespertilio ciliatus of Blasius, vide Méhely, 330.
1898. MYOTIS DAUBENTONI, Oldfield Thomas, Zoologist, 100; Collett ; Johnston ;

Méhely ; Cabrera; Muillais.

1910. MYOTIS (LEUCONGE) DAUBENTONI, E,-L. Trouessart, Faune des Mammifcres
ad’ Europe, 27.

Die Wasserfledermaus of German writers, a book-name merely,
there being probably no local names for the lesser-known small bats
either in Britain or elsewhere.

Distribution :—This Water Bat, or very close allies, attains a
comparatively northern latitude throughout boreal and transitional
Europe and Asia, ranging from south Sweden and Norway (Collett),
Finland, and middle Russia, to Sardinia, Sicily, and Galilee (Tristram) ;
and from Ireland to the Altai, and south to Tenasserim (Dobson).
It reaches at least 2000 feet in the Hartz Mountains, and 4000 feet in
the Alps. It is not mentioned by Cabrera as occurring in Spain.

It is probably an abundant species in every part of England and
Wales, affording suitable combinations of water and woods. There
exist reliable records of its occurrence in every county of England
except Wiltshire, Somerset, Gloucester, Buckingham (doubtfully),
Huntingdon, Rutland, Nottingham; but, inasmuch as each of these
counties is bordered by others in which the bat is well known, there
can be little doubt that it will be found in all of them. It ascends to
at least 1000 feet, having been observed by Oldham in the ornamental
cardens at Buxton, Derbyshire, at that elevation. Its presence in
Wight, long ago reported by Bury (see More, Zoologist, 1894, 148), is
now confirmed by Wadham.

In Wales it is definitely known only from Carnarvon, Denbigh,
Flint, Merioneth and Montgomery (Forrest). But it is common in
several at least of its known haunts, and since the discovery of these is
chiefly due to the recent work of Caton Haigh, Oldham, and Forrest, no
doubt further exploration will reveal its presence in the remaining
counties.

In Scotland it is reported from the mainland as widely distributed,
although local (Alston), and it occurs at least as far north as Fochabers,
in Banff, whence a colony was reported in 1892 (Eagle Clarke,
Ann. Scott. Nat. Hist., 1892, 266). In parts of Aberdeen Sim found it

1 Vespertilio capucinellus and Vespertilio minutillus are cited under this species
as of “Koch, Bayr. Fauna,” but I cannot find the original reference ; see Fitzinger,
Sitzungsberichte der Katserlichen Acad. der Wissenschaften (Vienna), Ixiii., 1, 206.

DAUBENTON’S, OR THE WATER BAT 145

“by far the most common and abundant Bat”; and in Elgin it is not
uncommon in some parts of the valley of the Spey (Kinnear); and it
may be expected further north. A colony has been recorded from
Kinlochaline Castle, Morven, Argyll (Charles Campbell, /ze/d, 31st July
1897, 222); it was found abundantly in Perth by Grieve (Aun. Scott. Nat.
Hitst., October 1894, 193-195), and in Solway is commoner than P. /zp7-
strellus (Service), but definite records are still needed for many counties.
In the Highlands it reaches at least 512 feet above sea-level (Grieve).

In Ireland the bat is as yet but little known. The first known
specimen, obtained in Londonderry, was submitted to Jenyns for identi-
fication by the Ordnance Collectors (Thompson), the date being given
by Jameson as 1838 (/rzshk Naturalist, 1897, 39). It was next encountered
by J. R. Kinahan in Kildare in 1853, and was at first identified with J/.
natterert (Nat. Hest. Review (Dublin), i., 23-25 and 87, 1854), but assigned
by Bell to the present species (of. c7t., i., 148-149; also vi., 383, 1859).
Subsequently Jameson (/7zsh Naturalist, loc. cit., also 1896, 94) secured
specimens in both counties Fermanagh and Louth; in the former
case in Bohoe Cave, in the latter at Branganstown on the river Glyde.
Barrington received one from the Lucifer Shoals Lightship, nine miles
off the Wexford coast (see below, p. 153). It had been caught on 21st
April “flying low over the deck,” at 7.30 P.M. Lastly, E. B. Knox sent
one to Alcock from Bray, Co. Wicklow (/vzsh Naturalist, 1898, 256). No
other reputable records exist, that given by Knox in his History of
Down being unsupported by evidence, and that by Lydekker for Donegal
having apparently -been copied from Bell, in whose second edition
Thompson’s Londonderry record is credited to Donegal. The species
thus ranges from north to south of the island, and, since it was found
numerously in its Kildare, Fermanagh, and Louth localities, and was
established in a distinct colony when found in Wicklow, there can be no
doubt that our knowledge of its distribution is defective, and it may yet
prove to be a widely spread and abundant species.

Distribution in time:—Bones from the Pleistocene deposits of
Ightham Fissure, Kent, have been provisionally associated with this
species, or with A/yotzs mystacinus, by E. T. Newton (Quart. Journ. Geol.
Soc., August 1899, 420; see also Abbot, Journ. czt., May 1894, 171-211),
as have leg-bones from superficial deposits in the caves of Co. Clare,
Ireland (Scharff, Zrans. Roy. Irish Acad., Feb. 1906, 53); but, in the
absence of skulls, it were well to accept the identifications with caution,
and in the latter case, as Scharff points out, no conclusions affecting
the age of this bat in Ireland can be based on such fresh remains.

The period of gestation is unknown, but probably does not differ
widely from that of Pzpistrellus pipistrellus.

Breeding season and number of young:—A single young one is
born, most frequently in June and July.

146 VESPERTILIONIDAZ—MYOTIS

Description :—In general form and appearance this bat is typical
of the group Leuconde. In size it is somewhat larger than Pzpzstrellus
pipistrellus, but distinctly smaller than Wyctalus lecslert.

The ear (Fig. 2, No. 5, p. 7), when laid forward, reaches about to the
nostril; the outer margin is very slightly notched, the upper two-thirds
being straight or only slightly concave, the lower third abruptly convex ;
a deep emargination opposite the base of the tragus gives rise to a
distinctly rounded basal lobe; the inner margin is convex almost
throughout its length, most markedly at its centre, whence it passes
upwards to a broadly rounded tip; there are four cross-folds.

The tragus, which is about half as long as the ear, reaches its greatest
breadth at about its centre, is straight, and not sickle-shaped, and tapers
to a more or less acute point; the inner margin is about straight, the
outer gently convex, with a distinctly rounded triangular lobe projecting
just above the base.

In the wing (Plate VII., Fig. 3, p. 86) the most noticeable features
are the extremely long lower leg and large foot ; a well-developed spur
or calcar extends fully three-quarters of the distance from the ankle to
the tail, the tip of the spur projecting from the posterior border of the
interfemoral membrane as a small, but distinct, lobe. The third digit is
short, so that the wing as a whole is of less than average length. The
wing-breadth is moderate, the fifth metacarpal being distinctly shorter
than the third. The tail is short, and has usually two free vertebre.

The fur is short, but soft and plentiful. The,hairs on the ear are
small and inconspicuous. The face is half-naked and rather tumid before
the nostrils, but the muzzle carries a moustache composed of numerous
long hairs (Plate X1., Fig. 2, p. 140). Both surfaces of the wing are furred
as far as a line running from the centre of the humerus to slightly below
the head of the lower leg; the upper surface and the posterior border
of the interfemoral are ciliated, the latter inconspicuously. The toes are
provided with whitish hairs.

The colour above is warm brown, of some shade between “mars
brown” and “mummy brown,” the bases of the hairs darker and the
tips lighter, resulting in a grizzled appearance; below, near light
“broccoli brown,” the hairs plentifully tipped with dirty white, or
yellowish. The line of demarcation is moderately distinct, and runs
approximately from the angle of the mouth to the thigh. The wing is
dusky, with a reddish tinge, the interfemoral membrane whitish beneath.

I have had no series from which to study variation, seasonal or
otherwise, but the general colour is somewhat variable, perhaps owing to
age; the young are sometimes described as darker (Bell, Millais), some-
times as greyer (Coward and Oldham). Jenyns’ description of Vesper-
tlio edilis was based on an albino specimen.

In the skull (Fig. 8, No. 2, p. 101) the profile of the cranium is almost

DAUBENTON’S, OR THE WATER BAT 147

horizontal ; that of the face descends in a slightly concave line at an angle
of about 15°. The upper incisors are about equal in size: they are
short and broad (Fig. 14), the breadth of the

crowns about equal to their length, bicuspid,

and with the cusps strongly divergent. The ————

two anterior upper premolars are both in the

tooth-row, but the central, which is much the

smaller, is slightly displaced internally; its

conical tip reaches about to the cingulum of

the posterior. The lower outer incisors are

oval in section, being about one and a half

times as long as they are broad, and less than

half as thick as the canines.

Geographical variation has not been
worked out in this species, but there is
little doubt that had JV. carisstma of North
America been taken in England, it would
have passed muster as daubentonz. ee ae once a

Myotis daubentoni (enlarged

Dimensions in millimetres (see page and diagrammatic).

148).

Proportionate lengths (males) :—Foot, without claws, about -58 of
lower leg; fifth metacarpal, about -94 of third; lower leg, about -44
of forearm and about -33 of head and body.

Skull :—Greatest length, 14-5; basal length in middle line, 11;
palatal length in middle line, 6; from posterior border of 7° to anterior
border of canine, about 5; same in lower jaw; greatest breadth at
zygoma, 9; posterior breadth, 7-5; breadth between orbits, 7-25;
breadth at constriction, 4.

The weight is given by Couch (Zoologzst, 1853, 3942) as 97 grains
=6-44 grammes. The same writer (p. 4012) measured one having an
expanse of 10-5 inches = 251-20 mm.

Distinguishing characters:—Of preceding species, the Pipistrelle,
which is slightly smaller, can alone be confused with this bat. But the
pointed ears; lanceolate tragus; little developed post-calcarial lobe;
light under side; large foot, lower leg and calcar; and grizzled upper
surface, are quite distinctive.

FIG. 14.—FRONT VIEW OF IN-

Daubenton’s Bat, although one of the most abundant and
characteristic of British mammals, remained for long unrecog-
nised in this country, and affords a striking instance of the
light thrown upon our smaller species by the more exact studies
of the last years of the nineteenth century. It was first made
known to science by the labours of Leisler and Kuhl in Germany.
In England, Bingley, who had certainly observed it in its well-

148

DIMENSIONS IN MILLIMETRES :—

VESPERTILIONIDA—MYOTIS

MALES.
Gavel gat g : .
oO a ° . _ . Oo
eye C8l hele ae |e | od ea
~ 2o see = Eo 5 ue} s 3S S an
a) 2 | S23) 8.) 8 lef] 2 -) 8 |e eal eee
3 3 R@2 a ES om q a 2 = 5) +e
us! 2 AS } ge Se 8 =) 3 s a
$ 80 3 Hols 3 i} 5 2 =
Be) sel 8 & A 2a eee
a
cs a o
Male (from spirit), North-
ampton . 6 Ee 17 9 87-5 5 57 31 29°5
Male, Middlesex 4 W755 8 38 6 58 82 30
Male(from spirit), Temple
Combe, England, Oct.
1907 > 50°4 | 12°6 6 80°6 | 15 9 35°6 iG 59 32 29°2| 244
*Male, Merioneth, Wales,
31st March, 1891 . - | 41 12°5 6°75 | 35°5 33 202
*Male, Strathspey, ‘Scot-
land, 5th Sept. 1891 . | 43 9°5 7 36 17 1lt 36 226
*Male, Loch Dochart, W.
Perth, Scotland, 25th
July 1894 - 46 12°5 74 | 35 17 7-5 | 388 227
*Male, do., 25th July 1894 46 14°25] 7:4 |383°5 | 15°75) 8 37 226
*Male, do., 15th July 1894 | 43 14°25] 6°4 | 37 16 10°5t | 38 228
*Male, do. . 43 14°25| 5:8 | 37 16°5 | 10t 38 227
*Male, do., 15th July 1894 49°5 |17°5 64 | 31°5 ae 10t 34°25 201
Male, Bohoe Cave, Fer-
managh, Ireland, 11th
July 1895 (Dublin
Museum), from dried
skin 26 15 10°25 | 35 5°95 | 57 31 30
Male, Co. Louth, 6th July
1895 (Dublin Museum),
from dried skin 31 15 10 34°25 | 5°5 | 60°5 | 382 30 me
Two males (from spirit), ‘]
Bohoe Cave, Fer- || 44 30 18 10°25 | 38°5 iL 64 34 32
Sey aaa | 46 pe apveo! | 19 37 7 60 32 | 380
lin Museum) é
Average of males . | 49°1 | 138°4 6°6. | 32-7 | 16:3 9°5 | 36°4 61 | 58°8 82 30°3 | 222°5
FEMALES.
Female, Yorkshire, mea-
sured by A. Whitaker. | 43 34 227
Female, Cheshire, 29th
June 1897 (dried skin
at British Museum) 50 12 31 9 35°5
Female, do. do. 5 51 13 35 10 35
Female, Hampshire (mea-
sured from spirit at
British Museum) . ote 17 9 38 6 63 34 31°5
reemale, Gordon Castle,
N.-W. Banff, 14th July
1892 : 45°5 |13 6°5 | 37 16°5 | 85 | 36 10°5 ate 50 An 227
*Female, Loch Dochart,
W. Perth, 15th July ®
1894 3 44 13 6°5 | 37 16°75 | 10°5t | 39 a4 a0 Ac 227
Average of females . | 46°7 | 12-7 34°8 9:4 | 86°7 3: St 50
*Female, Yorkshire, one
day old(A. Whitaker) . | 29°25 15°75 we ee 76 82
*Female, Loch Dochart, approx.
W. Perth, young, 15th
July 1894 5 - 29 18°5 9°5 8:5 | 18°5 aa ae 140 |} 101

* Measured by Eagle Clarke in inches, and converted to mm.

+ With claw.

WINGs (4 natural size and diagrammatic) OF

1. Nyctalus noctula. 2. Vespertilio serotinus.
3. Rhinolophus ferrum-equinum.

PLATE IX.

DAUBENTON’S, OR THE ‘WATER BAT 149

known haunt at Christchurch, Hampshire, and who describes
its flight with accuracy, was not aware of its distinctness from
the Pipistrelle, and the same remark is probably true also
of Gilbert White," who once met with “myriads of bats” on
the Thames between Richmond and Sunbury. Fleming, who
obtained it in Fifeshire, and Jenyns, who procured one from
Milton Park, Northamptonshire, supposed that they had before
them the Notch-eared Bat,’ a species said by its describer
to have been obtained near Dover by A. Brongniart, while
an albinic example from Durham was believed to be a new
species* by Jenyns. Yarrell possessed three taken at Islington,
and it was from a study of these, together with others belonging
to Jenyns, that Bell was led to add the present species definitely
to the British list. Yet the Water Bat was for years regarded
as one of the rarest of British Chiroptera, whereas we now
know it to be, in its own peculiar haunts, one of the com-
monest and most widely distributed of them all; indeed, the
late R. F. Tomes wrote that at certain spots on the Avon,
near Stratford, there could not have been fewer than one to
every square yard, and this abundance extended over a very
considerable space.

So peculiar are the vespertinal habits of this species, that
although it is locally abundant, an ordinary observer may be
quite unconscious of its existence. It is essentially aquatic,
if such an expression be applicable to an animal which never
enters the water. It haunts that element continually, flying
so close to it that it is difficult to distinguish between the
creature itself and its reflection. The flight, quivering and
slow, is performed by very slight but rapid strokes of the
wings ; it may, indeed, be said to vibrate, rather than to fly,
over the water. It could not well fly in any other manner so
near the surface without often striking it, and this it seldom,
or perhaps never, does, although it often pauses to dip its
nose into the liquid, whether to drink or to pick up some
floating food, has never been ascertained with certainty. Mr
G. H. Caton Haigh,* however, was on one occasion so for-

1 Letter xi. to Pennant, dated 9th Sept. 1767, original edit., 1789, 32.
2 V. emarginatus of Geoffroy. 3 Vespertilio edilis.
4 Zoologist, 1889, 434.

150 VESPERTILIONIDAA—MYOTIS

tunate as to observe one, after hovering repeatedly over a
particular spot, suddenly drop flat with its wings fully extended.
There it remained motionless for several seconds, and then
rose with apparent ease to continue its flight; the action
appeared to be voluntary, and not the result of accident.

The flight of the Water Bat, as it pursues the tenour of
its untiring way in easy circles, now over mid-water, now close
to the bank, has excited the wondering admiration of many
naturalists, and has been by Macpherson compared with that
of a sandpiper, and by others with that of a swallow or
martin. Its preference for certain areas of streams or lakes
to the exclusion of others has, however, hidden it from many
eyes, and, although it may be abundant in certain haunts, it is
often absent from the surrounding and less favoured localities.
Broadly speaking, although with conspicuous exceptions, it
prefers pools or reaches where the current is slack or the
water stagnant, and where there is ample shelter. One such
resort was known to Tomes, near Alcester, in Worcestershire.
A small stream called the Arrow passes near the mansion of
Coughton Court, and in the grounds is a chain of deep and
melancholy ponds overhung by alders and other moisture-loving
trees. Here Tomes found the bats abounding. Their diurnal
retreat was the roof-chamber of a neighbouring out-building,
where the floor was covered an inch or two thick with the
evidences of the presence of a numerous colony, and where
every crevice overhead was crammed with occupants.

Its peculiar habits occasionally bring this bat under the
notice of anglers, and it is probable that many of the accounts
of the hooking of bats by artificial flies refer to it. Mr Gordon
Dalgleish’ was successful in securing one with a specially
designed bait of tissue- paper, but Mr Symington Grieve
found that at Loch Dochart, although several dashed after
the artificial flies, they discovered the true nature of the lure
in time to save themselves, although he was assured that such
was not always the case.

1 Zoologist, 1904, 345; see also F. Coburn, Journ. cit, 1892, 485, the bats being
caught by the mouth. In some cases the hook catches in a wing, as related of a
Long-eared Bat by C. G. Gray, Field, 28th May 1892, 810, the capture being

probably not accidental, but due to the bat’s having attempted to envelop the bait
with its wings.

DAUBENTON’S, OR THE WATER BAT I51

It does not appear that any study has ever been
attempted of the food of this species, but Mr Robert Service’
believes that it feeds entirely on caddis flies. Judging from
the reluctance of captive specimens to attack large insects,
not excepting the succulent mealworm, Mr Arthur Whitaker
suggests that its ordinary food must be of minute size. Mr
Charles Oldham’s experience is different, since he found that
both mealworms and large moths were accepted without the
slightest reluctance. The struggles of the former were over-
come without any difficulty, the bat being both bigger and
stronger than either the Whiskered or the Pipistrelle. Large
and powerful moths were, however, pouched with such prompt-
ness and efficiency as to suggest that insects of consider-
able size are occasionally at any rate captured under natural
conditions.

During the day the Water Bat resorts indiscriminately to
buildings, trees, or caves, in which its habit is to congregate
in hanging clusters, somewhat like swarms of bees, but where
there are nooks or crannies in its retreat it wedges itself into
them on a system which can only be based upon the utmost
economy of space. These colonies, at least in summer, include
both sexes, as Mr J. G. Millais found to be the case in two
assemblages from which he received specimens in June. Mr
T. A. Coward, too, finds the summer colonies composed of both
sexes, young and old. It should be noted, however, that
five specimens taken by Mr H. Lyster Jameson,’ on 11th
July, in Bohoe Cave, Fermanagh, were all (perhaps only by a
coincidence) males.

The diurnal retreats are often, but not necessarily, situated
in the immediate neighbourhood of water, and it is no doubt
for the convenience thus afforded that this bat loves districts
where the woods grow close to the water’s edge. It does not
object to the companionship of other species, and J. R.
Kinahan* found it in company with the Pipistrelle in crevices
in Tankardstown Bridge, on the river Barrow, Ireland, in
1853. The stones at the entrance were so smooth and

1 Ann. Scott. Nat. Hist., 1896, 201. 2 Trish Naturalist, 1896, 94.
3 Proc. Dublin Nat. Hist. Soc. Dec. 9, 1853, reported in Mat. Hist. Review
(Dublin), i., 1854, 23-25 ; Zoologist, 1853, 4012-4013.

152 VESPERTILIONIDA:—MYOTIS

polished as to indicate a large colony, and the little animals
voiced their protests by a loud outcry, described as chirp-
ing, squeaking, and clicking. It was perfectly possible to dis-
tinguish the two bats, ‘both on the wing and when coming
out of the hole—Daubenton’s coming to its mouth, and thence
flying straight out; whilst the Pipistrelle crept to the edge of
the pier, and letting himself fall from thence, flew off. This,
added to the latter’s being more wary and active in dodging
the net, rendered him more difficult to capture, and to retain
when captured, as out of three, supposed to be of this species
captured, I was only able to retain one. Daubenton’s Bat
does not fly as rapidly nor make such quick turns as the
Pipistrelle, and when struck into the water, floundered in it

so as to enable me to catch him, while the Pipistrelle, under —

similar circumstances, just touched it and was off. The cry
of the Pipistrelle is much shriller than that of the other.
There was also an unmistakably fetid odour from the Pipi-
strelle, which I did not remark from Daubenton’s Bat... .
On confinement, their manners are very different; the
Pipistrelle being impatient, squealing, and biting like a little
fury, and running up and down the sides of the net; while
Daubenton’s Bat was gentle, submitting to be handled, merely
gaping with its mouth, and uttering a soft, low chirp; .. .”?

One of the most picturesque summer breeding colonies
of bats in the British Isles is that described by Mr
Grieve,’ as discovered by him at Glen Dochart, Perthshire.
Here the spurs of Creag Liuragam descend at several points
in precipitous rocks into the waters of Lochs Ure and Dochart,
and in these perpendicular and fissured faces are the chosen
homes of Daubenton’s Bat. So numerous are they that their
presence was in one case detected by the “strange wail” of
their voices, which, rising and falling intermittently, attracted
attention from the opposite side of the loch, about three
hundred yards away.

As is the case with so many other species, the Water
Bat often changes its domicile for the winter, at which season

1 In the quotation, Daubenton’s has throughout been substituted for Natterer’s
Bat, for which species Kinahan mistook it (see p. 145, under “ Distribution ”).
2 Ann. Scott. Nat. Hist., 1894, 193-195.

.
a

PLATE

(Natural size.)

b
<
fea)
a
=
=
fe)
4
|
Ww

DAUBENTON’S, OR THE WATER BAT 153

it seems to prefer the greater security afforded by a cave or
building to the somewhat uncertain protection of the branches
of trees, or of open rock-fissures like those of Loch Dochart,
which were deserted by their occupants in autumn.’ In its
winter retreats, in contradiction to the sociability of summer,
it is not unusually met with hanging singly in complete dark-
ness. Mr Heatley Noble’s cavern at Henley-on-Thames, and
the mine tunnels at Alderley Edge, Cheshire, are said to be
only resorted to in winter.

Possibly the seasonal migration suggested above may at
times reach a somewhat extended scale; otherwise, unless it
had a wish to emulate the performances of the petrels, it is
difficult to account for the capture of a bat of this species, as
related by Mr R. M. Barrington,’ at the Lucifer Shoals Light-
ship, nine miles off the coast of Wexford, on 24th April 1891.

As a rule, this species is supposed to be late—distinctly
later than the Pipistrelle—in making its vespertinal appearance,
and to delay the commencement of its flight until the shades
of night are well advanced. It is further usually assumed
that on emerging it flies straight to the water; but observa-
tions on these points are rarely recorded, since it is difficult
to detect a bat actually issuing from its retreat. The prob-
able truth is that it is an early flier, but that it begins
the evening with some evolutions in mid-air, as has been
independently suggested by Messrs Coward and_ Lyster
Jameson.? The latter naturalist, at Branganstown, Ireland,
and Kinahan, at Tankardstown Bridge, found the Pipistrelle
the later of the two, at Tankardstown Bridge by a full half-
hour. Lastly, Mr Oldham writes me that, although the average
time of appearance at the water in Cheshire is fifty-six minutes
after sunset, he has actually caught four bats as they emerged
from their den thirty-four and twenty-nine minutes earlier.
At the same place, and nearly, although not exactly, at the
same date, he caught a Pipistrelle as it emerged some thirty
minutes after the Water Bats. The point is one of some
interest, and deserves attention.

1 Ann. Scott. Nat. Hist., 1896, 57-58.
2 The Migration of Birds, 284: R. H. Porter, London, 1900.
3 Trish Naturalist, 1897, 39.

154 VESPERTILIONIDA—MYOTIS

The average time of appearance at the water, as computed
by Mr Oldham, for Cheshire, is corroborated for Ireland by Dr
N. H. Alcock,’ who gives it as fifty-four minutes after sunset.
At Tankardstown Bridge, Kinahan states that bats began to
leave their den at 9.30 p.m. in the last week of June-—that is,
at about seventy minutes after sunset. In July mornings,
also in Ireland, Mr C. B. Moffat found them flying frequently
up to forty-four minutes before sunrise, less frequently after
that time, and disappearing altogether six minutes later.

That Daubenton’s Bat continues its flight all night, although
known to Mr Service’ and others, was first demonstrated by
Mr Moffat,? who, by many painstaking vigils, has been able
to detect its flight before sunrise,* as well as after sunset, and
at most of the intervening hours, a feat of no easy accomplish-
ment considering the size of the bat and the nature of its haunts.

In spite of its apparent objection to light, this bat has,
like others, been observed on the wing by daylight, and
William Borrer® states, not, however, from his own know-
ledge, that in the boathouses of Ullswater and Grasmere it
flies throughout the day. Cold or wet nights may cause it
to suspend its flights altogether, although Mr Coward tells
me that it does not object to slight rain. Kinahan found it
returning before bad weather at 10 o'clock; and on a wet
and stormy 25th June, only one bat issued from the strong-
hold, in which, at 10.30 p.m., all was quiet, and there was no
sign of animation.

Very little is known with certainty concerning the length
of the hibernatory sleep, but it appears that roughly speaking
it lasts from the latter part of September to the middle of
April. Mr Oldham finds the bat active in Cheshire up to
23rd September, and in Hertfordshire to 13th October, while
in Ireland it has come under the notice of Mr E. B. Knox, in
County Cavan, on the same day, and of Dr Alcock on 26th
September. Mr J. Steele Elliott gives the earliest date for its
spring appearance at Sutton Coldfield, near Birmingham, as

1 Trish Naturalist, 1899, 31 and 124. 2 Op. Cit, 202:

3 Trish Naturalist, 1905, 106-107.
* Oldham has seen it flying at 2.45 A.M., or seventy-five minutes before sunrise,

on 13th July.
5 Zoologist, 1874, 4128. 6 Trish Naturalist, 1899, 33.

DAUBENTON’S, OR THE WATER BAT Iss

27th April. Mr Oldham tells me that he has seen it on
the wing in Cheshire on the roth, and Mr Barrington’s
Lucifer Shoals specimen was caught on the 21st of the
same month. Mr Moffat finds it abroad in Ireland at least
from 29th March to 29th October, on one occasion with
a temperature so low as 42° Fahrenheit, and, judging from
its northern distribution in continental Europe, the bat ought
to be at least as hardy as the Pipistrelle; in fact, faecal matter
was found by Mr Oldham in the intestines of one taken at
Alderley Edge, Cheshire, on 5th December 1894.

All observers agree that when on the wing this species is
silent, but it uses its voice freely for some little time before
commencing its evening flight, or when annoyed. Mr Caton
Haigh? writes of its notes as “very weak and shrill, some-
times prolonged into a sort of chatter.” Mr Oldfield Thomas
thinks them of particularly high tone, while Mr Whitaker noticed
of a newly born one that its utterances were very soft and
musical, so faint that they were hardly audible at a distance of
a foot.

Daubenton’s Bat, like others of its genus, is not known to
produce more than one young one at a birth, and it were well
to have confirmation of Monsieur Henri Gadeau de Kerville’s
statement, that the number is even rarely two.* The earliest
young are probably born in June, the latest in July. Kinahan
states that one of the females taken at Tankardstown Bridge
in the last week of June contained a large embryo. He
makes no mention of the other females taken at the same
time, so that we are left to suppose that either they were
not examined, or that their young had already been born.
Mr F. Coburn‘ took a large embryo almost ready for birth
from the body of a female taken on 14th June. Mr
Whitaker had a young one born in captivity® on the 19th,°
while Messrs Coward and Oldham, on the 28th, received two
fledglings, as well as an adult female, recently a mother.

1 Journ. Birmingham Nat. Hist. and Philosoph. Soc., Jan. and Feb. 1896, II., 1.5 7.

2 Zoologist, 1887, 293.

3 Faune de la Normandie, i., 149: Paris, 1888. 4 Zoologist, 1892, 403.

5 In an instance observed by Professor R. Collett, the mother hung head down-
wards (2 /iz.).

6 Naturalist, March 1907, 74: unfortunately this bat died almost immediately.

156 VESPERTILIONIDAZ—MYOTIS

Borrer, Mr William Evans, and Mr Grieve refer to recently
born young which they encountered in July, in the last-
named instance on the 4th. That mentioned by Borrer was
clinging to the nipple of its mother, while Mr Grieve’s
description of a young one at first hanging on to its mother’s
back, but afterwards lying rolled up in her left wing, recalls
the known habits of the young of other bats.

Although gentle, fearless, and readily taking food, this
species is delicate, and seldom lives for long in captivity. A
good flight round the room in the evening is, according to
Tomes, necessary for its health, whereas the Pipistrelle and
the Noctule will live in a box without exercise for a long time,
if well supplied with food.

Mr Oldham has remarked that its flight in a room is
wavering, uncertain, and much slower than that of the
Pipistrelle, the tail is but slightly decurved, and the bat alights
frequently on the furniture or pictures. Sooner or later
it settles down to the peculiar motions which are so
characteristic in a state of nature, skimming the surface of
the floor, with typical shuddering or vibratory action, and
threading its way amongst the legs of chairs and tables with
unerring certainty. Mr Oldham liberated some in a_bath-
room in the expectation that they would skim over the water
in the bath, but they did not do so. One fell into the water,
but seemed to be none the worse for the accident, since it
paddled along the whole length of the bath, constantly striking
the water with its wings on its way.

Captive bats of this species readily drink milk or water, and
accept flies, mealworms, and moths, of which Mr Oldham men-
tions several.’ Like other bats, they must be taught to eat
in their novel surroundings, and a fluttering moth appears to
have no meaning for them until it passes close before the face.

As stated above, Mr Oldham finds that the larger moths
are pouched in the orthodox manner, the struggle even
causing the bat to fall right over on its back. Contrary to
Tomes’ statement in Bell, not the slightest use was made of

1 Xylophasia polyodon, Urapteryx sambucata, Triphena pronuba, Mamestra
brassic@, Cidaria populata, and Porthesia aurifiua, the latter accepted with reluctance :
Zoologist, 1899, 472-473.

PLATE XI,

I
SS
I
2 3

HEADS (natural size) OF

1. Myolis mystacinus. 3. Myotis nattereri.
2. Myotis daubentoni. 4. Pipistrellus pipistre/lus.

THE ROUGH-LEGGED WATER BAT 157

foot or carpus as a means of securing a firm hold of the prey,
and it is possible that on this point Tomes, who certainly
did not altogether understand the meaning of the pouching
manceuvre, was mistaken, especially as he states that it was not
easy to make an accurate observation, from the operation being
so much concealed by the body of the animal.

Unlike some other species, Mr Oldham’s bats were par-
ticularly averse to light, running beneath objects on the table
and getting into the shade as much as possible.

Daubenton’s Bat is fortunately one of those the identifica-
tion of which when in its typical haunts is easy. A careful
search for it in suitable localities ought to reveal its presence
in every county of England and Wales, as well as very widely
in Scotland and Ireland. The use of the remarkably large
foot is not understood, and is a question well worthy of further
study. The suggestion that the bats of this group are cave-
dwellers, while the remainder of the genus live mainly in
trees,! appears to be untenable. They may well be called
Water Bats, from their habit of flying over streams, ponds,
and lakes in the peculiar manner of the sole British member of
the group.

[THE ROUGH-LEGGED WATER BAT.
MYOTIS DASYCNEME (Boie).2

The Rough-legged Water Bat was included in the British
list by Sir Harry Johnston and Mr Richard Lydekker, although
excluded by most other naturalists. Its claim to be considered
a British species rests upon the authority of R. F. Tomes,”
who thus identified a specimen taken upon the banks of the
Stour by G. B. Buckton. Buckton‘* himself regarded his
specimen as a variety of Daubenton’s Bat, and alluded to it
under the name emarginatus, a name not infrequently applied
to that bat by British naturalists of the period. It was
probably Buckton’s record which caused Dobson® to include
southern England in the area of distribution of the present

1 Dobson, Catalogue of Chiroptera, 285.

2 F. Boie, Oken’s /szs (Jena), 1825, 1200.

3 Zoologist, 1854, 4361 ; 1856, 4938.

4 Proc. Linnean Soc. (London), 1853, 260 (pub. 1855).
5 Catalogue of Chiroptera, 296.

158 VESPERTILIONIDA-—MYOTIS

species, but no evidence supporting the statement has ever
been printed.

This bat, which is an inhabitant of temperate and boreal
Europe and Asia, at least from the shores of the North Sea to
the Altais, differs from Daubenton’s Water Bat, amongst other
characters, in its larger size, smaller central upper premolar,
more pointed ear, and forward curving tragus. The forearm
measures about 45 millimetres. |

THE WHISKERED BAT.
MYOTIS MYSTACINUS (Kuhl).

1819. VESPERTILIO MYSTACINUS, Heinrich Kuhl, Meuwe Ann. der Wetterauischen
Gesellschaft fiir die gesammte Naturkunde, i., \1., 202; described from Leisler’s
MSS., from Hanau, Germany; Bell (ed. 1); Jenyns; MacGillivray; Blasius ;
Clermont ; Fatio; Bell (ed. 2); Dobson; Harting, Zoologzs#, 1888, 161-166, pl. 2,
and 441, pl. 3; Blanford ; Flower and Lydekker ; Winge.

1821. VESPERTILIO COLARIS, H. R. Schinz, Cuvier’s Das Tihierreich, i. 177;
described from the collared phase (see description below), from Meisner’s MSS.,
from Mont Blanc, Switzerland ; type lost, vzde auct. cit., Europaische Fauna, 17,
1840.

1827. VESPERTILIO SCHINZII, C. L. Brehm, Ovazs (Jena), iii., 27; described from
Renthendorf, Germany.

1833. VESPERTILIO HUMERALIS, L. A. F. Baillon, /ém. de la Soc. Roy. d’Emula-
tion a’ Abbeville, 50; described from the phase with black shoulder-patches (see
description below), from Abbeville, France.

1838. VESPERTILIO EMARGINATUS, William MacGillivray, British Quadrupeds, 96 ;
not V. emarginatus of Geoffroy.

1841. SELYSIUS MYSTACINUS, C. L., Prince Bonaparte, Fauna Jtalica, i., introduc-
tion (3).

1844. VESPERTILIO LATIPENNIS, J. Crespon, Faune Meéridionale, i., 17 ; vide Troues-
sart, Bull. de la Soc. d’Etude des Sci. Nat. de Nimes, 7, i., 35-39, 1879: described
from Nimes, France.

1845. VESPERTILIO BRANDTII, Eduardo Eversmann, Bull. Soc. Impériale des Nats.
de Moscou, xviii., 505, pl. 13, fig. 8, 1845; described from the Samara River, and
province of Kasan, S.-E. Russia,

1856. BRACHYOTUS MYSTACINUS, F. A. Kolenati, Allgemeine deutsche Naturhist.
Zeitung (Dresden), Neue Folge, ii., 131, 174-175; not Vespertilio ciliatus of
Blasius, 1853 = V. emarginatus of Geoffroy.

1862-63. BRACHYOTUS MYSTACINUS, var. NIGRICANS, var. RUFOFUSCUS, and var.
AUREUS, Carl Koch, /ahrbiicher des Vereins fiir Naturkunde im Herzogthum
(Nassau), xvili., 440-448 ; see also Zoologische Garten, xi., 368-9, 1870; described
from Nassau, Germany.

1869. VESPERTILIO MYSTACINUS, var. NIGRICANS, Victor Fatio, Vertébrés de la Suisse,
i., 92; described from Switzerland: see Mottaz, loc. cz¢. infra.

1869. VESPERTILIO LUGUBRIS, auct. et loc. cit.; see Mottaz, Bull. Soc. Zool. (Geneva),
Nov. 15, 1908, 152-153.

THE WHISKERED BAT 159

1870. VESPERTILIO MYSTACINUS, (I) var. NIGRICANS, (2) var. RUFOFUSCUM, (3) var.
AUREUM, Carl Koch, Zoologische Garten, xi., 368-369, 1870; all described from
three coloured plates, from Frankfort, Germany.

1871. VESPERTILIO MYSTACINUS, NIGRO-FUSCUS, L. J. Fitzinger, Sztzungsberichte
der Katserlichen Acad. der Wissenschaften (Vienna), Ixiii., 1, 217; renaming
Brehm’s Vesfertilio schinziz.

1871. VESPERTILIO MYSTACINUS SCHRANKII,! auct, et op. cit., 219.

1871. MYOTIS BRANDTII, auct. et op. czt., 284.

1898. MYOTIS MYSTACINUS, Oldfield Thomas, Zoologist, 100; Collett; Méhely ;
Johnston ; Cabrera; Millais, 99, pl. 12.

1910. MYOTIS (MYOTIS) MYSTACINUS, E.-L. Trouessart, Faune de Mammiféres
@ Europe, 33.

Le Vespertilion Moustac of the French; de Bartfledermaus of the
Germans; but these are book-names only, the bat being probably
unknown to popular observation either in this country cr in continental
Europe.

Distribution :—This bat, or closely allied representatives, ranges
through boreal and transitional Europe and Asia, from sea-level to
at least the summit of the Hartz Mountains (Blasius), and in the Alps
to about 5500 feet (Fatio), from about 65° N. latitude in Skandinavia
(Collett), Finland, and middle Russia to Spain, Syria, and Trans-
caucasia, where it is very common (Satunin); and from Ireland to
Ferghana, the Altais (Kaschtchenko), the Amoor (Schrenck), Pekin
(Dobson) and Sakhalin. J7. szdzgorensts (Horsfield) of Nepal and
Sikkim is, perhaps, sub-specific. The species appears to have no near
representative in America.

M. mystacinus is probably a common species in every part of
England, except portions of the east and possibly the north. But it is
so little known and so often confused with Pzpzstrellus piptstrellus
that the information at our disposal is still very meagre. It
was until recently regarded as rare or unknown in many
localities where it has since been ascertained to be plentiful. In
Yorkshire, for instance, the first record dates from 1882 (Roebuck,
Zoologist, 1882, 147), yet six years later Roebuck described it as
“one of our common species,” about equally common with Plecotus
auritus and Puprstrellus pipistrellus (Journ. cit., 1888, 164-165).
Similarly, Oldham’s Shropshire record of 1890 (/ourn. cit., 1890,
349) was, I believe, the first, but Forrest now writes me that he
has since received many examples from all parts of that shire. Re-
membering these facts, and its occurrence far to the north in Sweden
and Finland, it seems reasonable to suppose that the regular range of
M. mystacinus extends northwards in Britain, at least to the Highlands.

1 A. Wagner (Wiegmann’s Archiv fiir Naturgeschichte, ix., 25, 1843) mentions
V. schrankit as of Koch, but I cannot find the original description.

160 VESPERTILIONID#—MYOTIS

To come to details; in spite of scarcity of information, Harting,
who collected the available data up to 1888 (Zoologtst, 1888, 161-166),
was able to cite occurrences in Somerset, Dorset, Hampshire with
Wight (see also More, /ourn. cit, 1894, 148; Wadham), Sussex, Kent,
Essex, Cambridge, Northampton, Warwick, Worcester, Stafford, and
one or two other counties (see also Cocks, Journ. czt., 1906, 186) to be
more particularly noticed. Millais added Devon, Berkshire, Surrey,
and Middlesex, but without details; Aplin, Oxford (Journ. cit., 1901,
315; 1904, 311); Oldham, Derby (/ourn. czt., 1889, 68-69), and
Lancashire (/ourn. cit., 1890, 349); and Clark Cornwall. Many of
the above are bare records only, the distribution of the bat not
having been worked out; but for a few counties there is more
detailed information, as, for instance, for Cheshire and Derby, where
Coward and Oldham find it widely distributed; Somerset (see
also Zoologist, 1907, 193), Gloucester, and Wiltshire, where Jenyns
(quoted by Harting) found it commoner than P. fzpzstrellus at Bath,
and Charbonnier and Lloyd Morgan report similarly from the dis-
trict around Bristol; Shropshire as noticed above; Yorkshire, where,
as already shown, it is one of the commonest species; and Essex,
where Laver, who has seen it wherever he has looked for it, believes it
to be much more plentiful than is generally supposed. These records,
added to the fact that in Wales it has been taken in Carnarvon
(Oldham, Zoologist, 1896, 255), is probably not uncommon in Denbigh
(Oldham, /ourn. ctt., 1906, 70), is very likely the most abundant
bat of Merioneth (Caton Haigh, /ourn. cit, 1887, 294), and has a
general reputation for being not uncommon in the north of the princi-
pality (Caton Haigh, Journ. czt., 1887, 144), probably indicate a wide
distribution. But the reports from East Anglia are _ strangely
different, since Jenyns, in contrast to his experience at Bath,
comments upon its rarity; and Caton Haigh, Rope, and South-
well have quite failed to find it in Lincoln, Suffolk, or Norfolk.
The bat may clearly be regarded as common in all worked counties
of England and Wales, except those in the east, to at least as far
north as the York and Durham boundary, and the remaining counties
will probably be added to the list in due course; they are Hereford,
Leicester, Rutland, Buckingham (where Cocks, although unable to pro-
duce a specimen, is sure that it occurs), Hertford, Bedford, Huntingdon,
Nottingham, Montgomery, Radnor, Brecknock, Cardigan, Pembroke,
Carmarthen, Glamorgan, Monmouth, Anglesey, and Flint.

The distribution in the north of England may be treated separately.
So far as I know, no records exist for Westmorland or Northumber-
land; Durham has but one, and that not so satisfactory as might be
(see Harting, of. czzt., 165), but Macpherson mentions four for Cumber-
land, including one near the Scottish border.

THE WHISKERED BAT 161

In Scotland this bat has been taken twice—in the first instance
by Hardy, on the road to Pitlochry, about four miles from Ran-
noch, Perth, in June 1874. The specimen, which is in the Owens
College Museum, Manchester, long lay unrecorded, but, happening
to attract the attention of Kelsall, was by him mentioned to Harting
and alluded to by the latter writer (0p. czz, 165), a clue which
enabled W. Evans to work out its entire history (/ammalan Fauna,
Edinburgh District, 23-24). For the second record we are again
indebted to the activity of Evans, who received from George Pow an
example taken at Dunbar, Haddington, on 20th March 1893 (Azz.
Scott. Nat. Hist., 1893, 146). No other Scotch specimens are known,
Millais’ allusion to a third record being an error, so there is as yet
no means of estimating the exact status of this species in North
Britain.

In Ireland the species was first taken by J. R. Kinahan, by whom
one, caught by a cat at Treakle, Co. Clare, in August 1852, was
presented to the Dublin Natural History Society, in February 1853
(Wat. Hist. Review (Dublin), i., 24, 1854; see also, vi., 383, 1859). This
was at first recorded as an example of J/. daudentoni, but the mistake
was corrected at a meeting of the same Society on 12th May 1854
(Wat. Hist. Review (Dublin), i., 148, 1854). In 1897 risk Naturalist, 38-
39) Jameson was able to add the counties of Fermanagh and Louth as
within its habitat, it having been taken in three separate localities
within the former county, and being, in his own experience, prob-
ably not uncommon in the latter. It has since been detected in
Dublin (Welland, Journ. cit, 1898, 272; Alcock, Journ. cit., 1899, 57),
including Lambay Island (Baring, Journ. cit., 1907, 19): Down (Lett,
Proc. Belfast Nat. Field Club, 20th March 1900; R. Patterson, /rish
Naturalist, 1900, 162): Wexford (Moffat, Journ. cét., 1902, 103; Barrett-
Hamilton, Journ. ctt., 1908, 207): and Carlow (Pack Beresford, Journ.
cit, 1906, 16); in most cases more than once in the same county, a
fact which, as in the case of so many other species, seems to indicate
defective human knowledge rather than rarity.

M. mystacinus is found in the Yorkshire hills to a height of 1400 feet
in the Washburn Valley (Storey, MSS.), and in Cheshire it ascends to
heights little inferior in the Longdendale and Goyt Valleys, as I am
informed by Coward and Oldham.

Distribution in time :—See under MV. daudentoni.

The period of gestation is unknown, and the number of young, in
Britain at least, is believed not to exceed one, usually born in June or July.

Description :—The general form and appearance of this bat are
those of its genus, exclusive of the group Leuconve, and it is thus readily
distinguished from the Pipistrelle, than which it is larger. It is smaller
than 17. daubentonz,

162 VESPERTILIONIDAE—MYOTIS

The forehead is elevated, the occiput prominent, the muzzle pointed,
the nostril moderately tumid (Plate XI., Fig. 1, p. 140).

The ear (Fig. 2, No. 6, p. 7), when laid forward, reaches well beyond
the tip of the nose; the outer margin is rather deeply notched, the upper
half being deeply concave, with wavy outline, the lower half abruptly
convex ; a slight emargination opposite the base of the tragus gives rise
to a small but distinct basal lobe; the inner margin is convex through-
out its length to the rounded tip; at its base it bends inwards almost
at right angles to its former course; there are four to six cross-folds.

The tragus (Fig. 2, No. 6, p. 7), which is broadest at a point opposite
about one quarter of the height of the inner margin, is straight, does
not curve outwards, and tapers to a point at slightly more than half the
height of the ear ; the inner margin is straight, the outer margin roughly
triangular, with a small rounded basal lobe surmounted by a concavity.

The wing (Plate XIII., Fig. 1, p. 174) arises from the base of the toes,
and, the third and fifth metacarpals being more nearly of equal length, is
broader than that of WW. daubentont. The relations of the lower leg and
foot are remarkably different, the length of the latter being only about half
that of the former as against nine-sixteenths in JZ. daubentonz. Rela-
tively to the forearm the lower leg is longer than in any other bat already
described, and creates a large interfemoral membrane. The calcar is
small, extends along the free border of the interfemoral only about half-
way from foot to tail, and does not terminate in a noticeable projection.

The fur is long and thick, burying the eyes, covering the face to the
upper lips, and giving it a short, thick appearance ; the lips are fringed
with a moustache of bristles, and there are a few on the chin and a row
across the forehead. On the wing, above and below, the fur extends to
a line connecting the central points of humerus and femur, and on the
interfemoral membrane to the end of the third caudal vertebra above
and to the root of the tail beneath. The ear is very sparsely haired.

The colour is somewhat variable, but the most frequent type is
probably that having the upper surface somewhere between or near
grizzled “raw umber” and “wood brown,” the lower surface lighter and
nearer dirty white. These colours are those of the tips of the hairs, the
bases being everywhere dusky. The line of demarcation is not abrupt,
and runs approximately from the angle of the mouth to and along the
line of attachment of wing and body. The face and chin are often
dusky, in contrast to the brownish forehead and whitish throat, and
there may be conspicuous patches of dusky hairs at the inset of the
arms, a pattern which no doubt suggested the technical names colarzs
and humeralts.

The bare skin of the muzzle, face, nose, lips, ear, and limbs, is silky
black, the wing and tail brownish black ; the nails dusky with paler tips.

The young are darker above and lighter below; but old females,

THE WHISKERED BAT 163

especially after having reared their young, are said to have the upper
side faded or rusty (Bell).

In the skull (Fig. 8, No. 3, p. 101) the profile is saddle-shaped, or like
a shallow S; the frontal region is depressed, and the narrow muzzle
about horizontal. The bullz are small.

The teeth are similar to those of Jd/.
daubentonz, but the outer lower incisors are
not so long in section (Fig. 15).

Individual variation runs frequently ‘ani
towards melanism, and some specimens give
the general impression of being sooty black,
but it is not known whether this phrase is
due to true melanism, or is merely exhibited
at seasons when the brown hair-tips may
have worn away. C. Koch (Zool. Garten, xi.,

1870, 369) and Fatio state that in Germany
and Switzerland this form, which they name
“var. nigricans,’ is an inhabitant of the
mountains; the former distinguishes also
two other forms—a “var. aureum” from FIG. 15.—FRONT VIEW OF In-
south Germany and the plains, and the ©IS08S AND CANINES oF {yous

- mystacinus (enlarged and dia-
typical “var. rufofuscum.” Probably all grammatic).
three occur in Britain.

Koch mentions also an albinic example, a young male; otherwise,
information on the variation of this species is very deficient and
demands attention.

The geographical variation is also, as yet, unknown, but Blanford
states that Nepalese specimens are of large size.

Dimensions in millimetres (see page 164).

Charbonnier and Lloyd Morgan state that newly-born bats of this
species have an expanse of about three inches (75 millimetres).

Proportionate lengths :—Foot (without claws) about -48 of lower
leg ; fifth metacarpal about -96 of third; lower leg about -45 of forearm
and about -34 of head and body.

Skull:—Greatest length, 13-5; basal length in middle line, 10-5 ;
palatal length in middle line, 5-75; from posterior border of 7° to
anterior border of canine, 5; same in lower jaw, 5-5; greatest breadth
at zygoma, 8; posterior breadth, 7; breadth between orbits, 6; breadth
at constriction, 3-75.

Distinguishing characters :—The Whiskered Bat is most frequently
confused with the Pipistrelle, a bat of similar size, from which it differs
in many important respects, notably its thirty-eight, not thirty-four,
teeth, its hairy face, longer than broad, notched ear, and lanceolate
tragus. The forearm, third and fifth metacarpals, and lower leg are

164 VESPERTILIONIDA!X—MYOTIS

all distinctly longer. Relatively to the forearm, the lower leg, foot,
and third metacarpals are longer, but the longest digit is shorter.
Daubenton’s Bat, on the other hand, is distinctly larger than the
Whiskered, and, apart from its shorter, hardly-notched ear, and
shorter tragus, broadest at about its centre, is easily marked out by
the proportions of its large foot and calcar.

DIMENSIONS IN MILLIMETRES :—

a 2 : S
SPE tos 2 |3 Bi |b [OS Sine aes
coe eae ees 2 Eo 8 3 cs) 3 Be ol esacoea ee
Col Ea to = ba es 3 a » i=) =) be OD fers)
EO SP || eee 8 o 2a o s 2 LS s pais ‘oO
sila) AZ = 5 35 5 Ss Sp 5 8 | #F
g/e|es Sir Geel sey Ft ces 3) gate e
Ss | & $ 4 a) 3 g 2 Seliac
Hla] & 2 Bt eal So eae
3 q o
Male (in spirit), Burring-
ton, Cheddar, Jan.1907 | 50 | .. ae a 18 75 37 6°0 | 55 30 | 30
Male, Macclesfield (dry si
skin - British Mu-
seum Ba ON) ate 30 14 50 35 Sond Wee: 30 | 28
Male, Co. Louth, 18th
July 1905 (Dublin
Museum) . Beal eacelllke é i Py 6 82
Male (in spirit), Co. Fer-
managh, llth July
1895 (Dublin Museum) | 42| .. | .. | 31:5] 16 7 83 | 5:25|53°5 | 28°5 | 27
Male, do. (do.) (no date) | .. | ..] .. .. TERS 6 84 | 5°25| 58°25] 28 | 26°25
Female, Burrington,
Cheddar, Jan.1907 ./|..] .. ae ae 16 75 34 5 55 29 PH
Female, Merioneth, 8th
July 1892 OW, E.
Clarke) 4 -| 48) 14] 7:5 85 14 8 33 5'5 se a oF 228 | 5°45
grms.
Average (approximate) | 46 | .. as 55) vb) 184 5:5 | 54 29 | 27-5

Male, young (in spirit),
Burrington, pecader,
Jan. 1907 . aS ah i oe ip 16 T5 34 55 | 54 28 26°75

The Whiskered Bat is probably the commonest British
representative of its genus. It was described by the German
naturalist Leisler early in last century, and was first recorded
as British by J. E. Gray’ in 1826. Gray supposed that
Montagu confused it with the Barbastelle, and stated that
the specimen thus named in the British Museum, which
belonged to Montagu, was really a Whiskered Bat. The bat
was afterwards obtained by Jenyns, Yarrell, and Bell, the latter
of whom figured it in his first edition. In Ireland, J. R.
Kinahan found it in County Clare in 1853, and it has since
been shown to be anything but rare. In fact, in the
west of England and Wales it probably outnumbers the

1 Zoological Journal, ii., 109.

ee ee eS eee

PLATE XII.

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S1Ze.

(Natural

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THE WHISKERED BAT 165

Pipistrelle, the popular ‘“‘Common Bat” of this country gener-
ally. It is hardly known in Scotland.

But although not rare in England, the Whiskered Bat is
little known and seldom seen, a fact which has given it a repu-
tation for scarcity, or, at least, for solitary habits. It might,
indeed, be inferred from the published accounts that it is quite
unusual to find it in company, an inference strengthened by
the fact that, in its winter retreats, each individual hangs, asa
rule, in its own corner or crevice, in marked contrast to the
strongly gregarious character of the Noctule and other species,
which crowd closely together. On the other hand, assemblages
have been described, as by R. F. Tomes,’ who knew of a
colony consisting of more than a hundred, inhabiting the roof
of his house at Littleton, Worcestershire, and by Mr Oxley
Grabham,? who took five from behind a shutter in Yorkshire ;
and it may be that further examination will prove it to be
more sociable than is commonly supposed.

' The flight of this bat so closely resembles that of the
Pipistrelle that the most acute observers often fail to distinguish
the two when on the wing, a fact due rather to their small size,
which prevents accurate observation and comparison, rather
than to any real resemblance. In the confined space of a room
the two are very different, and, apart from the Whiskered Bat
being larger, the slow and steady rhythm of its flight, always
prone to “skim” floor or ceiling, the interfemoral membrane
with a downward curve, is, as in Daubenton’s Bat, very
characteristic.’ If there be other distinctions, they lie, perhaps,
in the general silence of the present species, the Pipistrelle
being very noisy, and its supposed preference for the branches
of trees, from the leaves of which it picks off its prey; in fact
Tomes thought it the most arboreal of all English bats.
According to Mr G. H. Caton Haigh, the Whiskered Bat
comes abroad earlier in the evening than the Pipistrelle, “and
usually selects for its hunting-ground the sheltered ends of a
high hedge or plantation, or even a cliff, along which it
flies to and fro, seldom rising as high as the tops of the
trees or rocks nearest to it. When crossing an open space it

1 Worcestershire, i., 174. 2 Naturalist, 1899, 74.
3 As remarked also by Caton Haigh, Zoologist, 1887, 294.

166 VESPERTILIONIDA—MYOTIS

generally keeps close to the ground.” He has never observed
this species frequenting the open places in woods, of which the
Pipistrelle is very fond. In Worcestershire, Tomes’ has seen
the bats drop out of a hole under the slates of his house,
either singly or in twos or threes. They lost no time in getting
into the top of a large walnut tree, through which they passed,
and scattered off to other trees to feed, passing with quivering
flight through the branches and leaves. Other naturalists
such as Mr Arthur Whitaker, remark on their preference for
the neighbourhood of water, and Mr Charles Oldham has
described his observation” of at least one hundred feeding along
half a mile of the river Dane at Danebridge, near Macclesfield.
“Their flight,” he writes, ‘“was slow, steady, and silent—I
have never heard this species squeak on the wing. Individuals
did not appear to wander far, but confined their attentions to
single pools or short stretches of the stream, where they flitted
about the alder bushes or threaded their way with marvellous
precision through the lower branches of the sycamore trees.
I never saw one rise to a greater height than twenty feet,
and often they flew within a few inches of the ground, or
skimmed the surface of a pool for a yard or two, only to rise
again and resume their flight around the alders. Even when
close to the surface of the river their flight could never be
mistaken for the continuous flight at the same level, just above
the surface, of the narrow-winged” Water Bat. That this
bat may sometimes wander to comparatively exposed situations,
is shown by its occurrence—the second time on record for
Scotland—on the links east of Dunbar, by the shores of the
North Sea, and again on Lambay Island, off the east coast of
Ireland. Like that of the Pipistrelle, its flight is often so
lowly as to lead to its destruction by means of sticks and caps.
One which came under the notice of William Borrer* had
flown in broad daylight against a man’s white frock, white
being particularly attractive to bats. Another, which came
into the hands of Mr Grabham,* had struck against a
policeman’s helmet in the streets of York, while Mr Oldham’®
found one hanging in a moribund condition impaled upon a

1 Worcestershire, \., 174. 2 Naturalist, 1897, 242. 3 Zoologist, 1874, 4128.
4 Naturalist, 1899, 74. 5 Zoologist, 1899, 475.

THE WHISKERED BAT 167

briar, a thorn of which, despite its usual agility, it had evidently
failed to avoid.

During its season of activity the Whiskered Bat chooses
varied situations for its diurnal resting-place, and, as a rule,
sleeps apart from its fellows. Holes in walls, in roofs of houses
and buildings, and the spaces behind shutters and sign-boards,
are its customary home. Less frequently, it resorts to some
convenient hole or crevice in a tree, or creeps behind a loose
piece of bark. Among more exceptional retreats may be
mentioned the ivy on a wall,’ an auger-hole in an old gate-
post,” and a crevice between two boulders in a wood;* one
has even been found asleep in bright sunlight on the top of
a stone wall.* During the summer months it rarely resorts
to caverns and similar situations, although Mr Oldham has
taken one near the entrance to an old mine-tunnel in June,’
but in the winter such places are favourite hibernacula.
Messrs T. A. Coward and Oldham have frequently found it
at that season in disused copper-mines at Alderley Edge,
Cheshire, and in old lead workings in the Derbyshire dales,
places which are deserted in spring. Of Alderley they
write that: ‘The red sandstone rock is pierced in many
places by horizontal tunnels about six feet high and as much
in width, From November to March these tunnels are
resorted to by Whiskered, Long-eared, and, more rarely,
Daubenton’s Bats, which hang suspended by their feet from
the roof and walls. We have found all three species in the
same tunnel, but they cannot be said to associate with one
another, and the individual bats are, with the rare exception of
the Long-eared, always solitary. At times a bat may be found
near the mouth of a tunnel where there is sufficient light to see
it without a candle, but as a rule they retire to the deeper
recesses, sometimes more than a hundred yards from the tunnel
mouth, where they are in total darkness. For the most part
the tunnels are dry, but the bats sometimes hang in damp
places, and their fur then glistens with beads of water. It is
possible that they feed during their retirement, for, although

1 Zoologist, 1874, 4128. 2 Tomes in Bell.
3 N. H. Alcock, /rish Naturalist, 1898, 272.
4 W. D. Roebuck, Waturalist, 1886, 113. 5 Zoologist, 1896, 255.

a
4
‘

168 \. VESPERTILIONIDA—MYOTIS

we have never seen one on the wing, we have often found food
in the alimentary.canal on dissection, and, indeed, food in
abundance is close at hand, as the tunnels are resorted to in
winter by a spider, two species of Moth . . .,) and myriads of
flies.2 No bats are to be found in these tunnels during the
warmer months of the year.” In Mr Heatley Noble's big
artificial cave near Henley-on-Thames, Mr A. H. Cocks and
Dr E. A. Wilson® found seven on 14th February, all near one
of the entrances, but in summer this cave is deserted by all bats.

The Whiskered Bat is frequently seen abroad during the
day, probably more so than any other British species, and it is
also one of those which appear on fine days—at least in the
south—throughout the winter. Mr ©. B. Moffat records it as
flying at a temperature of 48° Fahrenheit on 21st January,
in Ireland; Borrer‘* received one shot near Dover in the mild
January of 1853; Mr William Evans’ Dunbar specimen was
taken on 20th March, and Mr T. W. Proger sends me word
that he has observed it abroad in south Wales on 21st December
1903, and in Wiltshire at the end of November of the same
year. But the too frequent confusion of this species with the
Pipistrelle makes a series of reliable observations impossible.
Little is known about the duration of the nightly flight.
Probably, however, it begins early in the evening, Mr Oldham
having noticed one on the wing at 7.30 p.m. in broad daylight
in the latter end of May:° it may also last all night, since I
caught one in my house between one and two o'clock on the
morning of 26th July.

Only the most vague information is available about the breed-
ing habits of this bat. Tomes’ statement that its single young
—the number verified by dissection—is brought forth about
the end of June or in July, corroborates, as regards the number
at least, the experience of Blasius, for Germany. Messrs H. J.
Charbonnier and C. Lloyd Morgan® found that of some
hundreds obtained at Willsbridge, near Kevnsham, Bristol,

1 Gonoptera libatrix and Scotosia dubttata.

2 Of which the most abundant are Blepharoptera serrata, Borborus niger, and a
species of Culex.

3 Zoologist, 1906, 186; also, Wilson, in lit. 4 Zoologist, 1874, 4128.

5 Naturalist, 1897, 242. 6 Charbonnier, see Somerset, i., 164.

— ee

THE WHISKERED BAT 169

on 2nd July, twenty or more examined were all females.
Several had young ones clinging to their fur, and others
became mothers the next day, the newly born bats being
of the usual naked appearance, with dusky heads and wings.
An immature male (now preserved in the Dublin Museum) was
found clinging to a house by Mr W. Garstin on 18th July, in
County Louth, Ireland; another taken at Kilmanock, County
Wexford, on 26th July, although immature, had attained the
usual size of an adult: both seem to show that the date of birth
may be much earlier in Ireland. Allowing seven weeks for
gestation and as many more for growth of the young, it may be
safe to calculate that the mothers of these two young bats became
alert enough to ovulate in the middle of April, and gave birth
to their young at the end of May or in the beginning of June.

Mr O. V. Aplin* suggests that the Whiskered Bat feeds
largely on moths, but the only definite information available on
this subject is the statement of Professor G. H. Carpenter,?
that the fragments of insects in the excreta of an Irish specimen
seemed all referable to the two-winged flies.» Mr Oldham found
a small staphylinid beetle in the mouth of one which he captured.*

In captivity, this species has been the subject of study
by Dr Alcock® and Mr Oldham,°® the latter of whom has
written a detailed account of its habits and demeanour. It
appears that, although naturally of fierce temperament when
first taken, it may be readily tamed, and when the difficulties
at first attending its feeding have been overcome, it displays a
marvellous appetite, swallowing with eagerness moths,’ spiders,
and raw rabbit’s liver, and lapping up milk or water even more
readily. Dr Alcock’s specimen also partook of fish, while Mr
Oldham’s executed an elaborate toilet—a habit common to
other species—but, true to the character of its race, it was the
host of external parasites, a tick and two fleas having been
caught upon its person by its captor.

“Tt evinced,” writes Mr Oldham, ‘“‘little disposition for

1 Zoologist, 1901, 315 3 1904, 311. 2 Trish Naturalist, 1902, 103.
3 Diptera, and that some pieces of legs and wings clearly belonged to a small
Tipulid—very possibly a species of 77ichocera.
4 Naturalist, 1897, 242. ® Irish Naturalist, 1899, 56.
° Zoologist, 1899, 49-53. ? Scotosia dubitata and Gonoptera libatrix.
Q

170 VESPERTILIONIDA—MYOTIS

flight, especially after feeding, and if compelled to take wing
would, after one or two turns round the room, drop on to the
floor, or pitch on a curtain, chair, or my head or body... .
Although loth to fly, it seemed never tired of running about
among the papers and other objects on the table, and was
seldom stationary unless it was eating. The bell-jar in which
I kept it was raised above a stand on supports rather more
than 4 in., or, to be exact, just 7 mm. in height, and whenever
the perforated zinc guard was removed from the intervening
space the Bat would creep out at once. The bright light of the
lamp on my table seemed to cause it no inconvenience, for it used
to sit, supported on feet and wrists, eating mealworms within a
few inches of the flame, and never showed any desire to retire to
dark or shaded places. Sometimes it would creep under my
hand, or up my sleeve, but this, I think, was on account of the
sensation of warmth it experienced in nestling against my skin.

‘“The sense of sight seems to be but feeble in the Whiskered
Bat. The example under notice could not see, or at all events
recognise, a mealworm or wet paint-brush, if more than an inch
from its face. As this species is more diurnal than any other
British bat, and may frequently be seen abroad at midday in
summer, the inability of my captive to see objects an inch
away cannot be attributed to the dazzling effects of too strong
a light, especially as this inability existed equally in the daytime
and in the artificial light of alamp. Its hearing also appeared
to be dull, as it never showed by any movement of its head
that it perceived a sudden noise, such as the snapping of my
fingers, or the click of a watch-lid being closed. It sometimes
slept prone upon the floor with wings folded and pressed
closely to its sides, at other times suspended by its toes to the
rim of a wooden box. During sleep, which was always pro-
found, its temperature fell considerably, and it felt, as all Bats
do in this state, extremely cold. It usually wakened in the
evening, but exceptionally in the daytime without being roused ;
while, as a rule, it was necessary to warm it into activity by
holding it for a minute or two in my hand if I wanted to feed
it by daylight.”

Its voice, often used, is described by Mr Oldham as a feeble
squeak, less shrill than that of the Long-eared Bat, and by

THE WHISKERED BAT 171

Dr Alcock as lower in pitch than that of other species. Frank
Norton’ likened it to the clicking of a cogwheel and chain.

It was in this species that Mr Oldham first observed the
habit of pouching the prey’ already alluded to in the account
of the Pipistrelle (see Plate VIII.), and his remarks thereon
may fittingly conclude this article. ‘My captive,” he writes,
“used to tuck its head away under its body directly it had seized
an insect, at the same time bringing its feet forward, so far indeed
that it sometimes lost its balance and toppled over on its back.
This habit, practised from the very first, was evidently one of old
standing, and not a trick acquired in confinement. By feeding
the Bat on a sheet of glass so that I could see it from beneath,
or, better still, by giving it an insect as it hung suspended by
its toes, the reason of its action was at once apparent. The
tail being directed forward beneath the body, the interfemoral
membrane formed a pouch into which the Bat thrust its head,
and was thereby enabled to get a firmer grip of its prey without
any danger of dropping it. When the Bat was on a flat
surface the lower side of this pouch was pressed closer to its
belly than would be the case during flight, so that it sometimes
failed to get its head into the pouch, and let a mealworm drop.
When this was the case it never made any attempt to seize its
prey again, and the mealworm would escape by crawling out
from beneath its wings or tail. When the Bat was suspended,
however, the bag was wide open, and the insect never escaped.
Experience seemed to teach it that the mealworms were
incapable of escape by flight, and latterly it did not always
thrust its head into the interfemoral pouch after seizing one,
but devoured it without this preliminary. Ina free state, Bats
capturing the greater part, if not all, of their food on the wing,
must often fail to grip large insects securely at the first bite,
and it would be a manifest advantage to have some means of
adjusting their hold without alighting. An insect accidentally
dropped during flight could hardly be recovered, and would
probably be abandoned without further thought, as was the

1 Midland Naturalist, 1883, 151.

2 Zoologist, 1899, 51-53. Norton (doc. cz¢., 152) had previously noticed that it could
be caught by the wing with a rod and line baited with a fly or moth, the wing, as
he thought, being used for striking at the bait.

72 VESPERTILIONIDA—MYOTIS

case when my Whiskered Bat dropped a mealworm. A Long-
eared Bat which I kept for a few days invariably thrust its head
into the interfemoral pouch on seizing a moth. Both Long-
eared and Whiskered Bats have the tail curved beneath them
during flight, although they are usually figured with it held
straight behind them ; and I have little doubt that when on the
wing they actually use the method I have described for securing
their prey. Further observation will probably show that this
curious habit is common to all our British species, with the
possible exception of the Horseshoe Bats, in which the inter-
femoral membrane is comparatively small, and the tail, during
repose at any rate, is carried in a very different way.

“Having firmly secured its prey, whether moth or meal-
worm, by the head or tail, my Whiskered Bat used to swallow
it lengthwise, crunching it thoroughly by rapid movements of
the jaws as it slowly disappeared. Neither foot nor carpus was
ever used in any way to assist it in capturing or holding an
insect. The use of either would, of course, be quite impossible
during flight. Moths and spiders moving near it were pounced
upon and captured, but mealworms disassociated from my
fingers seemed to puzzle it, and only once did I see it capture
one itself, although the creatures frequently crawled just before
its eyes and over its wings and feet. The wings and legs of
moths were always dropped, but once or twice a wing accident-
ally encountered in the Bat’s ramble about the table was
picked up and eaten. The mealworms were, as a rule, entirely
consumed, but sometimes the horny heads were left.”

BECHSTEIN’S BAT.
MYOTIS BECHSTEINI (Kuhl).

1818. VESPERTILIO BECHSTEINII, Heinrich Kuhl, Meue Ann. der Wetterauischen
Gesellschaft fiir die gesammte Naturkunde, \., i., 30, pl. 22 ; described from Hanau,
Germany, from Leisler’s MSS.; Jenyns; Bell (ed. 1); MacGillivray; Blasius ;
Clermont; Fatio; Bell (ed. 2); Dobson; Flower and Lydekker; Lydekker ;
E. T. Newton, Quart. Journ. Geol. Soc. (London), August 1899, 420; Winge.

1829. NYSTACTES BECHSTEINI, Jakob Kaup, System der Europidischen Thierwelt,
i., 106, 108-9.

1842. MYOTIS BECHSTEINII, J. E. Gray, Ann. and Mag. Nat. Hist. Dec., 258;
Fitzinger ; Thomas, Zoologist, 1898, 100; Méhely ; Johnston; Millais. .

1856. MYOTUS BECHSTEINII, F. A. Kolenati, Algemeine deutsche Naturhist. Zeitung
(Dresden), Neue Folge, ii., 131 and 179.

.
|

aren

' r
ral a

PLATE XIill.

WINGs (4 natural size and diagrammatic) OF

1. Myotis mystacinus. 2. M. bechsteini. 3. MW. natiereri.

BECHSTEIN’S BAT 173

1902. VESPERTILIO BECHSTEINII GHIDINII, Victor Fatio, Rev. Suisse de Zool., 10,
400; see Mottaz, Bull. Soc. Zool. (Geneva), Nov. 15, 1908, 160; described from
Gerso, near Lugano, Tessin, Switzerland.

1906. MYOTIS BECHSTEINI FAVONICUS, Oldfield Thomas, Aum. and Mag. Nat. Hist.,
Sept., 220; described from La Granja, Sierra de Guadarrama, Central Spain.

Ig1o. MYOTIS (MYOTIS) BECHSTEINI, E.-L. Trouessart, Faune des Mammiféres
ad’ Europe, 30.

Distribution :—The exact range of Bechstein’s Bat is little known,
but it certainly occurs from Sweden to central Spain and from England
to Hungary. In Britain it is only known from England, its occur-
rences wherein are detailed below.

Distribution in time:—Remains of bats from the Pleistocene
deposits of Ightham Fissure, Kent, were assigned by E. T. Newton to
this species, or, possibly, to JZ. murinus = myosotis (Quart. Journ. Geol.
Soc, August 1899, 420).

The period of gestation is unknown; the number of young is
probably one only, born in midsummer (Blasius; C. L. Brehm).

Description :—Bechstein’s is the largest British member of its genus.

The ear is long and unnotched (Fig. 2, No. 7, p. 7). When laid for-
ward, it stretches beyond the tip of the nose by about half its length ; the
outer margin is very slightly flattened beneath the tip, then convex to
a point very nearly opposite the base of the tragus, then slightly
emarginate and terminating in a small convex lobe; the inner margin
is convex for the lower two-thirds of its length, and then straight or
slightly concave to a rounded point: there are nine or ten cross-folds.

The tragus (Fig. 2, No. 7, p. 7) attains its greatest breadth at a point
close to its base; it is curved outwards above, and tapers to an acute
point at about opposite to the middle part of the ear; the inner margin
is straight below, slightly convex above; the outer margin is quite
distinctly concave above, markedly convex below, terminating after an
emargination in a small triangular basal lobe.

The wing (Plate XIII., Fig. 2) arises from the base of the toes.
In the two New Forest specimens the third and fifth metacarpals are
about of equal length, but in those from the Isle of Wight the third is
slightly the longer of the two. The thumb is long; the area of wing
between digits two and three is large; the tail-tip is free of the inter-
femoral membrane. The most conspicuous features are the exceptionally
long lower leg, but comparatively short longest digit. With a forearm
about equal in length to that of Myctalus lezsleri, the lower leg is but
little shorter than that of Vespertzlio serotinus, but the longest digit
and expanse are far short of those of either, while the third and fifth
metacarpals are about equal to those of 4. darbastellus.

As compared with the forearm, the lower leg is only equalled or
exceeded in Plecotus or Barbastella. The third metacarpal and longest
digit are the shortest amongst British vespertilionid bats.

174 VESPERTILIONIDA—MYOTIS

The wing is therefore of expanse under the average, broad at its
base, with an ample interfemoral membrane. It is poorly developed

F1G. 16,—SIDE VIEw (diagrammatic and enlarged)
OF SKULLS AND TEETH OF

1. Nyctalus leisleri. 2. Myotis bechsteint.
3. WZ, natterer?.

at the extremity. The thumb
is long.

The fur is soft and woolly.
The face (Plate XIV., Fig. 1)
is thinly haired except on the
forehead, and there are a few
moustache -like hairs on the

nc
/

FIG. 17,—FRONT VIEW OF INCISORS
AND CANINES OF Myotis bechsteini (en-
larged and diagrammatic).

muzzle. The wing is compara-
tively free of hairs, the hair of
the body not extending farther
than about to a line joining
the middle points of humerus
and femur, and very slightly to
the interfemoral membrane.
The ear is free of hair except
externally at the base and a
fringe on the lower third of the
outer margin.

The colour is very similar to that of J/. xattereri, the bases of the
hairs being everywhere dusky, with the tips some shade of wood-brown

BECHSTEIN’S BAT 175

above, whitish beneath. The moderately distinct line of demarcation
passes along the line of junction between wing and body and thence
to the base of the ear.

The skull, as compared with that of JZ. nattereri, is larger, but
narrower ; it carries more prominently developed cranial ridges and
much bigger bullz (Fig. 16, No. 2, p. 174).

The teeth are similar to those of J/. mystacinus, but the tip of the
middle upper premolar reaches far beyond the cingula of the neighbour-
ing teeth (Figs. 16, No. 2, and 17, p. 174).

The geographical variation of this rare bat is almost an unknown
quantity, but Thomas has rightly separated central Spanish examples
from Hungarian as the sub-species favonzcus on the ground of their
markedly smaller ears and bulle. I cannot distinguish the few British
specimens available from the type of /avonzcus, but await further
material before identifying the British Bechstein’s Bat with that form.
Both are distinctly smaller than the Hungarian.

DIMENSIONS IN MILLIMETRES :—

Z| oo 3
i tp a 5 3 s : e oD
=
cB alk gl) ~ |S | | ates tiple! | eee
a ms ae 2 See FI SSeo'|| ce 3 3 SS
“ Bivens || fs Bi Sie Wc || eisai. B= ay | bo
PS feel S bE rear Bias: syle eae
ae ae ee 6 (ee jae [ao | ot Sy soe
iy & qa ~~ me) oO
g op $ Se ee | 3 a 2 |s
td En Eb & a A = =| o
3 i = bs
| o
ee rd = of Biase NP icio® ie 35 |20 |10-75| 48 | 7 | 64 | 36 | 845] 248
from old skins i 4 18°5 | 9 35 19 10 42 tf 64 33 33 252
Male, Henley-on-Thames,
10th March 1901 (J. G.
Millais, British Museum),
from dried skin . : te 18 e ae 19 10 39 Ui 59 80°5 | 31
Male, Isle of Wight, 14th
Aug. 1909(P. Wadham) | [52] |] 19 | 8 w= | 20 < |) 490M) Weel ey) le s5iy eee
Female, do. 31st July
1909 c : 2 : [52] | 19 8 34 20 85 39 7+ 65 85 34
Average . C : An 18°7 | 8:7 85 19°5 2 41 7 64 83°5 | 83 [250]

Proportionate lengths:—Foot, without claws, about -53 of lower
leg; fifth metacarpal about equal to third; lower leg about -46 of
forearm.

Skull:—Greatest length, 17; basal length in middle line, 12-25;
palatal length in middle line, 7-5; from posterior border of 7° to
anterior border of canine, 6-5 ; same in lower jaw, 7-25; greatest breadth
at zygoma, 9; posterior breadth, 8; breadth between orbits, 7; breadth
at constriction, 4-5.

Distinguishing characters :—The large ear of Bechstein’s Bat marks
it off very distinctly from every other British species except the Long-

eared, in which the ears meet at their anterior margins. In other

176 VESPERTILIONIDAQ—MYOTIS

respects it approaches nearer to Natterer’s, but the ciliated posterior
margin of the interfemoral membrane and the relatively longer tragus
of the latter are diagnostic. Its closest ally is, perhaps, JZ. myosotzs of
continental Europe, a much larger bat, with relatively shorter ear,
cigar- not sickle-shaped tragus, and wing arising slightly above the
base of the toes.

This handsome and striking species is probably the
rarest of undoubtedly British bats, and was for long known in
this country solely from a specimen now in the British Museum,
taken in the New Forest before 1837 by Millard. Its occur-
rence in this locality was verified by Mr E. W. H. Blagg,
who in a somewhat laconic note,’ written two years after the
event, described his good fortune in having in July 1886
discovered a colony of about a dozen in a hole made by a
woodpecker. Of these he secured two, and has since presented
them to the British Museum. Another, an old male, was shot
by Mr W. C. Ruskin Butterfield near Normanhurst, Battle,
Sussex, on 28th July 1896.2. This bat was examined by
the late Sir William Flower, and was lent to the Corpora-
tion Museum at Hastings in 1898. Unfortunately it has
been mislaid, and is not at the present time forthcoming; Mr
Butterfield writes me that he fears that it has been acciden-
tally destroyed. Then, in March rgor, Mr J. G. Millais found
an adult male in Mr Heatley Noble’s chalk cavern near
Henley-on-Thames.? Lastly, on 31st July 1909, Mr Percy
Wadham captured an adult male near Newport, in the Isle
of Wight.t It was sent to Mr H. G. Jeffery for preservation,
and was identified by that gentleman and by him sent to the
British Museum for verification, with a female taken by Mr
Wadham at the same place on 14th August. I am indebted
to the owner and to Mr Jeffery for the opportunity of examin-
ing these two bats, which have been set up by Mr Jeffery for
Mr Wadham’s collection. Other British records have proved
to be erroneous, viz., that of two examples taken at Preston,
near Brighton, and identified with this species by Frederick

1 Zoologist, 1888, 260. 2 SUSSEF, 11.5, BOM.

3 In Berkshire, not Oxfordshire (see A. H. Cocks, Zoologist, 1910, 74).

4 Recorded by J. E. Harting, /ze/d, 20th Nov. 1909, 889; and by J. E. Kelsall,
Zoologist, 1910, 30.

BECHSTEIN’S BAT 077

Bond, but later assigned by Messrs Millais and J. E. Harting
to Natterer’s Bat;* and that of a specimen in the Oxford
Museum, taken at Godstow, in Berkshire, by Frank Norton,
which was mentioned by H. A. Macpherson as Bechstein’s,?
but referred by the Rev. J. E. Kelsall® to Natterer’s Bat.

The habits of Bechstein’s Bat have never been studied in
detail. Blasius writes that in Germany its haunts are woods
or large orchards and the neighbourhood of buildings, where
it lives in small, dry holes in trees. It commences its flight late
in the evening, flying slowly and at a low elevation over lanes
and forest roadways, and may be recognised while on the
wing by its long ears. It is said not to appear until late in the
spring, to fly only in fine, calm weather, and not to venture
abroad in winter. In Germany, Blasius on two occasions
found one young one with its mother, and, about eighty years
ago, on 9th June, C. L. Brehm* took a party of twenty-
two females, of which seventeen proved to be pregnant with
young almost ready for birth. Kuhl also found as many as
thirteen females together. The occurrence of so many
examples of one sex together in the breeding season suggests
that in this respect the habits of this bat may resemble those
of the Noctule. Mr Millais’ example was lying in a crevice
close to a couple of Natterer’s Bats. It was wide awake, and
when handled remonstrated by biting and uttering a series of
querulous screeches, not unlike the cries of a very young
child. Mr Wadham’s bats were taken at dusk near a withy
bed and small group of trees, from which the Lukely runs for
about a hundred yards by open meadows to a large mill pond
and reed bed.

The long unnotched ears of this bat differentiate it
absolutely from all the other British members of its genus.
Natterer’s Bat approaches it most nearly, but the relatively
longer tragus, slightly notched ears, and ciliated posterior
margin of the interfemoral membrane in that species are
quite distinctive.

1 Harting, Zoologist, 1887, 162; Millais, British Mammals, 1., 98.
* Midland Naturalist, 1883, 153. 3 Zoologist, 1884, 483 ; 1885, 146.
4 Ornis (Jena), 1827, iii, No. 3, 20.

178 VESPERTILIONIDAI—MYOTIS

NATTERER’S BAT.
MYOTIS NATTERERI (Kuhl).

1818. VESPERTILIO NATTERERI, Heinrich Kuhl, Meue Ann. der Wetterautschen
Gesellschaft fiir die gesammte Naturkunde, \., 1, 33, pl. xxiii. ; described from Hanau,
Germany ; Jenyns; Bell (ed. 1) ; MacGillivray ; Blasius ; Clermont; Fatio; Bell
(ed. 2); Flower and Lydekker; E. T. Newton, Quart. Journ. Geol. Soc. (London),
May 1894, 192 ; Lydekker ; Winge.

1842. MYOTIS NATTERERI, J. E. Gray, Aun. and Mag., Nat. Hist., Dec., 258;
Fitzinger ; Thomas, Zoologist, 1898, 100; Johnston ; Méhely; Millais.

1856. ISOTUS NATTERERII, F. A. Kolenati, Algemeine deutsche, Naturhist. Zeitung
(Dresden), Neue Folge, 11., 131 and 177.

1862-63. ISOTUS NATTERERI, var. TYPUS, and var. SPELAEUS, Carl Koch, Jahrbiicher
des Vereins fiir Naturkunde im Herzogthum (Nassau), xviil., 426 and 430; described
from Nassau, Germany.

1910. MYOTIS (MYOTIS) NATTERERI, E.-L. Trouessart, Faune des Mammif eres
a’ Europe, 29.

Reddish-grey Bat of Bell and others, but there are no local names.

Distribution :—Natterer’s Bat ranges through boreal and temperate
Europe and Asia from southern Sweden to Seville, Spain, and Arezzo,
Tuscany (specimens in British Museum), and from Ireland to Kiushiu,
Japan, where the representative sub-species is JZ. x. bombinus of Thomas
(Proc. Zool. Soc.(London), 28th Nov., 1905, 337). It ascends to 2300 feet
in the Caucasus (Satunin).

M. thysanodes of Miller (Vorth American Fauna, 13, 1897, 80-85),
of the lower Sonoran zone from near the southern border of the
western United States to San Luis, Potosi and Michoacan, is probably
the representative in America.

Thanks to Harting (Zoologist, 1889, 245-47), the distribution of WZ.
natterert in the British Isles is now fairly well known. In England it is
found from Cornwall and the Isle of Wight to Durham in the north and
to Norfolk in the east. When Harting wrote there were fourteen
counties within the above area in which it had not been found. Five
only of these now remain, absolutely without records, viz., Wiltshire,
Hereford, Buckingham, Hertford, and Rutland. It has twice been
taken close to the Buckingham border (see Cocks; also Steele Elliott,
Zoologist, 1903, 349), and our ignorance of it in this as in the other
counties is due almost certainly to want of observation.

In Wales it probably occurs in every county, having been recorded
by William Thompson (Proc. Zool. Soc. (London), 13th June 1837,
52), from Harlech Castle, Merioneth, in the extreme west, a locality in
which J. Backhouse, jun., still finds it (Forrest, MSS., and Zoologzst, 1898,
493, and specimen in British Museum); and in 1903 from St David’s,
Pembroke (Forrest). In 1905 Coward and Oldham identified one while
flying in broad daylight near the Cefn cave, St Asaph, Denbigh; in the

NATTERER’S BAT 179

same year Mills examined one shot in Pembroke, while in 1907 Oldham
saw in the possession of Owen two live ones taken in Cardigan (all in lit.).

Within these limits it is probably numerous, although no doubt
overlooked, in most, if not all of the wooded localities, being reported,
for instance, as plentiful at Stainborough, Yorkshire (Armitage,
Naturalist, 1905, 37), and as one of the commonest species at Colchester
(Laver). It has even been taken in Thayer Street, Manchester Square,
London (Harting, Zoologzst, 1888, 25).

In the extreme north of England it has never been reported from
Westmorland or Northumberland, but the occurrence of a numerous
colony at Castletown, near Carlisle, Cumberland, probably indicates its
abundance right up to the Scottish border (Macpherson),

From Scotland two specimens only are recorded, neither of which,
unfortunately, is regarded by Scottish naturalists as absolutely satis-
factory. The first of these,an immature male in the British Museum
(mo¢ an adult female as in Dobson’s Catalogue of Chiroptera, 308), is
registered as from Inveraray, Argyll, August 1858; it had escaped the
recollection of the (supposed) donor, the late Duke of Argyll, but there
seems to be no valid reason for doubting its authenticity! The second,
a roughly preserved skin, unfortunately without a label, from the collec-
tion of the late Robert Gray, is now in the possession of W. Evans: it
may with much probability be connected with Gray’s statement to
Harvie-Brown that he had found this bat “in some plenty” near
Dalkeith, Midlothian, “in dozens in the hole of a tree,” this was in
1880 (W. Evans, Proc. Roy. Phys. Soc, Edinburgh, xvi., No. 8, 388,
27th November 1905). The whole matter has been fully discussed by
W. Evans (Vammalian Fauna of Edinburgh, 22-23, and Ann. Scott.
Nat. Hist., 1901, 129-131), and at any rate there would seem to be no
inherent improbability in the occurrence of this species in Scotland.
Coward, it may be mentioned, has a specimen in alcohol, which he
believes he received alive from Montrose in 1895; but an undoubted
error on the label of the bottle prevents him from making a definite
statement.

In Ireland the species was first made known from an example
procured by Mangan at the Scalp, Enniskerry, almost on the Dublin
border of County Wicklow, in 1845, and presented through M‘Coy to
the Dublin Natural History Society (Proc, 12th February 1845 ;

J. R. Kinahan, Proc., 9th December 1853, published in Dublin Vaz.
Betdis?, Review, i., 23, 1854; see also ii, 8, 1855, also vi, 383, 18590;
M‘Coy, Ann. Mag. Nat. Hist, April 1845, 270). Dobson had a
specimen, still in the British Museum, and dated 1876, from Co, Long-
ford, and the bat has since been detected in Donegal, Fermanagh,

' | have searched in the Duke’s correspondence at the Museum, but can find no
reference to this specimen ; the entry in the register is, however, quite clear.

180 VESPERTILIONIDAZ—MYOTIS

Louth, Longford (Jameson, /7ish Naturalist, 1897, 39), Cork (Folliott
Darling, Zoologist, 1883, 294), Galway (Kane, Jrish Naturalist, 1894,
116, and 1897, 88; also a specimen from Woodpark, in the British
Museum), Antrim (Patterson, /ourn. cit, 1900, 274), Carlow (Pack-
Beresford, Journ. cit., 1905, 228), a combination of counties indicat-
ing a wide distribution throughout the island. (Jameson suggests
that Kinahan’s error in identifying the Tankardstown Bridge, Co.
Kildare, colony of J/. daubentoni with this species may have been
founded upon an actual mixture of the two—see Journ, cit., 1897, 40.)

Distribution in time:—Of very numerous remains of bats found
by Lewis Abbott in the Pleistocene deposits of Ightham Fissure, Kent,
the greater number are referred to this species by E. T. Newton, who
states that they are sufficiently perfect as to leave little doubt as to the
correctness of the determination (Quart. Journ. Geol. Soc, May 1894,
192).

The period of gestation is unknown. The number of young at a
birth does not appear to exceed one (Blasius), born probably in the
latter half of June (Proger): they have been found not long after birth
on 25th July (J. Backhouse, jun.).

Description :—Natterer’s Bat resembles Bechstein’s, but is smaller
and with quite distinct ear and wing.

The head (Plate XI., Fig. 3) is proportionately small, the upper
jaw overhanging and naked at its extremity, the face is hairy and
moustached, and there are two prominent glands, one on each side
above the lips.

The ear (Fig. 2, No. 8, p. 7) is large, and when laid forward reaches
slightly beyond the tip of the nose. It is nearly oval in shape, with con-
vex inner margin, and the outer also as a whole convex, but its regularity
interrupted by a notch at about one-third of the distance from the tip
and ending in a distinct basal lobe. There are only five cross-folds.
The tragus is long, narrow, and pointed, and attains its greatest
breadth near its base, where it is deeply emarginate and conspicuously
lobed ; its tip lies about opposite to the upper cross-fold at about two-
thirds the height of the ear. The inner margin is nearly straight, or
slightly convex, the outer slightly serrated and concave above.

The wing (Plate XIII., Fig. 3) is the longest found amongst British
bats of this genus, both forearm and longest digit being of exceptional
length. It is also broad, since the third and fifth metacarpals are about
. of equal length. On the other hand, both foot and lower leg are small,
relatively and actually, and the interfemoral membrane is narrow and
inconspicuous. Its posterior margin is crenated and furnished with a
fringe of stiff short hairs, most conspicuous between the tail and the
calcar, which is almost as long as in JZ. daubentont. The thumb is weak.

The fur is long, thick, and soft, and extends to the face so as to

PLATE. XIV;

HEADS (natural size) OF

1. Myotis bechsteini. 2 and 3. Barbastella barbastellus (front and side views).
4. Plecotus auritus.

NWATTERER'S 'BAT 181

conceal the small eyes. On the wing it extends above and below
about to a line joining the central points of humerus and femur, and
is not conspicuous on the interfemoral membrane. The foot is ciliated
with dusky hairs.

In colour this bat is very distinctly bicolored, being greyish brown—
between “wood brown” or “isabella” colour, and “drab”—above and
whitish below, the bases of the hairs always dusky or dark brown,
and the tips grizzled and shining above. The line of demarcation is
distinct, and passes approximately from the angle of the mouth to the
junction of the wing and body. The membranous parts are dusky.

In the young the grizzled hair-tips are wanting, and the colour
contrast is greater, the upper surface being darker—* mummy brown”
almost to “seal brown”—and the lower sur-

face a clearer white. The skin of the foetus
is dusky (Proger).

In the skull (Fig. 16, No. 3, p. 174) the
profile resembles that of MZ. mystacinus, but
the brain-case is far more prominently raised
above the face-line and more rounded.

In the teeth (Fig. 16, No. 3, p. 174, and

Fig. 18) the middle upper premolar (there is
some variation) is usually about half the size
of the anterior, but sometimes almost equals
it in diameter and length.

Geographical variation has not been
studied, except in the sub-species MZ. x.
bombinus of Japan, which is stated to be
characterised by more abrupt and consider-
able inflation of the frontal region of the Fic. 18—FRonr ViEW oF
skull and by darker colour. INCISORS AND CANINES OF

= z - Sar : Myotis nattereri (enlarged and

Dimensions in millimetres (see following ub Saar
page).

Proportionate lengths :—Foot, without claws, about -48 of lower
leg ; third metacarpal, about -99 of fifth; lower leg, about -41 of fore-
arm, and about -36 of head and body.

Skull :—Greatest length, 15-5; basal length in middle line, 11-25 ;
palatal length in middle line, 7; from posterior border of 7? to anterior
border of canine, 6; ditto in lower jaw, 6; greatest breadth at zygoma,
9:5; posterior breadth, 7-5; breadth between orbits, 7; breadth at
constriction, 4.

Distinguishing characteristics:—The light colour, large notched
ear, elongated tragus, and particularly the ciliated interfemoral
membrane, distinguish this bat absolutely from any other British
species.

182 VESPERTILIONIDAZ—MYOTIS

DIMENSIONS IN MILLIMETRES :—

MALEs.
B& S 8 3 2
ao (o} ° — . o
2 | 8 | |) 2) eu) a] eal Ee) oe
Sle feels joe | ee) Bee eS | ao
= 2 ae s Oo) | se & ~8 | @ = a 3.8
s Ss | Re] & Be area NERO te ree ee 8 2g | Se
cs) 2 | Hs SYS |S a 3 fey || 1S
g oo $ H | = cS) a) Sin ee
ll os bp # A A Be ll |
Cy fen] 3
* |
Male (British Museum) . | 42 16°5 | 10 38 17 8 40 6 66°5 | 33 32°,
Male, Bedfordshire (mea-
sured from dried skin) . | 42 LBS ci eee 41 ee ee:
Male, Salwar: (Dublin
Museum) . 5 : is oe a be) | itee 8 39 6 68°25 | 34 33
FEMALES.
Female, received from A.
Whitaker, Yorkshire, 10th
October 1906 (measured
inthe flesh) . 43 we ac 82,
Female in alcohol (Tomes’
collection, in British
Museum) 14 10 AO 16°5 8°25] 40 5 64 33 32
Female (do.) 14 9 se HPL: 8 40 6 65 33 32
Female (do.) 14°5 9°5 50 16 8 89 6 66°5 84°5 | 34
Female (do.) 14 8 =o) 6 8 40 | 4:75 | 69 85°5 | 33
Female (do.) 14 9°75)|) wee 16°5 8°5 88 6°5 | 63°75 | 38 82
Female (do.) 17°5 9 ae 16°5 8 89 6°75 | 65°5 33 32
Female (do.) ne ed etal ea! 8 Be alhitir 75 | 89 | 6 65°5 | 34:5 | 38
Female, Northants . . | 46 14 9 36 17 75 | 39 6°5 | 65 35 33
Female, do. 47 14 9 33 16 8 39 6 65 34 33°5
Female, England (Bartlett:
British Museum) . 42 15 9°5 16°75| 8 39 5 65°75 | 84°5 | 34
Female, Berks, measured
from dried skin 3 - | 49 165 | 9 40 as ae oh oa Ae oe Ae 267
Female, Ilminster . 46 17 Ao 43 Se a5 42°5 as aC avs ‘i 285
Female, N. Wales (the last
threein British Museum) | 43 16 8 42 ate an ais aie Af as a 272
Female, Co. Fermanagh
(Dublin Museum) . oh cas sis Se ee 225) 9 39 6:0 | 67°5 | 85 | 34°75
Average of females i i! : F r
(approximate) 45°1 | 14:9 9 87°6 | 16°5 8 39°4 | 5°8 | 65°6 BBP | bs! [274]

Note :—The above measurements, except where otherwise stated, were taken from specimens in alcohol.

This species, to which the inappropriate name of Reddish-
grey was applied, it is believed, in the first instance by Bell, was
described by Kuhl in 1818, and by him named after the
Austrian naturalist, Natterer. In Britain it was recognised by
Jenyns (who gave it a place in his W/anuaZ), by Yarrell,* William
Thompson of Belfast, and Bell, but it for long retained the
reputation of being, if not of unfrequent, at least of purely local
occurrence. It is now known to be widely distributed in some
numbers throughout the greater part of England, having been
found in almost every district where bats have been the subject of
expert study.

1 In Jenyns.

————

NATTERER’S BAT 183

Natterer’s Bat is as sociable and gregarious as the Whisk-
ered Bat was formerly reputed solitary. It is often abundant
in wooded localities, where it haunts, usually with slow and
steady flight at no great altitude, the open spaces amongst the
trees or the neighbourhood of old gardens. By day it retreats
to holes in walls! or trees,? or to caverns, and in the latter
is often found in company with other species, such as the
Whiskered and Long-eared Bats, with which it will even share
the same crevice. The recesses of caves are, however, more
favoured for purposes of winter hibernation than for the briefer
retirements of summer. Occasionally it breaks its rule by
hanging alone in hibernation.

A favourite winter resort is Mr Heatley Noble’s cavern
near Henley-on-Thames. Here it was found numerously in
company with other bats on 14th February 1906. By the
15th of the following month, however, all except four, and
they, strange to say, numbering as many species as individuals,
had departed, and by the 25th of the same month, only one,
a Daubenton’s, remained.* Hibernation is, therefore, not
more profound than in other species, and in South Wales,
Mr T. W. Proger has known this bat to fly abroad for its
food in February.

The late R. F. Tomes graphically described for Bell’s work
a colony of these bats which he discovered in 1848 in the
church of the village of Arrow, near Alcester. ‘‘ Between the
ceiling of the church and the tiled roof was a dark retreat,
accessible by a low arch from a floor in the tower. Here the
Bats were seen adhering, by all their extremities, to the under
surface of the row of tiles which forms the crest or ridge of the
roof (partly supported, however, by the upper tier of roof-tiles
on which the ridge-tiles rested), and others clinging to them,
until a mass was made up three or four inches thick, six or
seven wide, and about four feet in length. It would be wrong
to call this their place of repose, as they presented a most
singular scene of activity, the constant endeavour of those

1 Alfred Newton, Zoologist, 1853, 3804; J. Backhouse, zézd., 1898, 493.
2 William Borrer, zdzd., 1874, 4127.

3 Joseph Armitage, Waturalzst, 1900, 114, and other references.

4 A. H. Cocks, Zoologist, 1906, 186-187.

184 VESPERTILIONIDAZ—MYOTIS

outside being to penetrate the mass, probably for warmth, and
to do this they were continually poking their noses between
those nearest to them, and then forcing in their bodies, to be
in their turn again pushed to the outside. In this manner a
regular bickering was kept up in the whole mass. However,
they seemed to be very gentle, and to have no idea of biting or
otherwise annoying each other.

“On the boarded floor in the tower adjoining this retreat
many dead ones were lying about, in a dried condition, all of
them very small and hairless. These probably had fallen from
their mothers when on the wing, as they were themselves too
young to have flown there, and the parents could not have
rested in this chamber, and at that time let fall their young.

“After watching this remarkable assemblage for some time,
about sixty were secured in a bag (only a very small proportion
of the number there), and the bag was opened in a lighted
room in the evening. They were soon flying about in all
directions. On the window being thrown open, those nearest
to it at once flew out; but so completely gregarious are these

Bats, that after taking a turn or two outside they re-entered’

the room, and being joined by others, again went forth, and
again returned, until all had become aware of the means of
escape, when the whole company left the room in a cloud. We
may add, as further showing the gregarious nature of the
species, that a few which were retained, exhibited great uneasi-
ness when separated from each other, which disappeared when
permitted to be together.”

The gentle nature of this bat was also remarked upon by
Bell, who kept three in captivity for some little time, and found
that they readily took food from his hand, yet their apparently
friendly disposition, both to himself and to their companions,
did not prevent two of them from devouring one night the body
of the third. They were active in their habits, “running about
the cage and climbing with great agility.”

Mr J. E. Harting,’ who has seen a good deal of this bat in
west Sussex, finds it in the habit of flying about the oak
trees on the outskirts of the woods, appearing earlier in the
day—even before sunset—than the other local species, and

1 Zoologist, 1889, 241-248.

NATTERER’S BAT 188

allowing so near an approach as to be identified while on the
wing. Its flight when feeding was by no means rapid, but on
leaving one tree for another at a little distance it flew much
faster, yet never so rapidly as the Pipistrelle or high-flying
Noctule. So far as could be ascertained without actual
examination of the prey captured, its food appeared to con-
sist principally of small flies and moths, which it captured not
only on the wing, but snatched off the leaves on the outside
branches of the trees with great dexterity. Just as a dog will
“bolt” a rabbit and catch it before it has gone many yards, so
one would disturb a small moth and seize it within a few inches
of the leaf or twig on which it had been resting. Apparently
the bat is noisy while on the wing, since its voice recalled to Mr
Harting the well-known lines in Collins’s “Ode to Evening ” :—

“ Now air is hushed, save where the weak-eyed Bat
With short shrill shriek flits by on leathern wing.”
At Colchester the peculiar squeak is heard more frequently by
Dr Henry Laver than the voice of any other species.

Mr Harting has noticed Natterer’s Bat on more than one
occasion feeding in bright sunshine, as early as 3 P.M. in
August. His observations are confirmed by Mr J. Ffolliott
Darling,’ who once caught one in April in County Cork flying
about a wood in bright sunlight at about the same hour, and
by Messrs T. A. Coward and Charles Oldham, who obtained
a good view of one outside the Cefn Caves, near St Asaph,
Wales, again at 3 p.m. on 2nd December 1905. The last,
however, had possibly been disturbed from the caves.

The young, the number of which, according to Blasius, does
not exceed one each year, are found in summer in the mixed
colonies of their elders. Mr J. Backhouse, jun.,” received several
already well-grown from the ruins of Harlech Castle, north
Wales, in July (Mr Oxley Grabham writes me that the exact
date was the 25th), and this agrees with Mr Proger’s state-
ment that in south Wales they are usually born towards the
end of June, on the 22nd of which month he once caught a
female bearing her young on the wing.®

1 Zoologist, 1883, 294. 2 Tbid., 1898, 493.

3 Paper read before the Biological and Geological Section of the Cardiff
Naturalists’ Society, March 1905, 5; also in MSS.

186 VESPERTILIONIDAZK—MYOTIS

The best account of this bat while in captivity is that of
Mr Coward,’ based upon an example which he had in his
possession for some little time. He corroborates previous
writers in regard to the docility and gentleness of the species,
which readily acquired the habit of eating mealworms after
two had been placed in its mouth. Mr O. V. Aplin? alone
accuses it of fierceness: “having bitten, it retains its grip with
the tenacity of a bulldog.”

According to Mr Coward, the two most remarkable features
were the method of carrying the tail in flight and the manner
of alighting. Unlike many other species, such as its congener
the Whiskered Bat, it carried its tail in an extended position
behind it, not half-bent beneath the body. In spite of this,
the tail resumed the curve usual in bats when the animal rested
on a flat surface, and troublesome insects were pouched in the
orthodox manner if it was fed when hanging or resting in a
horizontal position.

This bat devoured—besides mealworms—flies, bluebottles,
crane-flies, spiders, wood-lice, beetles and moths, in the latter
case rejecting the head and wings. Though usually attempt-
ing to eat anything that was offered to it, it was more particular
about its food than some other species, and it invariably
dropped cooked or uncooked meat or shreds of fish. ‘ Most
spiders,’ Mr Coward writes, ‘‘were eaten rapidly, but one
or two with conspicuous markings were snatched and then
rejected ; a carnivorous beetle* was not only treated in the same
way, but the Bat, by spitting and shaking its head, showed
evident signs of disgust.”

‘‘ At times it used the carpus to hold a struggling mealworm,
and would stand, when engaged in eating, with one wing
slightly raised, as if ready to hold its prey if it proved too
powerful; it never used the thumb in any way to tear its prey.
In its normal position when feeding, its head was held rather
low and its shoulders were somewhat hunched up. When
searching for food or flying round, it either held its mouth open
or chattered, opening and shutting its mouth with great
rapidity.” Its sight appeared to be bad and it made many wild
snatches at nothing when an insect was moving in its cage, yet

1 Zoologist, 1905, 51-56. * Ibid., 1889, 382. 3 Carabus.

NATTERER’S BAT 187

it generally managed to clear its cage of living insects during
the night. |

When sleeping this bat did not always hang by its feet, but
often lay prone on the floor of the cage. It did not as a rule
remain on the wing for more than ten or twenty minutes at a
time, after which period it would alight for rest. In doing so
“it frequently seized hold first with its thumbs, and then did
not shuffle round so rapidly as those Bats do which invariably
settle in this manner; a Noctule, Long-eared or Whiskered
Bat clutches the object to which it intends to hang with its
thumbs, and with great rapidity twists itself round so as to
gain the usual reversed position. The Lesser Horseshoe per-
forms a more remarkable feat: it flies to within an inch or so
of the object, and then reverses itself in the air, catching first
with its feet ; in this way it is ready at once to drop from its
hold and fly. When my Natterer’s Bat wished to settle on
certain objects—especially on the tip of one of a pair of Fallow-
deer antlers over the door—it sometimes turned in the air after
the manner of a Horseshoe. This action was not so clean or
certain as in the case of a Horseshoe, and occasionally the Bat
missed its hold and fell, generally recovering itself before it had
fallen many inches. It was somewhat remarkable that it shared
the habits of both the Vespertzlionzde and the Rhznolophide
(for the Greater Horseshoe reverses in the air) in this respect,
but did not perform either action with the same celerity or
ease.”

Its voice is described as consisting of two notes, of which
‘“‘one—pitched much lower than the other—was a low chatter
rather than the usual high-pitched cry of a Bat.”

Mr Coward suggests that “if some of the habits of this
example were typical of the species, we may see in Natterer’s
Bat the first traces of habits which have become constant in the
specialised Rhznolophide. In the Horseshoes the short tail is
carried recurved over the back; in Pterygzstes,' Pzpistrellus,
Plecotus, and some species of JZyotzs, it is usually carried
curved beneath the body; in JZ. natterert we find the tail,
although used as a pouch, is carried extended behind the body.
Again, the habit of turning in the air before alighting appears

1 Le, Myctalus.

188 VESPERTILIONIDAE—MYOTIS

to be constant in the Akzxolophide, and seldom noticeable in
the Vespertzlionide, except in this species, where we find the
Bat sometimes alights in one way and sometimes in the other.”
Thus, in habits, Natterer’s Bat shares some of the character-
istics of each family, and it is noticeable that the small
interfemoral membrane and weak thumbs recall those of the
Horseshoes, so that structure and habits go together. In
other respects, such as the position of the wings when sleeping,
Natterer’s Bat is not known to differ from its near relations.
Mr Arthur Whitaker’ has described an interesting ex-
periment, which, without resorting to the cruel artifices
of Spallanzani and other continental naturalists, enabled him
to demonstrate that the flight of a bat of this species was in no
way dependent upon the uninterrupted use of its eyes. The
animal having been rendered temporarily stone-blind by means
of wax, was released in a room in which it had never been
before and with which it was consequently quite unfamiliar.
Usually when captive bats are allowed to exercise themselves in
this room they fly in circles close to the ceiling, but the
behaviour of this blinded bat was somewhat different. ‘“ When
released,” writes Mr Whitaker, ‘it commenced to fly in a rather
slow and hesitating manner, but with rapidly-growing con-
fidence. It went first straight for the closed door, and, I
thought, was about to fly right against it, but it suddenly
turned itself when but a few inches off, and hovered slowly once
or twice along the top edge and down the side, still without
touching, but following, I feel convinced, the slight draught of
air admitted. Having apparently satisfied itself that there was
no exit large enough for it there, it turned round, and flew the
length of the room, straight for the fireplace, still, I believe,
following the draught. When it got near the fire it turned,
warned, no doubt, by the heat, and then commenced to fly
slowly and cautiously about the room at a height of about six
inches from the floor, and I noticed it repeatedly pause and
hover in front of the wainscot at one point where it had sprung
slightly from the wall and admitted a distinct current of air.
Although it flew fairly quickly, and kept passing underneath
the chairs, of which there were over a dozen in the room, it

! Naturalist, 1906, 148-149.

lt Si

Se ee eee ee le

—_—

THE NOTCH-EARED BAT 189

never once, so far as I could see by lying down to watch it,
even touched anything with the tip of its wings. An attempt
on my part to catch it caused it to fly up to the ceiling, and
just below this it commenced circling round and round rapidly,
repeatedly dipping to pass under a beam crossing the centre of
the ceiling. I tried holding a walking-stick perfectly still in
its path, but it would swerve suddenly when but a few inches
from it. After flying for over twenty minutes it suddenly
settled on a chain supporting one of the weights of the gas
chandelier, and that it could settle in such a place is in itself a
wonderful proof of the accuracy of this ‘second sight.’

“T stood on a chair and approached my hand very slowly in
order to catch it again, but when my hand was within about a
foot of it, it commenced to turn its head nervously and jerkily
from side to side (an action characteristic of a bat when dis-
turbed), and flew again before I could get hold of it.

“Eventually I was obliged to get out my butterfly net to
catch it, and even then had some little difficulty. . . .

‘“When I caught my bat again I found the wax still adher-
ing properly and quite covering the eyes.”

Natterer’s Bat, should a tolerable view be obtained of it, is
one of the easiest of all British species to identify while on the
wing. The light colour of the under parts, the long ears—
longer than those of any species except the rare Bechstein’s and
the Long-eared—and the general size, which is about a quarter
as large again as the Pipistrelle, Daubenton’s, or Whiskered Bat,
form an unique combination amongst British species.

[THE NOTCH-EARED BAT.
MYOTIS EMARGINATUS (Geoffroy).

A few lines are necessary in regard to other reputedly British
species of this genus.

Of these the present was described and figured by Isidore
Geoffroy in 1806,' and redescribed in error by J. H. Blasius
in 1853.” Geoffroy’s types came from Abbeville, France, and
the inclusion of the species in the British list is due to his

1 Ann. du Museum @ Hist. Nat., viii., 198.
As ciliatus, Wiegmann’s Archiv fir Naturgeschichte, X1X,, i., 288-293.
R 2

190 VESPERTILIONIDA—MYOTIS

statement, which forms part of the original description, that he
received from Alexander Brogniart a single specimen taken
“sur sa route” near Dover. From this scanty information
he concluded that the species must be ‘‘assez commune en
Angleterre.”

Many naturalists have expressed conflicting opinions as to
this bat, which is found so near Britain that its occurrence
within our boundaries would not appear to be very unlikely.
De Selys’ appears to have been the first to accept it as a
distinct species allied to JZ. nattererz, and R. F. Tomes,?
although he believed that Geoffroy’s type-specimen was a
M. mystacinus, after examination of specimens in continental
museums, pointed out that it was a form quite different from
anything British.

This bat, which ranges at least as far east as Hungary,
combines an ear even more fully notched than that of JZ.
mystacinus, with a narrow interfemoral membrane, its posterior
border ciliated but less thickly so than in AZ. natterert. The
forearm measures about 40 mm., the foot about 9 mm.
The middle upper premolar is exceptionally small and slender,
and scarcely rises above the flesh in which it is imbedded. |

[THE MOUSE-EARED BAT.

MYOTIS MYOSOTIS (Bechstein).

The Mouse-eared Bat was hesitatingly included by Bell in
the British fauna on the strength of specimens supposed to
have been taken in the gardens of the British Museum.
Since then it has been occasionally reported from other
localities, including, as collected by Mr J. E. Harting,’? Sher-
borne, Dorset (C. W. Dale), Epping (F. Doubleday), Fresh-
water, Isle of Wight (Hadfield), and Ireland (Blake Knox):
no doubt Mr Harting’s expression of misgiving as to the
correct determination of the species was not over-cautious.

All the above records are of little, if any, value, and that
for the Isle of Wight was expressly contradicted by A. G.

' Faune Belge, 1842, 20. * Zoologist, 1856, 4938-4939.
3 Jbid., 1887, 161-162, footnote.

THE MOUSE-EARED BAT I9I

More.’ In 1903, however, Mr J. Lewis Bonhote’ noticed a
genuinely British-killed specimen in the University Museum of
Cambridge. This was taken at Girton in 1888 by a lady who
brought it alive to Dr Hans Gadow.

This species has suffered much at the hands of slipshod
nomenclaturists. It is clearly the Vesfertzfo myosotis of Bech-
stein,®> and is as certainly the type of Jakob Kaup’s genus
Myotis of 1829, so that there can be no doubt about its
correct title, according to modern usages. Unfortunately,
however, the majority of continental naturalists, beginning
with Schreber in 1775, finding it their commonest or most
conspicuous bat, supposed that it must be also the ‘‘Common
Bat,” Vespertzlio murinus of Linnzeus, and thereupon trans-
ferred the latter name to it, whereas, as has been already shown
(p. 51), the bat thus designated is a very different species.
Unfortunately the confusion did not stop at this point, since
British naturalists, following the lead of their continental
brethren, applied the name murznus to the Pipistrelle, the
Common Bat of their country.

Considering that this bat is reputed to be one of the most
frequent on the European continent, even in those countries,
such as Normandy, which are separated from England only by
the breadth of the channel, it is very remarkable that it should
be unknown as a regular inhabitant of Britain. But it cannot
be doubted that Bell was correct in believing that, were it really
British, its large size and the comparative ease with which
specimens can be obtained, would long since have brought it
under notice. Mr Bonhote repeats this argument in regard to
the Cambridge example, which, although undoubtedly killed in
Britain, must, he thinks, have been brought over accidentally
from the continent.

The Mouse-eared Bat, should it occur again in Britain,
can hardly be mistaken for any other. In size it is at least
the equal of, perhaps superior to, our three largest species, the
Noctule, Serotine, and Greater Horseshoe. In form it is
rather, however, a large Bechstein’s Bat, with relatively smaller

1 Zoologist, 1894, 148. 2 Jbid., 1903, 387.
° Der Zoologe, v.-viii., 46, 1797, corrected to V. myotzs by Bechstein, in Gemeznut-
sige Naturgeschichte Deutschlands, &c., I., ed. 2, 1154, 1801.

192 VESPERTILIONIDAZ—PLECOTUS

ears extending, when laid forward, only just beyond the tip of
the muzzle; shorter and roundly triangular, not sickle-shaped,
tragus reaching not more than half the height of the ear; anda
very small middle upper premolar lying in a position completely
interior to the tooth-line. |

GENUS PLECOTUS.

1816. MACROTUS, W. E. Leach, Systematic Catalogue of the Specimens of the Indi-
genous Mammalia and Birds that are Preserved in the British Museum, 1; based
on Macrotus europeus of Leach, the “ European Longear,” from Devonshire, Eng-
land, presented by G. Montagu ; named, but without description, hence Plecotus
has priority.

1818. PLECOTUS, Etienne Geoffroy, Description des Mammiferes qui se trouvent en
Egypte, ii., 112, 118-119, pl. ii, No. 3 (for date see Sherborn, Proc. Zool. Soc.
(London), 1897, 287-288; based on /orezllard of Daubenton, Za darbastelle, and
an undescribed species from Timor.

1829. PLECAUTUS, Francois Cuvier, in Dictionnaire de Sciences Naturelles, \ix., 415, a
misprint for P/ecotus ; no species mentioned.

This genus, although small in known species, has a wide
distribution, mainly in the Palearctic region, where it ranges
from Ireland to Sakhalin, and from about 60° north latitude in
Skandinavia to north Africa. There is one British representa-
tive, the well-known P. auritus. P. puck, which I described
from Murree, northern India, differs in cranial characters, while
my LP. ¢eneriffz, from Teneriffe, is larger, having the fore-
arm measuring about 44 mm. (see Aun. and Mag. Nat. Hist,
December 1907, 520-521); G. M. Allen’s P. sacrimontzs, from
Mt. Fusi- Yama, Japan, has a long thumb (11°7 mm.). Several
other specific names have been instituted, but are of unknown °
value. They are mentioned below on p. 195. In North
America it is represented by the closely allied Corynorhznus,
differing in the shape of the nostril, conspicuously glandular
muzzle, and in the proportions of the wing.

Generic characters:—These are bats of medium size, with
immense ears (Plate XIV., Fig. 4), meeting by their inner
margins on top of the head, the outer margins terminating
just behind the angles of the mouth. The tragus is large and
elongated.

The muzzle bears the elongated, narrowly crescentic nostrils

PrLECOTUS 193

at its extremity, in front of deep grooves separating the hairy,
flat, and depressed central region from the glandular sides.
There is no groove in front of the nostrils.

The wing (Plate XVII., Fig. 1) is broad and arises from the
base of the toes. The antibrachial membrane is well developed.
The fifth metacarpal is a little shorter
than the third and fourth.

The calcar extends about half-way
from foot to tail. There is no post-
calcarial lobe. The tail is almost as I
long as the head and body, and has Anh)
the tip slightly exserted.

ithe skull (Fig. 8, No. 4, p. 101)
is weak ; the profile descends gradu-
ally from the inflated brain - case to
the depressed, broad, not saddle -
shaped facial region; the zygomata ae
are flattened; the cranial crests are
very slightly developed ; the auditory
bullz are very large, fully equalling, or,
perhaps, exceeding, in size those of
much larger bats, such as Wyctalus

noctula, Vespertilo serotinus or Myotzs Fic. 19—DIAGRAM OF ARRANGE-

On MENT OF TEETH IN GENUS
my OSOLLS. t Plecotus.

There are two less teeth (Fig. 8, (2) Upper and (o)ilowen eee
an 4, p: 101 ; Fig: 19; and Fig. 20,
p. 197) than in JZyotzs, one pair of upper premolars being absent ;
two more than in Barbastella, the formula being :—
6 4 ee = I

2-
Seeec rsa) 3

2 3-
2m 373 = 36.
=

The upper incisors point directly inwards; the inner is
bifid, with inner cusps much exceeding the outer; the outer is
smaller and unicuspid. The canine is without accessory cusps.
The anterior upper premolar, although small, is distinctly visible
externally. The posterior upper premolar is large. The three
lower premolars graduate in size from the posterior to the
central, which is smallest.

194 VESPERTILIONIDA—PLECOTUS

THE LONG-EARED BAT.
PLECOTUS AURITUS (Linnzus).

1740. THE DOUBLE-EARED Bart, Eleazar Albin, 4 Natural History of Birds, iii., 95,
pl. ci.

1758. VESPERTILIO AURITUS, Carolus Linnzus, Systema Nature, x., 32 (6) ; xii, 47 (5),
1766 ; described from Upsala, Sweden ; Bingley ; Donovan; Jenyns ; Clermont.

1789. (?) VESPERTILIO AURIBUS, Gilbert White, Se/borne, Letter xi. to Thos. Pennant,
oth September 1767 ; named, but’ without description, from Selborne, England.

1816.(?) MACROTUS EUROPAUS, W. E. Leach, Systematic Catalogue of the Specimens
of the Indigenous Mammalia and Birds that are Preserved in the British Museum,
etc. (London), 1 ; named, but without description, from Devonshire, England, from
Montagu’s specimens.

1818, PLECOTUS AURITUS, Etienne Geoffroy, Description des Mammiferes qui se
trouvent en Egypte, ii., 118, pl. i, No. 3; Fleming; Bell (ed. 1); MacGillivray ;
Blasius ; Fatio; Dobson; Blanford ; Flower and Lydekker ; Lydekker ; Thomas ;
Méhely ; Johnston ; Cabrera; Millais ; Trouessart (1910).

1825. VESPERTILIO OTUS, Friedrich Boie, Oken’s Jszs (Jena), ii., 11, 1206 ; described
from Copenhagen, Denmark.

1826. VESPERTILIO CORNUTUS, —— Faber, Journ. cit., i, 5, 515; described from
Horsens, Jutland, Denmark.

1827. PLECOTUS COMMUNIS, R. P. Lesson, Manuel de Mammatlogie, 95, No. 232.

1827. PLECOTUS CORNUTUS, auct. et op. cit., 96, No. 234.

1828. PLECOTUS BREVIMANUS, Leonard Jenyns, Trans. Linnean Soc. (London), xvi.,
55, pl. i, fig. 2; described from an immature specimen from Grunty Fen, Isle of Ely,
England ; not P. drevimanus of Bonaparte, 1841 ; Bell, eds. 1 and 2 (figures).

1829. PLECOTUS VULGARIS, A. G. Desmarest, Faune Francais, Mammiféres,
livr. 19, 18; pl. 2, livr. 16, fig. 3, described from France.

1829. VESPERTILIO BREVIMANUS, J. B. Fischer, Syxopsis Mammalium, 118 and 553.

1832. P. PERONII, Isidore Geoffroy, Guerin’s Magasin de Zoologie, ii., Classe L.,
No. 2, pl. iii, fig. 1 ; described from a male and female obtained, locality unknown,
during the voyage of Péron and Lesueur.

1860. PLECOTUS KIRSCHBAUMII, Carl Koch, Achter Bericht der Oberhessischen
Gesellschaft fiir Natur- und Hletlkunde, 40 ; described from Oberhessen, Germany.

1862-63. PLECOTUS AURITUS, TYPUS, MONTANUS ET BREVIPES, Carl Koch, Jahroiicher
des Vereins fiir Naturkunde im Herzogthum (Nassau), 406-408 ; described from
Nassau, Germany.

1873. PLECOTUS LEUCOPHEUS, N. A. Syevertzoff, Trans. Soc. Nat. Moscow, viii., 2 ;
translated by Craemer in Anum. Mag. Nat. Hist, July 1876, 42: named, without
description, from north-western Turkestan.

L Oreillard*‘ of the French, de langihrige Fledermaus of the Germans.

Distribution :—The Long-eared Bat ranges throughout boreal and
transitional into tropical Europe and Asia, from sea-level to about
6500 feet in the Caucasus (Satunin); from 60° N. latitude in
Skandinavia and Russia (Blasius) to north Africa, including Tunis?

1 Horned Bat, of Cheshire (Coward and Oldham) ; see also under PIPISTRELLE,
p- 105. 2 Specimens seen.

THE LONG-EARED BAT 195

and Egypt at least to the fifth Cataract (Bell); and from Ireland to
Palestine (Tristram) and Hokkaido, Japan.! Its area of distribution
includes the Islands Malta,! Sicily (Dobson), Minorca and Ibiza
(Cabrera; Barcel6). Specimens from more than one locality have
formed the basis of specific names, eg., donapartit of Gray (Mag.
Zool, and Bot., ii., 495, 1838) from Sicily; egyptiacus of Geoffroy from
Egypt ;” christzz of Gray (loc. cit.) and ustus of Heuglin, from Wadi
Halfa, north Africa; and homochrous of Hodgson (Journ. Asiatic Soc.,
Bengal, XVI, ii. 894, 1847) from the central sub-Himalayas. The
status of these forms, if indeed they be distinct from typical auritus,
is quite unknown.

The Long-eared Bat is probably one of the commonest, if not the
commonest, and most widely distributed in Britain, but its habits neces-
sarily render it less conspicuous than many others. It is certainly to
be found everywhere in England, including the Isles of Wight (More;
Bury, Zoologzst, 1844, 777; Wadham), and Scilly (Clark), and almost
with equal certainty in every county of Wales (see Forrest, 7vams.
Caradoc and Severn Valley Field Club, 1900, 242). It is common in
all the Channel Islands (Bunting).

In Scotland there are so many records of its abundant occurrence in
the low-lying southern, central, and eastern counties to the Moray
Firth, that this part of the country may probably be reckoned with
England and Wales. Service, for instance, believes that in Solway it
“is much the most abundant [bat] that we have,” and in Aberdeen it
is stated to be commoner than the Pipistrelle (Sim; Kinnear). In the
highlands it becomes rarer, but details are, as a rule, lacking for areas
north of the Caledonian Canal. Its range may, however, include the
extreme north, since Eagle Clarke has a note from a correspondent
that in west Inverness, north-west of Fort William, it outnumbers
all others, and Sir Joseph Fayrer found it in Sutherland (Dobson,
Report British Association, Swansea, 1880, 183). W. Evans cites an
out-of-the-way locality in rocks at Dumglow, Cleish Hills, Kinross-shire
(Proc. Roy. Phys. Soc.(Edinburgh), xvi., No. 8, 388, 27th November 1905).
In the islands it is known from Arran and Mull (Alston), Islay (Alston ;
Gilmour, Ann. Scott. Nat. Hzst., 1897, 191), and North Uist (Millais),
which facts suggest a wide distribution.

It is included by Kermode (Zoologist, 1893, 62) in the Isle of Man
fauna, but is stated to be not so common there as Pifistrellus
pipistrellus.

In Ireland it is probably found commonly in every county, having
been known to Alcock and Moffat from all except eleven (/rzsh
Naturalist, 1901, 250-1), and there can be but little doubt that the

1 Specimens seen,
2 Or egyptius, as frequently cited, but original description not found.

196 VESPERTILIONIDAE—PLECOTUS

vacancies represent rather lack of observers than of the bat. It has
been taken on such desolate western islets as the Tearaght, Co.
Kerry, on 4th November 1901 (Barrington, A/zgration, 284), and by
Sheridan on Achill, Co. Mayo (Dublin Museum).

Distribution in time:—A portion of a humerus from layer 11 of
the Pleistocene deposits of Hoe Grange Cavern, Derbyshire, has been
doubtfully referred to this species by Arnold Bembrose and E. T.
Newton (Quart. Journ. Geol. Soc. 28th February 1905, 50).

The period of gestation is unknown. The number of young is one,
born most usually in June or July.

Description :—The general form and appearance are typically those
of its genus.

The head (Plate XIV., Fig. 4) is only slightly raised above the face-
line; the muzzle rather long and prominent, horizontal, and slightly
emarginate ; the nostril with prominent edges, the opening lateral, pass-
ing backwards and outwards into a distinct groove. The ear is so large
as to almost equal in length the entire head and body. It is oval,
oblong, semi-transparent, and transversely folded. The inner margin
is bent outwards, and forms a broad longitudinal fold, ciliated at its
edge as well as along the keel formed by the bending. Immediately
above the point where the two ears meet, their margins form an angular
notch; above this, on each side, a small lobe projects laterally, so that
when the ears are erect it touches its fellow; these lobes are hairy,
thicker, and more opaque than the rest of the ear. The tragus is
elongate, lanceolate, rather obtuse, bending very slightly outwards, and
having a length about two-fifths that of the whole ear. The con-
spicuous eye is placed slightly in front of the inner angle of the base
of the tragus.

In the wing (Plate XVII., Fig. 1) the interfemoral membrane
is conspicuous, and is supported by a remarkably long tail and lower
legs. The foot is also large, but the calcar, although strong, is not of
exceptional length. It terminates in a distinct lobe.

The fur is rather long, soft, thick, and silky; its abundance on the
shoulders gives the body a broad appearance. On the wing it extends
about as far as lines joining the centres of humerus and lower leg.
The ear, except for fringes on the folds, is almost hairless. The foot
is ciliated.

The colour is, above some shade between “wood brown” and
“broccoli brown,” below dirty white or yellowish. The basal portions
of the hairs are everywhere dusky. The line of demarcation follows
the wing and thence to the angle of the mouth, but is not very
clearly marked. According to Bell, young specimens are redder,
old ones greyer; but Dobson thought the young and females darker
dorsally.

THE LONG-EARED BAT 197

I have seen no evidence of seasonal variation.

The skull and teeth are typical of the genus.

Individual variation manifests itself not infrequently in albinism,
partial or complete. White or “albino” varieties (the colour of the
eyes not stated) are mentioned by New-
man, Fzel/d, 21st March 1874, 271; Borrer,
Zoologist, 1874, 4128 (two); Baring, Journ.
ctt., 1898, 261. I have also records of a
one with the foreparts paler than usual
(Bury, Journ. cit., 1844, 777); a buff one
(in Dublin Museum of Science and Art),
and one of a uniform cream colour, with
pink eyes and nails (Oldham, /Journ. cit,
1890, 349). Mlillais has seen a true albino
and a cream-coloured variety from Nor-
folk, and Eagle Clarke (Waturalist,1894, 68)
mentions a melanic one.

The extent of geographical variation
is quite unknown. Cross-channel speci-
mens resemble British, but Hungarian
and central European average larger and
are lighter in colour, with apparently larger
ear and smaller foot, and may possibly ee ee ee
represent a hitherto undescribed sub- (niarged and Ee oes vie i
species. Those from Africa and others
from sandy regions are said to be of pallid coloration (?P. christiz
of Gray), while Hodgson’s P. homochrous from the Himalayas has
a larger ear and smaller thumb. Specimens from Tor, Sinai, Ladak,
and Hokkaido appear to be of the latter form. For local English
difference, see Whitaker, Naturalist, 1910, 422.

Dimensions :—The female appears to have slightly the larger wing.

Proportionate lengths:—Foot (without claws), about -48 to -50 of
lower leg; fifth metacarpal, about -95 to -96 of third; lower leg, about
-47 of forearm, and about -35 to -40 of head and body; ear, about -74 to
-77 of head and body, and about -96 of forearm.

Skull :—Greatest length, 16-5; basal length in middle line, 12;
palatal length in middle line, 7; from posterior border of 7° to anterior
border of canine, 5 ; same in lower jaw, 6; greatest breadth at zygoma,
9; posterior breadth, 7; breadth at constriction, 4.

Weight :—Charbonnier sends me the weight of a female as 2
drams 21 grains, or about 5 grammes.

Distinguishing characters:—The enormous ears, united at their |
bases in front, are a unique feature of this species amongst British
-mammals.

VESPERTILIONIDA—PLECOTUS

DIMENSIONS IN MILLIMETRES :—

(approximate)

MALES.
> Ea! 5 a suailies
313 - | ok ss i) ee
2 |$2| a2 $/88/ 4/48/3819) 4 \|ésl 4
Lo! oy — =] i= sO u g te 3 Q a3 a
a Hw | 3 S | 38 3 38 2 E a | +: “3
a} SP) e8) 8 |e (ee) 2 281813 | s eel 2
ee es 8 \Ba| = |Pa] 2 | a | 3 jee
Sa al ye e eine earch (ce
Male,Grimsby, Lincoln-
shire, 10th Sept. 1890
(W. Eagle Clarke) .| 47 35°5 | 18°25| 43°5 38 259* A
Male, Manifold Valley,
North Staffordshire,
16th March 1907 (C.
Oldham) . : -| 47 34°5 | 16 35 19 85 | 36 7 63 31 30 247
Male, Tring, Herts,
dried skin at British
Museum . 51 86 45 |17 9 37°75) 5° 62 81:5 | 30 nc
Male, London, do. 51 38 48 17 9 87°5 6 63 31°5 | 31
Male, Montrose, do. .| 45 36 45 | 16°5 Sb) | sre7o) |r 63 31 30°5 5
Male (dried skin), Co.
Louth, 38rd August
1893(Dublin Museum) s 18 9 38 7°25 | 64 82 30°25
Male (do.), do. 4th
August 1893 (Dublin
Museum) . ° : 17 9 87 7 62°75 | 31°75 | 30
Male (do.) do. 5th
August 1893 (Dublin
Museum) . 33°5 | 17 9 39 75 | 62°5 |\82 81 52 Ac
Male, Achill Island (do. ) ae 16°25 | 8:5 35 6 57°75 | 29°25 | 27°75 5
Male (in the flesh), Co.
Fermanagh (do.) . 18 9 39 a 62°5 | 32 29°5 +.
Average of males $ 3 2 7 f F 4 ,

(approximate) 48°2| 36 4. 41°6|17°3 | 88 | 87:5 6°6 |62°2 | 31°3 | 30 as
a SR Nee NS ee De A Mae a cr 2 I et
Male, young, Devon .| 50 37 ae 44/17 9 45 Se ba on
Male (dried skin), Co.

Louth (hardly ma-

ture) . e 18 9 38°5 7 61 31 29 :
Male, young, Co. An-

ein Oabtn Maseam,

in alcohol). ll 8 32°5 75 | 27:5 | 14:5 |13 Es aé

FEMALES.

Female (in flesh), Mani-

fold Valley, Stafford-

shire, 16th March1907

(C. Oldham) . 46 36 | 17 40°5 | 19 8 40 8 66 83 81 256 ae
Female, Devon, dried

skin, at Brit. Museum| 48 35 42 {17°56 | 9 87 if 61°75 | 30°25 | 29°75
Female, Essex, do. 42 36 45 18 9°5 89°5 | 6°5 | 65 33 82
Female, Herts, do. 49 35 42 18°5 8°5 39 6°5 | 65 82 31°5
Female (1), Montrgae,

do. 46 34 46 18 9°5 40°5 | 6:5 | 68 34 82°5 5
Female (2), do. do. 45 42 |185 | 8 40 6 67 83 31°5
Female (3), do. do. 44 40 |17°5 | 8 38 7 63 81 29°5
Female, North Ber-

wick, 16th Sept. 1894

(Ww. Eagle Clarke) 46 85°51 15 46°5 | 18 10 88°5 | 7:5 50 30 5c 260 |5 grms.
Female (dried skin), Co.

Louth, 5th July 1893 . STG 10 388 7 68 82 380°5 , ‘
Female ‘(do.) do. 5th

August 1893 (Dublin

Museum) . 41 |17 9 |()35°5 | 7 64 82 31°5 s
Female (do.) Co. Ar-

magh, 4th September

1893 (Dublin Museum) 18 9 389 7 63 82 30 5 C
Female (from spirit),

Co. Louth (Dublin

Museum) . : 0 19°75] 8 40°25] 7-5 | 67°65 | 88 81°5 zi 2
Female do. 5 ' 19 8°5 88 75 | 64 31 30°25 : i
Female do. 2 18°5 | 9 88°5 | 85 | 64°56 | 82 80 5
Female, Co. Meath do. ) 19 9 89 8 66°5 |81°5 | 81
Female, Co. Dublin(do.) 20 9 40 7 66°5 | 83 82:25| . A
Female, Co. Louth (do.) 19 9°25} 40 8 65°5 | 82 81

| | | | | — | | | | | | | | -———

Average of aan 45°7 | 35:2

a 42°2)18°3 | 8:8 88°38 | 71 | 65 82 30°9

* Average total expanse of eight individuals, 225.

THE LONG-EARED BAT 199

Few creatures have been more persistently misrepresented
by writers of books than this beautiful and abundant bat.
Although figured by Albin in 1740, and known to Gilbert White
at least as early as 1767, it met with very scanty treatment in
the works of our earlier naturalists, such as Pennant and
Donovan. Bingley alone described its habits at any length,
although, like his contemporaries, he fell into the picturesque
error, supposed to have originated with Edwards, that the
lesser ear may possibly serve as a valve to close the larger
in the sleeping state of the animal. It was not until Messrs
N. H. Alcock and C. B. Moffat’ undertook to write its
natural history that an adequate account of it from first-hand
study became available.

The large and beautiful ears are developed to such an
extraordinary degree as at once to strike the most incurious
observer, and yet probably their actual comparative magnitude
is not fully recognised. But it needed not the unnecessarily
cruel experiments of certain foreign naturalists * to show that
they must have some intimate connection with the animal’s
safe progress by night through the arboreal obstacles amongst
which it delights to wander, but whether as organs, directly
tactile or merely sensory, has not been exactly ascertained.

The ears are usually folded* under the arms during sleep,
especially if the sleep be profound, and this is also the case
during hibernation; the long traguses then stand up, and the
animal has the appearance of having short and slender ears.
Indeed, a person who had not seen it in the act of folding its
ears, would never imagine it to be the same species when they
are fully expanded. At other times they fold outwards and
sideways almost like the horns of a ram, the traguses in this
case reflexed. They frequently move independently in a
curious manner, a bat in an attitude wherein one ear projects
forwards and the other is folded beneath its arm, being a
somewhat remarkable object. In flight the ears are directed
forwards.

A great preponderance of observers have testified that
this bat is one of the most arboreal species, and not, as

1 Trish Naturalist, 1901, 241-251. 2 See Introduction, pages 4o, &c.
3 The folding is well described by A. H. Cocks, Buckingham.

200 VESPERTILIONIDA—PLECOTUS

R. F. Tomes supposed, an inhabitant of the open country.
It frequently seeks its prey, somewhat after the fashion of the
Whiskered Bat, amidst the intricacies of the branches, where
it plucks, snaps, and snatches insects of all kinds from the
leaves.

Among the first to detect it in the very act of thus
hunting was Tomes, who observed a bat of this species actively
engaged around the sprigs of a spindle tree which extended
across a window. ‘The tree was in bloom at the time, and was
surrounded by a cloud of micro-lepidoptera, on which the bat
was feeding at a distance of scarcely four feet from the open
window, so that it was easy to see the whole proceeding, and
to determine with certainty the manner in which the food was
taken. With scarcely an exception the moths were picked
from the leaves while resting there, only one or two being
taken on the wing. While thus occupied, the bat hovered
much after the manner of the kestrel, and the ears were bent
outwards so much as to curl down the sides of the face; they
thus suggested two large cheek-pouches rather than ears, no
part of them appearing at a greater elevation than the crown
of the head. This could be noted very accurately, as the
creature several times hovered scarcely a yard from the face
of the observer at the open window, as if desirous of entering.
This it afterwards did, and after flying round the room a few
times, returned to its feeding.

Similar observations had been made by Jonathan Couch,’
who, in broad daylight, happened to see one taking some-
thing from the surface of a leaf; he imagined that the long
ears might act as organs of quick sensation, as the bat flies
amongst leaves which stand thick on a tree.

Again, Mr G. H. Caton Haigh’ has watched it in a group
of silver-fir trees, which on warm nights in April ‘appeared
full of bats, sometimes flying with the greatest rapidity through
the branches and sometimes hovering like great moths at the
extremity of the twigs. On going underneath the trees the
bats presented a still more curious sight : generally upwards of
a score might be seen moving about in the space of a few feet.
They appeared frequently to come in contact with the branches,

1 Zoologist, 1853, 3937. * Tbid., 1887, 294.

THE LONG-EARED BAT 201

but whether by accident or not” he was unable to ascertain.
One which he shot at this place had a small leaf of the silver-
fir in its mouth.

Nowhere are these bats more in their element than when in
April the flowering sallows are the centre of a crowd of moths
attracted by the blooms. Round these Mr J. Steele Elliot?
has watched them circling, and when moths attracted their
attention, they would steady themselves in their flight, and
with quivering wings (which sometimes gave him the impression
that they were perching), would seize their prey, frequently
from off the bloom itself. So, too, Mr William Jeffery * has
observed them taking moths off the blossoms, “the bat closing
its wings, folding down the ears, and making its meal there
and then without quitting the tree.” The above description
represents the more normal habits of the species, but it is
nothing loth to catch its prey in full and open flight, gracefully
swooping upon the larger moths as Mr Peter Inchbald has
observed.’

The bats are wont to vary the activity of insect-hunting
by retirement with their prey to the seclusion of some barn or
outhouse, where the rejected wings of their victims falling to
the floor often betray their presence* Mr O. V. Aplin®
remarks that these retreats are not sleeping-places but merely
dining-halls, and in the neighbourhood of Banbury the
moths most extensively captured are the buff ermine,° yellow
underwing,’ and silver Y.*. Where no better dining-hall is
available, the bat is perforce content with such lowly sites
as a tree-trunk, where Mr J. Ffolliott Darling’ has observed
one, after catching insects, sit munching a large moth so
vigorously that he could hear the crackling of the moth’s
armour as it disappeared.

Messrs Alcock and Moffat find that in County Wexford
the tree most frequently selected for a hunting-ground is
the ash, amongst the branches of which it may be seen
every evening from May to September. In early May they

1 Zoologist, 1897, 231. 2 Jbid., 1890, 71.

3 Field, 23rd July 1887, 149. 4 Lord Lilford, Zoologist, 1887, 66.
5 Zoologist, 1889, 382. 6 Spilosoma lubricipeda.

1 Triphena. 8 Plusia gamma. 9 Zoologist, 1883, 294.

202 VESPERTILIONIDA—PLECOTUS

have found it also in the oak-woods. But trees are not by
any means an absolute necessity to it, for it will also hunt low
over the ground, probably in search of grass-loving insects,
and it has been hooked in the mouth by the Hon. R. E. Dillon
when fishing with an artificial fly. It appears to be a jealous
little creature, for it has often been seen to attack and repulse
a Pipistrelle whenever one chanced to approach its feeding-
grounds.

‘To observe this bat on the wing,” write these observers,”
“it is a good plan to wait at dusk under some tree whose
foliage is not too dense to be seen through—an Ash is probably
the best that can be selected—and watch for its appearance
amongst the branches overhead. From about thirty-two or
thirty-five minutes after sunset, its figure may, almost any
summer evening, be thus detected against the sky, gliding
and hovering in a stealthy manner among the outer sprays of
the tree. It threads its way with a beautiful facility among
the twigs and leaves, often seeming rather to swim than to fly,
so slight is the visible movement of the wings. Poising, at
times, like a humming-bird, it appears to be picking something
from the leaves; at other times it suddenly plunges into the
middle of a spray, and remains for several seconds clinging to
the twigs, no doubt securing or eating some insect. It is not
uncommon to see one Ash-tree occupied at the same moment
by five or six of these bats—though each comes and departs
by itself—all gliding in the same noiseless and lemurine
fashion among the leaves, and all to the casual bystander
practically invisible. The long ears are often thrown forward
so as to resemble a proboscis, and may be distinctly seen if
the observer is posted immediately below the bat.

‘“When one of these bats leaves a tree, if its object is
merely to pass to another quite near at hand, it darts through
the air with a swift arrowy flight ; but when a longer expedition
is contemplated the mode of quitting the tree is different.
The bat plunges headlong to within an inch or two of the
ground, and then skims away in jerking zigzag fashion—much
as a Nightjar does—over the surface of the field. The swift

1 The “ Detached Olive,” /rishk Naturalist, 1906, 278.
2 [bid., 1901, 242-244.

THE LONG-EARED BAT 203

plunging descent taken on these occasions is very remarkable,
and renders it a difficult matter to keep the animal longer
maysight.. . .

“This bat is decidedly later than the Hairy-armed* in
appearing on the wing. In August, 1900, the time of its
first appearance about a favourite Ash-tree was noted on nine
consecutive evenings, with the following result :—

August 4th. Long-eared Bat first seen 26 minutes after sunset.

” 5th ” ” 36 ” ”
” 6th 99 PP) 36 ” ”
” 7th ” ” 33 ” ”
” 8th ” ” 34 ” ”
» 9th 9 » Bae) tls s
” Toth ” >) ) 33 ” ”
33 I I th 93 ” 37 ” ”
3 I 2th »” ” 34 33 bbl

Thus, in eight evenings out of nine the first appearance was
between thirty-two and thirty-seven minutes after sunset.”

That the flight of the Long-eared Bat extends throughout
the night, was suggested by Tomes, who, as stated above, not
only observed it feeding at three o’clock in the morning, but
heard the shrill chatter of its easily recognised voice over his
head at all hours of the night, even the darkest, in the
open fields and elsewhere. Confirming this statement, Mr
Inchbald? writes that he has watched this species catching
moths between 2 and 3 a.m. in July, and Messrs Alcock and
Moffat state that it has been identified on the wing in County
Wexford in the third week of August, at midnight, at 1.30 A.M.,
and in the morning twilight.* At each of the earlier hours the
identification was effected by watching a bat flying in and out
of a conservatory, where it was possible to identify it with
certainty. The same writers suggest that this bat seeks its
diurnal hiding-places while the morning light is still dim, never
having observed it in view later than forty-five minutes before
sunrise.

The hibernation of the Long-eared Bat appears to be both
more profound and of longer duration than that of some other

Hohe. Leislers Bat, 2 Field, 23rd July 1887, 149.
3 Gordon Dalgliesh met with one on the wing at midnight on Ist December
(Zoologist, 1908, 178).

204 VESPERTILIONID/—PLECOTUS

species, but there is a dearth of precise data, and those avail-
able are contradictory. In Buckinghamshire Mr Cocks has
obtained specimens in nearly every month of the year. In
Wales it has been observed on the wing, on each occasion in
sunshine, on 18th March and 4th November,’ and Mr T. W.
Proger once saw one come out and drink at a fountain in
December?: in County Wexford it has not been detected flying
earlier than the 13th April, while in autumn it ceases to
frequent its favourite ash-trees in mid-October, yet is repeatedly
captured in November. But, as remarked by Messrs Alcock and
Moffat, these observations prove little more than that certain bats
were abroad on the dates stated, for the winter sleep is liable to
be broken by a moderate degree of warmth. They write :—

“On December 21st, 1900, a bat of this species was found
hibernating at Ballyhyland, in a convenient position for obser-
vation zz situ. The sleeping-place was a hole in a Beech-tree,
54 feet above the ground. On the insertion of a finger, the Bat
snarled savagely enough ; but when let alone it soon relapsed
into. an apparently profound slumber.

“On the next day (22nd December) it was gone, having
evidently flown during the night. This desertion must in fair-
ness be ascribed to its having been disturbed, and slightly
alarmed, on the previous afternoon. The remainder of De-
cember, and the first two nights of the ensuing January, were
characterised by cold and frost, and during this period the hole
continued unoccupied.

“The night of January 3rd was mild ; the thermometer until
nearly midnight remained at 46°, and the Pipistrelle was seen
flying. On the morning of the following day the Long-eared
Bat was found to have returned to its hole in the Beech-tree.

“For about three weeks from the above date this Bat was
looked at every day, and often with a lantern during the night.
From January 4th to 11th the observer detected no change of
attitude ; but on some of these days, when the temperature was
as high as 44°, the little creature fidgeted slightly during sleep.
On January 12th, at a temperature of 46°, it became very rest-

1 H. E. Forest in MSS.
2 Paper read before the Biological and Geological Section of the Cardiff
Naturalists’ Society, March 1905, 4. ;

XV.

LONG-EARED Bats, (3 natural size.)

THE LONG-EARED BAT 205

less, and after sunset crept out of its hole and sat for about
three hours in an exposed position on the trunk, with its eyes
open, but its ears still folded back beneath its wings. A high
wind was blowing, which probably prevented the bat taking
flight. On the following night—during a great storm, and at
a temperature of 49 —the animal was again found outside its
hole, this time with one of its ears unfolded and protruded in
front of it, while the other was still tucked below the wing.
Though the bat’s eyes were wide open, and it seemed in
attitude ready to fly, it manifested no concern at having the
lantern held over it. Flight in such a storm would at any rate
have been impossible, and before 9 p.m. it was back in its hole.
During the next seven evenings it was invariably found asleep
in its den at whatever hour visited; and on the night of the
21st of January it was still there at 6 p.m., one and a half hours
after sunset, though the temperature was high (51°) and the
wind light. Next day, however, the hole was empty. A long
spell of cold weather immediately set in, and the bat was not
seen again until March.

“Tt returned for a few days about March 17th, during the
observer’s absence from the locality, and deserted again on the
25th. On April roth it was once more in possession.

“‘ By this period of the spring one might have supposed that
its hibernation was over; but the Bat’s conduct proves the
reverse, for during the next three nights—April roth, 11th, and
12th—it did not fly at all. On the 1oth it was visited with a
lantern at 7.35, 9.5, and 11.5 p.m.; on the 11th, at 8.45 p.m.,,
and on the rath, at 8 and 10.30 p.m. ; and on all these occasions
was seen soundasleep. On the evening of April 13th, however,
it quitted the hole at 7.45 p.m. and did not again return. On
the same night, as already noticed, a bat of this species—
probably not the same individual—was seen flitting round a
willow.

“The respective temperatures for the four nights of April
roth to 13th, taken about 8 p.m., were 43°, 41°, 43°, and 45.

“Other observers have recorded similar facts. Mr C.
Oldham’ informs us that he has found individuals of this

1 See also Mem. and Proc, Manchester Literary and Philosoph. Soc., 49, ii., 9, 1-4,
31st March 1905.

206 VESPERTILIONIDA—PLECOTUS

species in winter in the caves at Alderley Edge, and that these
repeatedly shift their quarters. Many similar instances can be
adduced, so that hibernation in the case of the Long-eared
Bat, at any rate, is not as profound or as unbroken as was at
one time supposed, but is repeatedly interrupted; and, appar-
ently, this is liable to occur whenever the thermometer rises
above 46° F.”

The retreat of the Long-eared Bat is frequently beneath
the roofs of tiled houses in villages or towns, in which places
they may be found in summer—males, females and young—
suspended in clusters’ from the timbers, and during the winter
closely packed between the tiles or in holes, so that unques-
tionably the same haunts are occupied throughout the year.
But, as related above, solitary specimens, the cause of whose
lack of sociability is unknown, may be found in their own
private dens in winter and spring—in one case instanced by
Messrs Alcock and Moffat so late as 2nd May. In fact,
Mr Charles Oldham goes so far as to inform me that in his
experience solitary hibernation is the almost invariable rule,
at least as regards the copper-mine tunnels of Alderley Edge,
Cheshire, and the old lead-workings in the Derbyshire dales ;
whilst in the Henley-on-Thames cavern, Mr A. H. Cocks
and Dr E. A. Wilson?” found five, all resting singly, on 14th
February.

An interesting feature in the animal’s economy is its habit
of appearing for a brief season in summer in colonies or
‘‘swarms” in certain places to which it is at other times a
stranger. These gatherings, to quote again Messrs Alcock
and Moffat, ‘‘seldom remain more than a few weeks. In
two consecutive years, 1898-1899, a space between the wood-
work and wall of the farm-stable at Ballyhyland was occupied
by a swarm .. . during the first fortnight of August,
which disappeared soon after the middle of the month.
In the second year of their visit (1899) particular pains
were taken not to disturb them, but by August 2oth
none remained. In August, 1900, there was a swarm
over the doorway of a neighbouring forge, of which not
a trace could be found in September; and in July, 1901,

1 Compare Goldsmith’s “ Lazy bats in drowsy clusters cling.” 2 Tn lit.

THE LONG-EARED BAT 207

a smaller assemblage took possession for one or two days
only, of a space between two beams in the roof of an
old mill. This last- mentioned swarm probably consisted of
females, with new-born young; for an infant specimen, with
eyes unopened, was found on July rith crawling on the floor
immediately below the crevice in which the adult Bats were at
the time visible.” Since these swarms are so much more fre-
quently noticed in July and August than at other seasons ; and
since there is evidence, as in the instance last quoted, that they
are not composed exclusively of the young of the year, Messrs
Alcock and Moffat are led to infer that the social instincts of
this species are strongest in summer, and that individuals
which have lived solitary lives for the rest of the year become
at that season gregarious. It is probable, however, that this
“swarming” is in some ways comparable to the ‘‘camping-
out” for nursing purposes of the Noctule in summer, and it is
reasonable to suppose that, as in other bats, a seasonal change
of quarters is habitual. Mr Oldham finds the Cheshire and
Derbyshire tunnels tenanted only during winter, and the same
remark, although not of invariable application, appears to be
true also of the Henley cavern.

The cry of this species, although not loud, is shrill and
acute, but it is uttered with more vigour when the animal is
annoyed. Messrs Alcock and Moffat find it in use by the
young when only a few days old, and Mr J. G. Millais graphic-
ally dwells upon the considerable variation to which it can be
subjected, remarking that ‘‘one call is uttered when signalling
to its own species, and apparently the same note is uttered
during a quarrel; but a very different cry—a querulous, long-
drawn, childlike note—is used when it is handled or surprised.”

The Long-eared Bat gives birth to its young in June or
July, the number being usually one,’ but Bingley, on the
authority of a Mr Carlisle, incidentally mentions a female
flying about with two attached to her body. A French
naturalist, Monsieur E. Oliver, informed Monsieur H. Gadeau
de Kerville that the young clamber from one mother to another,
and are by them indiscriminately carried about and nourished,

1R. C. R. Jordan, Zoologist, 1843, 75 ; Alcock and Moffat, /rish Naturalzsé, 1901,
248 (11th July).

208 VESPERTILIONIDAZ—PLECOTUS

but the statement is contrary to the experience of Messrs R.*
Rollinat and E. L. Trouessart, and needs confirmation.’

No bat thrives better in captivity than the Long-eared.
It may, without apparent injury to its health, be removed
from its natural home in mid-winter when completely lethargic ;
in fact when in captivity it entirely ceases to hibernate,’ and,
although at first very shy and refusing food if offered in the
hand, it will readily capture and devour insects left in its cage,
and before long becomes a familiar and trusting companion.
When a number are kept together, they exhibit much happiness,
and are very playful, their gambols being not the less amusing
from their awkwardness. They run over and against each
other, pretending to bite, but never harming their companions
of the same species; though Bell knew some to exhibit a sad
spirit of persecution to an unfortunate Barbastelle which was
placed in the same cage with them. They may readily
be taught to eat from the hand; and one kept by James
Sowerby, when at liberty in the parlour, would fly to anyone
who held up a fly towards it, and pitching, would take the insect
without hesitation. If the insect were held between the lips,
the bat would then settle on its patron’s cheek, and take the fly
with great gentleness from the mouth; and so far was this
familiarity carried, that when one of its friends made a
humming noise with the mouth, in imitation of an insect, the
bat would search about the lips for the promised dainty.

Mr Oldham informs me that a Long-eared Bat which he
had in captivity, when its first grip was insecure, pouched its
wriggling prey in the orthodox manner in the interfemoral
membrane, but used neither foot nor carpus to adjust its hold:
on the other hand, it was frequently its practice to spring off
the table and eat its meals in the air as it flew round the room,
the crunching of its jaws being then distinctly audible. In
nature it probably finds the interfemoral pouch very useful,
and Mr J. R. B. Masefield informed Mr Oldham that, being
close to these bats when hovering over the sallows, he has
seen the tail bent upwards so as to form a receptacle for the
insect as it drops. ‘‘As you know,” he writes, “the sallow

1 Original not seen: quoted by Rollinat and Trouessart, vide supra, p. 36.
2 J. Armitage, Vaturalist, Feb. 1905, 39.

Hae History OF BRITISH BIRDS—continued.

Bi sf icieies of very many others, which were formerly little known,
have been fully elucidated, while, speaking generally, an immense
_imerease in our knowledge on such important subjects as Migra-

tion, Distribution, Habits, Nidification, Plumages, has accrued:

' And lastly, a new and important branch of study has been instituted
_» —namely, the recognition of the various Racial Forms or Sub-
| ‘species exhibited by certain birds in the British Islands, on the
Continent, and elsewhere.

A great advance has also been made towards a more satis-
factory system of classification of the Aves—always a difficult

ii ee subject—and this necessitates departures from the older views.

To bring this Standard Work thoroughly abreast of the most.
recent knowledge in all these departments is the object of the

present work.
.... It should be remarked that while it is not intended to go fully
_. into Synonomy, yet, where changes of nomenclature have been

necessary in order to conform with the Law of Priority—the only
method by which complete uniformity in nomenclature can ulti-

' mately be attained—the names used in the Fourth Edition of

Yarrell’s “British Birds” and in Saunders’ ‘‘Manual,” and the
Trinomial Names of the British Racial Forms, and of those
occurring in Britain as visitors from the Continent, will be quoted,

as will also the Original Name under which the species was

described.
In requesting Mr Eagle Clarke to undertake the duties of

Editorship, the Publishers desire to make it known that they are

acting under the advice of the late Mr Howard Saunders, who

placed all his collected notes for a New Edition at Mr Eagle

Bi Clarke’s disposal for this purpose. That Mr Eagle Clarke is emin-

ently fitted for the work is well known to all who are interested in
ornithological science. Through his investigations of the subject,
and contributions to its literature, he has long been recognised |

| : as one of the foremost authorities on all that relates to British

_ birds. He has studied our native birds in many portions of
the British Islands, and has visited a number of bird-haunts
in various parts of Europe in order to become acquainted
in their Continental homes with the visitants that seek our

shores.
On the important matter of the Migrations performed oe

British Birds, Mr Eagle Clarke’s knowledge is unrivalled—a

material fact, when it is called to mind how little has been said

on this most important subject in any published History of

British Birds.

A new and important feature of the New Work will be a
Coloured Plate of each species. These will be reproduced in the
best style from original drawings specially executed for the work
by Miss Lilian Medland, F.Z.S., an accomplished and well-
known bird artist.

GURNEY & JACKSON
| - PATERNOSTER ROW, LONDON, E.C.

ae HETERE RES

Ta ihe Press, to be published shortly

STUDIES IN BIRD-MIGRATION

BY WILLIAM EAGLE CLARKE, F.R.S. E., F.LS.

Member of the British Association Committee on the Migration of Birds as
Observed on the British and Irish Coasts, and Author of its Final
Reports, 1896-1903, etc.

With Numerous IMustrations and Maps

ITH the exception of the two initial chapters, this i
work is entirely original, being the result. of the

author’s investigations and personal experiences. These have
extended over many years, during which exceptional opportunities

have been enjoyed for acquiring knowledge on Bird-migration as

generally, and its British aspects in particular. —

In 1884 Mr Eagle Clarke was elected a member of the British si

Association Committee on the Migration of Birds as observed
on the British Coasts; and on the completion of that great

enquiry, he was requested by his colleagues to prepare the final a

reports on the results obtained—a difficult and arduous task, oh

which he. accomplished in 1903.

During the preparation of these reports (five in number), ‘Mr cs a
Eagle Clarke became much impressed with the advantages which

were likely to accrue from placing a trained ornithologist at

a number of the most favourably situated observing-stations — a

around our coasts. If this could be done, he believed that some
of the difficulties which the phenomena presented might be

solved, and our knowledge regarding the subject cence:

considerably advanced.

This conviction led him to undertake, by the special permis.

sion of the Elder Brethren of the Trinity House and the Commis-
sioners of Northern Lighthouses, a series of personal investigations
at various light-stations, each of whicn was selected for a special

purpose. In all, Mr Eagle Clarke has resided no fewer than forty-

two weeks in these isolated and remote observatories ; the stations

visited being the Eddystone Lighthouse, the Kentish Knock |.
Lightship (33 miles off the Essex coast), the lighthouses on the |
Flannan Isles and Suleskerry (both lying far out in the Atlantic), i
and the lighthouse at Fair Isle (the ‘(British Heligoland”). He
also visited the Island of Ushant—an important station—and —

Alderney for similar purposes; and spent a month or more in
the autumn of 1910 at St Kilda, for the purpose of carrying
the investigations to the outmost fringe of the British area.
With these unrivalled experiences for its foundations, the
book should not only prove a valuable contribution to the subject of

Bird-Migration, but should occupy a santa essentially its. own in.

ornithological literature.

GURNEY & JACKSON.
PATERNOSTER ROW, LONDON, EC.

Oliver and Boyd, Printers, ‘Palantege, Wit SHES teeth Wit ! Npabaten:

PLECOPUS 209

feeding NMoctue ... all drop immediately the flower or bush is
touched or shaken, and thus the head of the Bat and the
interfemoral pouch form a trap from which the moth cannot
escape. Mr William Evans has observed that in flight the
tail and interfemoral membrane are not stretched out behind,
but curved downwards and forwards at about half their length.

The toilet of Mr Oldham’s captives was no less elaborate
than that of other species, practically the whole body being
licked, scratched, or combed by the aid of one foot, the bat
meanwhile hanging suspended by the other. The long ears
were bent downwards by the wrists, and thoroughly licked
with the tongue.

Mr Oldham’s bats slept either suspended by the toes or
lying prone upon the floor. He noticed’ that of one of them,
after a few days’ confinement, although apparently in good
health, had undergone a change in the colour of the wing-
membranes from light grey to a dull muddy hue.

Neither in confinement nor when at large does this bat
manifest a special regard for any particular kind of food.
Many moths, besides those mentioned above by Mr Aplin, the
remains of beetles, dipterous flies, in particular the blue-bottle,’
have been found in its rejectamenta by Professor G. H. Car-
penter, while Mr J. E. Harting* mentions various moths of
the genus Zenzocampa, and Mr Jeffery the large spotted-winged
crane-fly. Mr Caton Haigh believes that it also takes
small caterpillars; Mr J. D. Batten has fed it with grass-
hoppers in confinement; Mr Oldham with moths* and meal-
worms, and Mr J. Armitage’ with flies, moths, pupz, small shreds
of beef, and mealworms. He has known one to eat twenty
of the latter at a single meal.

Like other species, this bat occasionally appears on the
wing in broad daylight,° and its activity on such occasions or
in mid-winter must be ascribed to the causes already recited.
That the species, like most others, is not a creature of unvary-
ing habits, is shown by the capture of one by a lighthouse

l Zoologist, 1899, 472. 2 Calliphora erythrocephala. 3 Zoologist, 1889, 245.

4 Gonoptera libatrix and Scotosia dubitata. 5 Naturalist, 1905, 39.

6 F. W. L. Ross, Zoologist, 1845, 1158; Couch, Journ. cit, 1853, 3942; Lilford,
Journ. cit., 1887, 66; Oldham, Journ. czt., 1890, 349 ; R. Newstead in Coward and
Oldham, 1895, 166; W. Evans; Forrest in MS., etc., etc.

210 VESPERTILIONIDA:—BARBASTELLA

keeper at the Tearaght, one of the most desolate and in-
accessible of all the Irish islets, some eight miles off the coast
of Kerry, at so late a date as 4th November 1891*; and Mr
J. F. Fortune, light-keeper at Aranmore, off the Donegal
coast, wrote me that one made its appearance at his lantern
on 24th September 1899.

The Long-eared Bat, once clearly seen either at rest or on
the wing, can never be mistaken for any other species. The
ears, although approached at a great distance by those of
Bechstein’s Bat, are unique amongst British mammals.’

GENUS BARBASTELLA.

1821. BARBASTELLA, J. E. Gray, London Medical Reposttory, xv.. 300; based on
Vespertilio barbastellus of Schreber.

1829. BARBASTELLUS, Jakob Kaup, System der Europaischen Thierwell, i., 96 ; based
on Vespertilio barbastellus of Schreber.

1839. SyNoTUS, A. Graf von Keyserling and J. H. Blasius, Wiegmann’s Archiv fir
Naturgeschichte, i., 305 ; based on Vesfertilio barbastellus of Schreber.

Synonymy :—The earliest generic name known to have been
applied to the Barbastelle is certainly Gardastella of Gray, a
name revived by Mr Miller in 1897 (Axx. and Mag. Nat. Fiist.,
Oct. 1897, 384-385). It clearly antedates both Larbastellus and
Synotus, the former of which has to meet the additional objec-
tion that, as pointed out by Dobson (Catalogue of Chiroptera,
175, footnote), it was in the first instance applied to a species
of Nyctophilus.

This small and distinct genus includes only one British
species, the Barbastelle, 2. darbastellus.

The distribution is imperfectly known, but extends at least
from Skandinavia to Abyssinia, Egypt (Heuglin, Rezse zx Nora-
ost Afrtka, ii., 30, 1877), and Arabia, where the local form was
named deucomelas by Riippell (Atlas zur der Reise 1m nord-
lichen Afrika, 73, pl. xxviii. B, 1826); and from Great Britain
to Assam, where the representative is Hodgson’s darvjeling-
esis, a common mountain bat of the Himalayas, ascending

1 R. M. Barrington, 7he Migration of Birds, 284; R. H. Porter, London, 1900.

2 Since the above article was written, Oldfield Thomas has described P. wardz,
a large pale-coloured form, from Leh, Ladak, Kashmir (Aum. and Mag. Nat. Hist.,
I9{I, 209); and P. ariel, allied to P. waraz but of much darker colour, from Ta-tsien-
lu, Szechwan, Western China (Aéds¢. Zool. Soc. (London), 7th February 1911, 3).

BARBASTELLA Zang

to at least 8000 feet (Blanford). It is said to have a larger ear
and longer forearm (44.5 mm.) than éarbaste//us. Satunin’s
sub-species caspica (forearm about 43 mm.), described from
Transcaucasia, is not available for examination. The genus is
unknown in America.

The generic characters are moderate size; ears not large, their
bases united at their inner margins on the forehead, their outer
margins encircling the eyes and terminating between them and
the upper lips ; a well-developed tragus
with an attenuated tip.

The short muzzle (Plate XIV., Figs. % :

2 and 3) has the upper surface naked and
nearly flat dorsally, the sides glandular
and tumid so as to form a raised border :
the nostril is terminal, with a broad shal-

low groove running across the upper lip. Ga)

The moderately broad wing (Plate
XVII., Fig. 2) arises from the base of
the toes. The calcar extends about
half-way from foot to tail. The post-
calcarial lobe is narrow and _ incon- 09 ?0,
spicuous. The tail is nearly as long as
the head and body ; its tip projects about
3 mm. from the interfemoral membrane,
which is ample, supported by long lower
legs, and extends triangularly to some
distance behind the feeble feet.

The skull (Fig. 8, No. 5, p. 101), AS Fic, 21.—DIAGRAM OF AR-
exemplified by B. darbastellus, is weak, RANGEMENT OF TEETH IN
with prominent rounded brain-case; “" ah ores

. : (1) Upper and (2) Lower Jaw.
broad, somewhat concave facial region;
weak, flattened zygomata; weak or no cranial crests ; and
moderately developed auditory bullez.

There are thirty-four teeth (Figs. 21 and 22) arranged as—

LS)

.2—2 I—I 2—2 1 3=3 3
Z b c if pm } quo

3-3 I— 2-2

The upper incisors are oblique, the outer nore small, the inner
large and bifid ; the upper canine is provided with small anterior

212 VESPERTILIONIDAE—BARBASTELLA

and posterior cusps at its base. The anterior upper premolar
is minute, and, lying in the inner angle between the canine
and posterior premolar, is invisible externally. The anterior
lower premolar is about half as high and broad as the posterior.

THE BARBASTELLE.
BARBASTELLA BARBASTELLUS (Schreber).

1760. LA BARBASTELLE, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s
Histoire Naturelle, viii., 119 and 130-131, 135-137, pl. xix., fig. 2; described from
France.

1774. VESPERTILIO BARBASTELLUS, J. C. D. von Schreber, Die Sdugthiere, i., pl. lv.,
168, evidently naming Daubenton’s Za Barbastelle; James Sowerby, British
Miscellany, pl. v., 9, 1804-6; George Montagu, 7rans. Linnean Soc., London, 1x.,
171, 1808 ; Bingley ; Pennant ; Jenyns ; Clermont.

1785. VESPERTILIO BARBASTELLA, P. Boddaert, Elemchus Animalium, 1., 69.

1821. BARBASTELLA BARBASTELLUS, J. E. Gray, London Medical Repository, xv., 309 ;
Miller ; Thomas ; Collett ; Johnston ; Millais ; Trouessart (1910).

1829. BARBASTELLUS BARBASTELLUS, Jakob Kaup, System der Europdischen Thier-
welt, i., 96.

1836. PLECOTUS BARBASTELLUS, Georges Cuvier, Ze Régne Animal, i., 74; Fleming.

1837. BARBASTELLUS DAUBENTONH, Thomas Bell, British Quadrupeds (ed. 1), 63;
(ed. 2), 81, 1874, renaming Daubenton’s Za Barbastelle; MacGillivray.

1838. BARBASTELLUS COMMUNIS, J. E. Gray, M/ag. Zool. and Bot. ii., 495 ; evidently
renaming V. darbastellus of Schreber.

1839. SYNOTUS BARBASTELLUS, A. Graf von Keyserling and J. H. Blasius, Wieg-
mann’s Archiv fir Naturgeschichte, i.. 305; Blasius ; Fatio; Dobson ; Flower and
Lydekker ; Lydekker ; Cabrera.

1900. BARBASTELLA BARBASTELLA, Méhely Lajos, J/onographia Chiropterorum
flungariaeé, 131 and 326, pl. v.

Barbastelle from the French daréastelle and Italian darbastello, from
the Latin darba, ze, a beard. There do not appear to be any local
names for this little-known bat, either in England, France, or Germany.

Distribution :—Barbastelle-like bats range throughout a great part
of the boreal and transitional regions of Europe and Asia from
southern Skandinavia and middle Russia to middle Spain, and south
Italy (Monticelli), the Crimea, and probably North Africa (Dobson) ;
and from England to Tiflis (Satunin). It is not known if Bb. darba-
stellus is preferably a mountain bat; as is B. davjelingensis, to which it
gives way somewhere in the East, perhaps in central and eastern Asia,
or, according to Satunin, in Transcaspia. Fatio, however, supposed so,
having found it at 5000 feet in the valley of Urseren, at the foot of Saint
Gothard, Switzerland. Robert took one at 3200 feet at Caterillo, Haute
Garonne.

THETBARBASTELLE 213

In Britain this bat is unknown from Scotland, Ireland, or the Isle of
Man, and in England the records of its occurrence are mainly from the
south. Sowerby’s first British specimen was taken by Peete at
Dartford, Kent, before 1804 (see synonymy), and Bell was indebted
to Waring for another from a chalk cavern at Chiselhurst, in the
same county; its occurrence there is corroborated by Millais, who,
however, believes that it is now rare; and it is also reported,
although not recently, from Cornwall (Clark). Montagu (see
synonymy) mentioned specimens from both Milton and Kingsbridge,
in Devon, thus completing its range across the south of England.
Montagu’s record has, indeed, been questioned by Gray (Zoological
Journal, ii., 110, 1826) on the ground that the individual in the British
Museum marked by him darbastellus, is undoubtedly referable to A/yotis
mystacinus. But this error must have arisen in the labelling and not in
identification, since Montagu’s description is very full and correct, and,
moreover, recent observers confirm the occurrence of the bat in Devon.
It was, for instance, reported as “scarce” at Teignmouth by Jordan
(Zoologist, 1843, 75), while a specimen from Torquay is, as Hollis
informs me, in the Exeter Museum (see de Hiigel, Journ. czt., 1869,
1768). Thus between Cornwall and Kent the sprinkling of occurrences
perhaps indicates a continuous distribution. There are several records
for Surrey (Mitford, Journ. cz¢., 1860, 6953; Millais, 42; Thorburn, edd,
19th July, 1902, 142; Dalgliesh, Zoologzst, 1907, 299; Bucknill and
Murray); and Sussex (Borrer, Zoologzst, 1874, 4128-29; Brazenor,
Journ. cit., 1887, 152; Millais, 42); for Hampshire, one or two (Hart and
Kelsall); for Wiltshire, one at Salisbury (Blackmore, Zoologist, 1860,
1558); for Somerset, a small colony in the roof of Wells Cathedral
(Berry in Millais, 43), and a breeding haunt in Mendip Caves (Lewis,
Journ. cit., 1906, 69). From Dorset the bat is reported by Borrer once
( Journ. cit., 1869, 4128-29), and by Dale as “not common” ( /ourn. cit.,
1887, 234). North of the Thames Valley it is known from Essex,
where Doubleday found it not uncommon in Epping Forest (/ourn. cit,
1843, 67), and where Laver, writing of the whole county, thinks that,
although “not so rare as it is usually believed to be,” it cannot be called
common; from Middlesex and Oxford Borrer received one each from
Hornsey and South Weston respectively (/ouwrn. czt., 1874, 4128); in Hert-
ford, Oldham found one on Berkhampstead Common ( /ourn. cit., 1908,
391); for Buckingham, Cocks knows of two occurrences, and one in
Berkshire; while from Gloucester two specimens came under the notice of
Charbonnier (zz “zt, also see Journ. czt., 1892, 329; Witchell, Journ. ciz.,
1892, 356): and others were known to Tomes from the Warwick border.
Alfred Newton met with it in Suffolk; the elder Gurney, in Norfolk
( Journ. ctt., 1857, 5420; see also for Suffolk, Rope, Journ. cit., 1891, 347) ;
and in Norfolk Southwell describes it as not rare and generally distributed.

VEZ

214 VESPERTILIONIDAZ—BARBASTELLA

Jenyns knew of its occurrence in Cambridge’ and Northampton ; Lord
Lilford in the latter county (/ourn. cit, 1894, 187), and in Hunting-
don (/ourn. cit., 1894, 395); Steele Elliott in Bedford; and Montagu
Browne in Leicester (Journ. cit, 1885, 215). Tomes found it not very
rare, although by no means abundant, in Warwick. Upton-on-Severn,
Arrow near Alcester, and Weston-on-Avon are mentioned as Wor-
cester localities (Jenkinson, /ourn. ct¢., 1857, 5590, see also Tomes). North
of the Wash, Caton Haigh’s description (/ourn. czt., 1887, 144) of a “ rather
large dark-coloured Bat,” frequently observed “flying low over grass-
land, so low as only just to clear the higher stalks of grass .. .” mov-
ing “heavily with slow flaps of its wings, and . . . generally seen in the
neighbourhood of trees”—suggests this species as a regular member of
the Lincoln fauna. A male in the British Museum, labelled Cheshire
but without further data (cited as a female by Dobson, Catalogue
of Chiroptera, 177), and two examples captured near Carlisle, are the
only known specimens from the north of England. The latter are
stated by Macpherson to have been skinned for T. C. Heysham; after
his death they were purchased by Bond, in whose house Macpherson
examined them in March 1886. Since August 1889 they have been in
the possession of Harting, who writes me that the details given above
are confirmed by Bond’s own handwriting on the labels.

The species was not known from Wales until, on 13th June 1904, Rev.
D. E. Owen sent two for identification to Forrest from Llanelwedd, near
Builth, on the Brecknock border of Radnor: they proved to be members
of a definite and well-established colony (Forrest, Zoologzst, 1904, 262 ;
Trans. Caradoc and Severn Valley Field Club, iv., 1, 52-54, 1905, Jan. 1906).

The interpretation of the records is a matter of some difficulty, but
the fact that the bat is well-known, although not abundant, in Essex,
Norfolk, and Warwick; that it has definite colonies in Somerset and
Worcester; and has been taken at intervals within the area of every
county south of the Wash and east of the Dee, suggests that further
observation will show it to be widely distributed in small numbers, at
least within the above limits, with outposts beyond them, as in mid-
Wales. The record from Carlisle may point to a gradual extension of
range or to wandering habits: Macpherson attributed it, although with-
out definite grounds, to migration. There is nothing in the bat’s
European distribution to suggest any inherent improbability of its
occurrence in the north of England or even in Scotland.

Distribution in time :—This bat is not known as a fossil.

The breeding habits are unknown.

Description :—The general form and appearance of this remarkable
bat are typically those of its genus.

The ear (Fig. 2, No. 9, p. 7) when stretched forward reaches slightly

' A record recently confirmed by A. Whitaker, who examined two caught at
Faversham on 17th May 1910 (Waturalis/, 1910, 424).

THE BARBASTELLE 215

beyond the tip of the muzzle!; it is nearly as broad as long and irregu-
larly four-sided; the inner edge is reflected, forming a longitudinal
groove just within the margin; the outer and superior angle is
prominent, rounded, and turned back ; immediately beneath this, on the
external margin, is a rather deep notch, from which five or six slight
transverse folds extend about half-way across the conch; the anterior
inner angle unites with its fellow immediately behind the muzzle. The
tragus has a length more than half that of the ear; it is of an irregular
lanceolate or semicordate form, with one conspicuous and two or three
smaller protuberances near the base of the superiorly concave outer
margin; the terminal third is straight and the apex rounded; the inner
margin is faintly convex.

In the wing (Plate XVII., Fig. 2) the most remarkable features are
the small foot and large lower leg, the proportions of which are unique
amongst British vespertilionid bats. The tail, longest digit, as well as
the third and fifth metacarpals, are, as compared with their forearm,
exceptionally long. The fifth metacarpal is distinctly shorter than the
third. The result is a wing amply, but not exceptionally broad, and a
large interfemoral membrane.

The fur is soft and long. The upper side of the muzzle is naked,
but the tumid cheek is thickly clothed with hair, forming a kind of
moustache and almost concealing the eye. The wing on the upper
surface is furred thickly to about as far as a line passing from the knee
to the centre of the humerus, and on the interfemoral membrane to a
line running from slightly above the knee-joint to a point one-third
of the distance along the tail. Beneath, the fur extends somewhat
beyond a line joining elbow and knee, and a sprinkling of hairs runs
along the membrane behind the forearm to the carpus. On the outer
side of the ear a thick column of fur clothes the central and lower
portions, but the outer margin is bare; the inner margin is fringed for
almost its whole length by a band of hairs. On the inner surface the
conch is sprinkled with hairs: a thick band of hair runs along the line
of the anterior folding almost to the apex, becoming thinner as it ascends.

The colour above is dusky, almost black, most of the hairs
having whitish or yellowish tips, thus causing a frosted appear-
ance, which may be most conspicuous in the region between the
shoulders. Beneath, the colours are similar, but the light tips are more
numerous, whiter and longer, and their influence predominates increas-
ingly towards the furred portion of the wings and interfemoral
membrane, which are lightest, the dusky basal portions of the hairs
being there absent. The wing, ear, foot, and nose are dusky.

1 At least in fresh specimens. Dobson’s statement that, “laid forwards the tips
extend to a point midway between the eye and the end of the muzzle” (Catalogue of
Chiroptera, 176), may have been based on stale specimens distorted in alcohol.

216 VESPERTILIONIDA-—BARBASTELLA

The nearly full-grown young are said to differ from the adults in
having the underside darker.

The skull (Fig. 8, No. 5, p. 101) and teeth (Fig. 22) are typical of
the genus.

No material exists on which to base a

study of the variation of this species.

Millais mentions a golden-brown form, and

Tomes two immature examples, one of

which was perfectly white ; the other (now

in the British Museum) had the body pure

white, but the head, part of the neck, and

lower dorsal region normal; in both, the

wings were almost white. A third, taken

at Alcester, Warwick, had the fur of the

under parts, from root to tip, strongly

tinged with purplish red or rose colour,

which was very conspicuous when the animal

was fresh, but faded considerably after pre-

servation. There is much variation in the

extent of the frosting of the hairs, some

| specimens being very dark; Lord Lilford

mentions a dark brown male taken on

22nd March, and a very grey female

oe ee oe taken on 4th September (Zoologzst, 1894,

stella barbastellus (enlarged and 187 and 395).

diagrammatic). As regards size, the forearm in eight

British specimens measured by me aver-

ages I-5 mm. less than in thirteen from the continent of Europe, but

the difference is too minute and the series too small to afford basis for
a definite conclusion.

Proportionate lengths :—F oot, without claws, about -30 of lower leg ;
fifth metacarpal, about -93 of third ; lower leg, about -49 of forearm, and
about -38 of head and body.

Skull :—Greatest length, 15; basal length in middle line, 10-6;
palatal length in middle line, 5; from posterior border of 7* to anterior
border of canine, 5 ; greatest breadth at zygoma, 7-7 ; posterior breadth,
7-5; breadth between orbits, 5 ; breadth at constriction, 3-5.

The weight is given by Fleming as 100 grains = 6-5 grammes.

Distinguishing characters :—The Barbastelle cannot be mistaken for
any other European bat. Apart from its dark colour, in no others
—except the Long-eared, the immense ears of which are very different—
are the internal margins of the ears joined together. The latter char-
acter is evident even in the unfledged young.

PLATE XVII.

WINGs (4 natural size and diagrammatic) OF

1. Plecotus auritus. 2. Barbastella barbastellus.
3. Rhinolophus hipposideros.

PLATE XVIII.

HEADS (natural size) OF

I, 2, and 3. AAinolophus hipposideros (front, three-quarter, and side views).
4. Rhinolophus ferrum-equinum.

THE BARBASTELLE 217

DIMENSIONS IN MILLIMETRES :—

2 2 a ® |
ean | ss g 2 | 2 ey hee
2 | & | ae 2/2/22) = oleate
ee. | Beh ail ete oe | eee Weel eee
| 2 | oO] ‘a Be Wissue g = ” at eae)
3 = BB | BS aie iS) S o s ° »'S
co} oD HS 5 ge? & “e &0 8 3S me |
Fy bi $ 4 a 3 ao! illy Soin ea
® ir) 2 us} = © & o
isa ay oD 3 g = a o |
3 eu) = o |
=a |
|
| |—_ |__| |__|
Male, Arrow Lodge, War- |
wickshire, 7th October
1848 (R. F. Tomes’s col-
lection), measured from
dried skin at British
Museum. Ss a a 42 | 18 ae leo 4 65 34 32 bee]
Male, Oundle, Northants, approx. |
5th April 1900 (Lord |
Lilford), do. . BOM) 2s be 48* | 18 Ui 38 | 4 68 35 33 |
Male, Milton, Hants,
7th April 1900 (Rev. J.
H. Kelsall), do. . 52* | 16* 6* | 44* | 18:5] 6:5 | 36 3:25 | 66 34 31-5 | 260
Male, Llanelwedd, Rad-
norshire, 6th June |
1904 (H. E. Forrest),
do. 43 19 6°5 | 38 4:5 68°5 85°5 | 88°5 | 254*
Male (received from E.
A. Wilson), 5th Oct.
1906, measured in
Heshy |: 3 5 si) 46) LO T5 oe 19*5 |) 7:5 1-88 5 63°5 35°5 | 34
Male, Cheshire (G. E.
Dobson), measured
from spirit in a
Museum . 45 14°5 6°5 40 19 6 39 5 67°5 35 31
Average of males \ : ‘ P ‘ i” ‘ ame c oe ; AS
(nmin er ice a .. | 48:4] 186] 6°7 | 37:6] 4:3 66°4 | 34:8 | 82:5
Female, Elton, Hunts,
4th Sept. 1894 (Lord
Lilford),measured from
dried skin . 60* So te oe 18 6°5 | 36 4 63°5 35 31
Female, Epping Forest
(H. Doubleday), mea-
sured from alcohol . 44 | 16 8 43 19 6 37 4 65 82 30
Female, Herts, received
in flesh from C. Old-
ham, 16th Sept. 1908 . | 51 | 13°5 9 54 | 20 7 41 4 67 37 35 268

* Collector’s measurements,

This curious and interesting bat was first described by
Daubenton in 1760, and subsequently by Buffon in his great
work. Kuhl, notwithstanding the extent of his researches and
his exertions to procure all the bats of Germany, failed to
obtain one of this species. It was first detected as a native
of Great Britain by Peete, who sent one to Sowerby from the
powder mills at Dartford in Kent. It was known to Montagu
from Devonshire, and subsequently obtained a place in all works
on British Mammals except that of Donovan.

Although quite unmistakable in the hand, many of the
earlier published notices of the habits of the Barbastelle
were clearly incorrect, and the meagre accounts available

218 VESPERTILIONIDA-—BARBASTELLA

made any attempt to write its natural history difficult until the
Rev. D. E. Owen and Mr G. E. Bullen sent Mr H. E. Forrest
an account of a colony in the porch of Llanelwedd Church,
Radnorshire. All that could be said for certain was that, of
the few recorded specimens which had been captured, the
majority were discovered either in houses, or resting on or
under the bark of trees! Others owed their detection to their
appearance on the wing in broad daylight,’ a proclivity which
Mr Eardley Hall, who, according to Mr J. G. Millais, has had
good opportunities for observing this species, believes to be the
rule and not the exception.

The first notice of its natural history came from the pen of
Bell, who received from Dr Waring a specimen taken during
a very hard frost in the latter end of December, in a large
chalk-cavern excavated at the bottom of a shaft seventy
feet deep, at Chiselhurst, in Kent. ‘In this cavern,” wrote
Bell, ‘during very severe frosts, several species of Bats are
found to retreat ; and on this occasion, with the Barbastelle was
received a specimen of the” Whiskered, three of Natterer’s,
and several of the Long-eared Bat. ‘These little prisoners,
when brought into a warm room, soon began to exhibit signs
of vivacity ; and the Barbastelle, with the others, fed readily on
small bits of meat, and drank water. He was a timid animal,
and did not evince the slightest disposition to become familiar ;
he would take his food, however, with his companions, and
was accustomed to rest with them in a cluster, at the top of
the box in which they were placed. The Barbastelle certainly
became torpid more readily than any of the others, and more
completely so; but when awake, evinced extreme restlessness,
and was incessantly biting with great violence at the wires of
his box. When suffered to fly about the room, he flew very
low, and less actively than the others under similar circum-
stances; and he was fond of lying before the fire on the
hearth rug, where he appeared quite to luxuriate in the warmth.
Whilst the Long-eared Bats evinced much attachment to each
other, and became very familiar with me, the Barbastelle

1 See A. Newton, Zoologist, 1857, 5421; Lord Lilford, zézd@., 1894, 187.
* See H. P. Blackmore, zézd., 1869, 1558 ; C. W. Brazenor, zd¢d., 1887, 152; A. H.
Cocks, Jenyns, etc.

ERE BARBASTELLE 219

remained sullen and apart; until at length I found that he was
an object of persecution on the part of his more active
companions, one of whom | detected in the act of giving him
a severe bite on the back of the neck. This occasioned his
immediate removal to another box; but this sharp discipline
probably hastened his death, which took place about a week
afterwards, though he continued to eat till the day before he
died.”

Nothing further of any importance was written on the
Barbastelle until R. F. Tomes met with it in Warwickshire,
and remarked’: that “if in a twilight stroll about midsummer
a person finds himself in a close proximity with a Bat of
somewhat thick and clumsy form, but of rather small size,
whose flight is so desultory that it appears to be flapping
lazily about, hither and thither, seemingly without purpose,
and intruding so closely that the flutter of its wings may be
heard, and even the cool air thrown by their movement felt
upon the cheek, it may with almost certainty be recognised as
the Barbastelle. Although there is no English Bat which
resembles the Barbastelle in its mode of flight, yet in choice of
situation there are several. Where the Whiskered Bat and
Pipistrelle are seen, the Barbastelle may be seen also, but
having been once observed, it will, probably, be useless to
make search again at the same place. Equally uncertain is
its diurnal retreat; most likely not the same place for long
together, as we have found it in places where it could not have
rested the day previously. A crevice in a wall or tree, the
spaces between the rafters and tiles of a cowshed, the timber
over a sawpit, the thatch of a shed in a brickyard, or behind
a cottage window-shutter, are suitable places of repose for the
Barbastelle, in all which situations we have met with it, and
always alone.”

In these few words Tomes seems to have hit off with
considerable accuracy the ‘flapping,’ almost aimless flight of
what would appear to be, at least occasionally, a wandering
species, living a life exempt from the routine which seems to
be so marked a feature in the daily existence of some others.
His estimate of its solitary habits not only when in its place

1 In) Bell.

220 VESPERTILIONIDAE—BARBASTELLA

of repose during the day, but during its evening flight, proves
to have been less correct, although supported by Dr Henry
Laver; for, besides the evidence of the Radnorshire colony, we
have Mr Millais’s statement, on the authority of Mr A. G. Berry,
that a small flock frequents the roof of Wells Cathedral. Again,
Mr J. H. Jenkinson’ met with a party of six or seven in their
hiding-place in the month of June; and in the spring on another
occasion discovered a mixed assemblage, consisting of a
Natterer’s Bat, a Barbastelle, and two or three Pipistrelles.
True, he does not say definitely that the bats were in the
latter case actually together, although undoubtedly inhabiting
adjacent crevices, but Lord Lilford expressly states” that he
“found this curious-looking little animal in great abundance ina
ruined monastery in Arragon, at the foot of the Pyrenees, .. .
and in smaller numbers in a similar locality at Pétes, in the pro-
vince of Santander,” so that there must be seasons, perhaps those
of procreation, when it courts the society of its own species.

It thus appears that the Barbastelle is no less sociable * and
gregarious, perhaps even more so, than the Long-eared and
other species. Some of those which Mr Owen found hanging
from a beam in the church porch at Llanelwedd, had formed
themselves into a small and compact ball. A week later
a bunch of ten were found occupying the same position,
so that it is evident that at least in summer this species
has rightly gained its reputation for wandering. When
disturbed early in the morning or in the afternoon, the Barbas-
telles were in a condition of deep torpidity, markedly distinct
from the state of the Long-eared Bat, but their actions were
very different at eventime, when if disturbed they exhibited
great agility in attempting to avoid capture.

In Radnorshire the Barbastelle is an early-flier, leaving its
diurnal resting-place at sunset, and preceding by some little
time the Pipistrelle, the Long-eared Bat, and even the Noctule.
Mr Owen has often seen the swifts and Barbastelles flying
about his rectory for nearly an hour before the former retired
for the night. Their habits and flight vary with the weather.

1 Zoologist, 1857, 5590. 2 Tbid., 1887, 66-67.

3 Two Cambridgeshire specimens were found hanging side by side on the wall of
an old shed much frequented by Long-eared Bats. They were “almost, but not quite,
touching one another” (Arthur Whitaker, Vaturalist, 1910, 424).

THE BARBASTELLE 221

When it is fine, they fly high and do not enter the church, prob-
ably sheltering themselves amongst the branches of the yew-
trees, but rain or wind drives them to seek the shelter of the
porch or tower, and to fly round the building with a slower and
more ponderous flight.

Mr Forrest is inclined to believe that this bat is silent
while on the wing: at all events he has never heard it cry
in flight. When captured, however, ‘“‘some of them made
the usual metallic squeaking noise, whilst one or two of them
made a noise very much like that of a huge bee held in a
handkerchief.” They repeated these two distinct cries when
recaptured after having been released in a room.

In captivity their demeanour varied, one readily consenting
to eat chopped steak, another refusing food, but none lived
for more than a few days. Mr Forrest remarks that they flew
against the window, but not with the blind dash of an
imprisoned bird. When walking or crawling they “straddled”
their limbs more than other bats, and their movements were slower.
He noticed that the head was habitually held downwards, so that
it was difficult to obtain a view of the curiously wizened face.

More recently Mr Charles Oldham! has kept a bat of
this species alive in captivity for a few days, and remarks
that it had two notes, a querulous squeal of the usual bat-
like kind and a peculiar subdued buzzing. In a room its
flight was slow and fluttering, and it generally preferred
the upper part of the room, but occasionally flew close to
the ground amongst the legs of chairs and tables. In flight its
tail was extended and only slightly decurved; the legs were
held wide apart, so that the interfemoral membrane looked very
large from beneath. Asa rule, but not always, it turned in the
air like a Natterer’s or Horseshoe Bat, before alighting. After
some preliminary difficulties it was induced to accept meal-
worms, but seemed to fear cockroaches. It preferred to eat on
the wing, but was not observed to call in the assistance of the
interfemoral pouch, the nature of its food probably making such
a course unnecessary. It was adroit in picking houseflies off
the ceiling.

Four facts are thus demonstrated for the first time:

1 Zoologist, 1908, 391-392.

222 VESPERTILIONIDA!A—NYCTERIS

that this bat is distinctly gregarious; that its flight begins
very early in the evening; that it is exceptionally active in
alighting and passing obstacles; and that it knows how to
catch its prey while at rest. The latter fact was indeed to
be expected from its peculiar facial attachments, but it must
be remembered that the behaviour of this, as of any other
species, may vary so much with the occasion, the individual, or
the season, that no hard or fast rule can or should be drawn
from the result of merely one or two observations.

According to Messrs R. Rollinat and E. L. Trouessart,
the Barbastelle is in France a hardy species, and that is what
might be expected of one whose range includes portions of
Skandinavia. Its hibernatory slumber is probably not deep,
and it has been captured abroad on a fine New Year’s Day.’
For the rest I can only find notes that it has been observed in
activity on various dates from 3rd March * to 3rd October.

The breeding habits are unknown, but the late Thomas
Southwell showed me a letter from S. Bligh to Henry Stevenson,
wherein the newly born young, picked up at Framlingham,
Suffolk, in 1865, are said to have been easily recognised by
their ears, a character which at all ages is infallible for the
distinction of this species.

[THE HOARY BAT.
NYCTERIS CINEREUS (Beauvois).*

The record* by John Wolley, of the occurrence in South
Ronaldshay, Orkney, about September 1847, of a specimen
of the Hoary Bat of North America, has long been relegated
to obscure corners of British faunal works on the ground that,
as indeed Wolley himself supposed, the animal must have been
conveyed across the Atlantic accidentally in a ship. The bat
was caught by some potato-diggers, and is stated to have been
kept alive for some weeks.

1 See H. P. Blackmore, Zoologist, 1869, 1558.

* R. Mitford, zd¢d., 1860, 6953 ; Lilford, z¢¢., 1894, 395 ; Owen, per Forrest.

3 For use of WVycéteris (replacing Las¢urus), see Miller, Proc. Biol. Soc., Wash-
ington, 17th April 1909, go.

* Under the name Vespertilio pruinosus. See Zoologist, 1849, 2343; 1850,
2695-96, 2813-14, for an accurate description, excepting only that the number of the

teeth is given incorrectly.

THE HOARY BA 223

The discovery by Dr C. Hart Merriam’ that the Hoary
Bat annually performs a migration’ no less extensive than that
of many birds seems to put a very different complexion
upon the matter, and makes Wolley’s record one of the most
interesting ever contributed to British mammalogy. This bat
is known to breed only in the boreal zones of North America,’
but in winter it occurs at least as far south as the most
southern border of the United States. It has been taken on
the Bermudas, showing that it is able to cross a strip of ocean
having at its narrowest extent a width of over 600 statute
miles, that is between the Islands and Cape Hatteras, which
lies very little north of due west of the Bermudas. It is,
however, much more likely that the bats commence their oceanic
passage at some point much further to the north, such as
Cape Cod, the distance from which to the Bermudas is
about 700 statute miles. They certainly pass Cape Cod on
migration, since Mr Gerrit S. Miller, jun.,* found them
there on passage in the autumns of 1890 and 1891, in
the former year from 26th August to 2nd September, in
the latter from 25th August to 12th September. The
distance from Cape Cod to the Orkneys is about six times
that to the Bermudas, but if a bat started its flight
from Nova Scotia or Newfoundland, the distance might
be reduced, in the latter case to four and a half times. It
seems, therefore, to enter the bounds of possibility that a
creature which habitually passes over 600 or 700 miles of
ocean should occasionally be blown the whole way across the
Atlantic, especially if its passage happened to coincide with
some great hurricane. The season at which Wolley’s bat was
found at South Ronaldshay corresponds with that of the
autumnal migration of its kind past Cape Cod, and it would
almost seem that the species deserves to be included in the
same category as the various American birds which from
time to time strike our coasts. But the case of a bat is

! Trans. Royal Soc. (Canada), 1887, iv., 85-87.

2 Some observations of A. H. Howell (Proc. Biol. Soc., Washington, xxi., 35-38, 23rd
Jan. 1908) suggest that, when migrating, bats may fly at high elevations, and by day ;
but further information is needed, and our knowledge of the subject is still very slight.

3 Miller, Morth American Fauna, No. 13, 112, 1897.
4 Sczence, New Series, v., 542-543, April 2, 1897.

224 VESPERTILIONIDAE—NYCTERIS

different from that of a bird. Even where the latter are
concerned, the proportion of American visitors detected
may be infinitesimal as compared with those which escape
recognition ; but the chances that a bat should be discovered,
or even that it should survive its passage, are remote.

Unfortunately, a new complication has arisen, and the
matter is not as clear as it might be. Wolley’s original
specimen is said to be preserved in the University Museum
of Zoology at Cambridge, and I am indebted to the late
Professor Alfred Newton for bringing the fact under my
notice. There is evidently, however, something wrong, since
the specimen which now does duty for Wolley’s belongs to a
species’ having its habitat in the Sandwich Islands, and does
not agree with Wolley’s very clear description. After so
many years it is impossible to explain the mystery. The
Cambridge specimen was dried and not skinned, and its condi-
tion would therefore suit the hypothesis that it was taken in
the Orkneys. But if so, it is an entirely unlikely species to
have arrived there alive, and doubtless never did so on its
own wings. On the other hand, it is known, as the Professor
informed me, that Wolley’s brother, George, returned from
the Sandwich Islands in 1856, so that it is possible that he
brought this specimen with him, and that it was accidentally
substituted for the bat taken at South Ronaldshay. This
does not look an improbable solution, for John Wolley only
wrote the label in 1858, and, being evidently doubtful of his
information, he was careful to mark it with a note of interroga-
tion. But if so, it is difficult to understand why George
Wolley should have brought home an unskinned bat all the
way from the Sandwich Islands, and it is to be feared that
the matter cannot now be decided with certainty.

Two other American bats, both easily recognisable species,
perform extensive migrations, and are at least as likely to
be blown over to this country as the present species, viz., the
Red Bat,? and the Silver-haired Bat,* but of these the latter,
at least, is stated to have powers of flight inferior to those of the
Hoary Bat. |

1 Nycteris semota (Allen). 2 N. borealis (Miiller).
3 Lasionycterts noctivagans (Le Conte).

RHINOLOPHID/E.
HORSESHOE-NOSED BATS.

Characters:—These bats have a conspicuous nose-leaf; no
tragus; a short tail and moderately developed interfemoral
membrane; premaxillz rudimentary, suspended from the nasal
cartilages and only partially bridging the gap between the two
upper incisors, which are quite rudimentary and widely separ-
ated ; nasal bones much expanded.

GENUS RHINOLOPHUS.

1798. LES RHINOLOPHES, Georges Cuvier, Zadleau élémentaire de ? Histoire Natur-
elle des Animaux, 105.

1799. RHINOLOPHUS, Le C. Lacepéde, Tadleaux ... des Mammiferes, etc., 15, No.
73; based on Rhinolophus ferrum-equinum.

1816. PHYLLORHINA, W. E. Leach, Systematic Catalogue of the Specimens of The
Indigenous Mammalia and Birds that are Preserved in The British Museum, etc.
(London), 1; based, without description, on Phyllorhina minuta of Leach= Vespertilio
minutus of Montagu, the Small Leaf-nose, from Torquay, Devon, presented by
G. Montagu ; see Blanford, Proc. Zool. Soc. (London), 6th Dec. 1887, 637-638.

1836. RHINOCREPIS, F. L. P. Gervais, Dictionnaire Pittoresqgue a’Histoire Naturelle,
iv., 2, 617, quoting from Cuvier and Geoffroy, Magazine Encyclopédigue, vi., 1795,
but the name does not occur in the paper referred to, nor, so far as is known, prior
to 1799.

1847. Aquias, J. E. Gray, Proc. Zool. Soc. (London), 9th March, 15-16; Ann. and
Mag. Nat. Hist. June 1847, 408 ; based on Rhinolophus luctus of Temminck, and
R. trifoliatus of Temminck.

1866. PHYLLOTIS, J. E. Gray, Proc. Zool, Soc. (London), 13th Feb., 81; based on
Phyllotis philippensis of Waterhouse, i.e., Khinolophus philippinensis ; preoccupied
by PHYLLOTIS of Waterhouse, 1837, a genus of JZuride.

1866. SPEORIFERRA, J. E. Gray, Proc. Zool. Soc. (London), 13th Feb., 82 ; based on
Speoriferra vulgaris, i.e., Rhinolophus vulgaris of Horsfield.

1866. C@LOPHYLLUS, W. Peters, Proc. Zool. Soc. (London), 22nd Nov., 427 ; based
on Rhinolophus celophyllus of Peters.

1887, PSEUDORHINOLOPHUS, Max Schlosser, Bettriige zur Paliontologie Osterreich-

225 U

226 RHINOLOPHIDAZ=—RHINOLOPHUS

Ungarns, etc., VI., i. and 11, 55, 61, pl. ii., fig. 42; based on Rhinolophus antiquus
of Filhol and other extinct species.

1901. EURYALE, Paul Matschie, Sztzungs-Berichte der Gesellschaft naturforschender
Freunde zu Berlin, 225 ; based on &. méhelyt of Matschie.

The genus is the only one of its family. There are two
British species, 2. ferrum-equinum and R. hipposideros. Out-
side Britain R. euryale of Blasius, occurring throughout the
Mediterranean Region to central France, has the lip with three
grooves, the sella with its sides parallel anteriorly and the
posterior connecting process projecting and triangular; it has
many sub-forms, such as /. méhelyz of Matschie and the
Spanish R. carfetanus of Cabrera. Peters’s 7. dZaszz of south-
eastern Europe has peculiar nose-leaf and lower incisors.

Distribution:—The seventy-one species and numerous sub-
species recognised by Andersen (Aun. and Mag. Nat. First.,
Dec. 1905, 648-662; Proc. Zool. Soc. (London), 17th October
1905, 75-145), range from Ireland through southern Europe
and Asia to Ceylon and Malaysia, the Celebes, Philippines,
Formosa, Japan, the Liu-kius, and Australia. They nowhere
attain to high altitudes, and are probably not found farther
north than the shores of the
Baltic (2. 4zppostderos) or
southern Korea (2. ferrum-
eguinum nippon and &. cor-
nutus). Andersen refers exist-
ing species to six “types,” all
Y, of which he believes to have
“4, had their origin in the Oriental

: = Region.
Ge ss = The genus is characterised
My Ais by eee or small size
M(({\ysse20))))

highly developed nose - leaf
with triangular posterior por-
Bie ane Nine °F tion, and deeply notched ear

with well-developed anti-

AU( WS

a, sella. d, division between the
4, lancet. nostrils. tracus.
Cc, ¢, eyes. é, ¢, horseshoe. §

The nose-leaf consists of
three distinct sections, of which the anterior, shaped like a
horseshoe, rests flatly on the muzzle, and encloses the nostrils

RHINOLOPHUS 227

within its arms. The median section or sella commences behind
or between the nostrils, and from a recumbent anterior portion
rises posteriorly into an erect, horn-like, connecting process,
standing vertically upon the face. From the sella there runs
backwards a process connecting it with the posterior section or
lancet, which is tongue-shaped and terminates posteriorly in a
single point.

The wing, which arises from the ankle, is very large
and broad, the fifth metacarpal being longer than the
third: the area between digits two and three is large, and the
base of the thumb is included in the antebrachial membrane.
The second digit has one joint only, the metacarpal; the third
three, the metacarpal and two phalanges. The first toe has
two joints, the remainder three each.

In addition to the single pair of pectoral mammez the
females are provided with two nipple-shaped prominences in
front of the pubis, to which the young
attach themselves by their teeth.

These false nipples produce no secre- (apy
tion, and are not in communication

with any gland: they are attached by

fibrous tissue to the pubis (see Rol-

linat and Trouessart, JA7ém. Soc. Zool.

de France, x., 1897, 124).

The teeth (Figs. 25 and 26) are
thirty-two, arranged as—

-I>1 Pa Z— 2 a4
eee a | PT 5 —— = 32. 4
2-2 I-1 3-3 3

The lower incisors are tricuspid.
The upper incisors, although minute,
are always present in the always
ossified premaxille. The anterior gic. 24—Fronr View oF In-
upper premolar is minute, and often gape ast Couns oF Wl
crowded out of the tooth-line exter-
nally by the large canine and posterior premolar. The central
lower premolar is often in similar position between the neigh-
bouring teeth. The molars are well-developed and their cusps
acutely W-shaped. The milk teeth undergo absorption in the
embryo.

228 RHINOLOPHIDAZ—RHINOLOPHUS

THE GREATER HORSESHOE BAT.

RHINOLOPHUS FERRUM-EQUINUM (Schreber).
RHINOLOPHUS FERRUM-EQUINUM INSULANUS (Barrett-Hamilton),

1760. LE FER-A-CHEVAL, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s
Histoire Naturelle, viii., 131-133, 135, 137, pl. xx., figs. I and 2; also, A/ém. de
2? Acad. Roy. des Sct., 377, 382, pl. 15, fig. 4, 1759, published 1765; described from
France.

1774. VESPERTILIO FERRUM-EQUINUM (in part), J. C. D. von. Schreber, Déze
Saugthiere, i., pl. \xii., upper of two figs., 174, 190; evidently naming Daubenton’s
Le Fer-a-cheval; Montagu, Transactions Linnean Soc. (London), 1x., 162, 1808 ;
Bingley ; Donovan ; Clermont.

1776. VESPERTILIO EQUINUS (in part), P. L. S. Miiller, Matursystems Supplements
und Register Band, 20, renaming Schreber’s Vespertilio ferrum-equinum.

1779. VESPERTILIO PERSPICILLATUS, J. F. Blumenbachs, Handbuch der Natur-
geschichte, 75 (part); included the leaf-nosed bats of Europe and South America.

1785. VESPERTILIO UNGULA (in part), P. Boddaert, E/enchus Animalium, i., 71,
renaming Schreber’s V. ferrum-equinum.

1788. VESPERTILIO FERRUM-EQUINUM, a major, J. F. Gmelin, Systema Naturae, i., 50.

1792. VESPERTILIO FERRUM EQUINUM MAJOR, Robert Kerr, Animal Kingdom, 99
(not V. molossus major of Kerr, of. cét., 97) ; described from France.

1797. NOCTILIO FERRUM-EQUINUM, J. M. Bechstein, Der Zoologe, 1, 1.-viil., 62.

1798. VESPERTILIO HIPPOCREPIS (in part), F. von P. Schrank, /auna Boitca, 1., 64.

1803. RHINOLOPHUS MAJOR, Etienne Geoffroy, Catalogue Mammifécres Mus. @ Hist.
Nat. (Paris), 56; described from Burgundy.

1807. VESPERTILIO FERRUM EQUINUM MAJUS, Lazare Spallanzani, Rapports de Pair
avec les étres organisds, etc., edited by Jean Senebier, ii., 77 (Geneva) ; described
from Nonantola, near Modena, or from Scandiano, Italy.

1813. RHINOLOPHUS UNI-HASTATUS, Isidore Geoffroy, Aum. du Mus. ad’ Hist. Nat,
xx., 257; described from Europe.

1817. RHINOLOPHUS FERRUM-EQUINUM, W. E. Leach, Zoological Miscellany, iii.,
2; Harting, Zoologist, 1887, 2, pl. i. (G. E. Lodge), ef Dlurimorum auctorum.

1829. RHINOLOPHUS UNIFER, Jakop Kaup, System der Europaische n. Thierwellt,
i., 104 ; without description, hence a zomen nudum.

1862-63. RHINOLOPHUS FERRUM-EQUINUM, var. GERMANICUS et var. ITALICUS, Carl
Koch, /Jahrbiicher des Vereins fiir Naturkunde tm Herzogthum (Nassau), xviii.,
522-523; described from Europe, north and south of the Alps respectively, but
evidently in error (see Andersen, Proc. Zool. Soc. (London), 16th May 1905, 113
footnote).

1872. RHINOLOPHUS LIBANOTICUS, CONCHIFER, e¢ RUFESCENS, “ EHhrbg. et Lichst.
Mots,” W. Peters, Monatsberichte der Koniglich Preussischen Acad. der Wissen-
schaften (Berlin), 1871, 310; without description, hence a momen nudum.

1885. RHINOLOPHUS UNIHASTATUS var. HOMORODALMASIENSIS, Daday Jeno, Orvos
—Természettudomdnyi Ertesité, etc., x. (3), 274; described from Homorédd—
Almas, Hungary, in error (see Mé¢hely, MWonographia Chiropterorum Hungaria,
120-122 and 323, 1900).

1887. RHINOLOPHUS FERRUM EQUINUM, var. HOMORODENSIS, Daday Jeno, Erte-
kezések A Természettudomdnyok Korébol, Budapest, xvi., No. 7, 13, pl., figs. 5 and 6,
1886 ; renaming his var. HOMORODALMASIENSIS.

THE GREATER HORSESHOE BAT 229

1905. RHINOLOPHUS FERRUM-EQUINUM TYPICUS, Knud Andersen, Proc. Zool. Soc.
(London), 17th October 1905, 113.

1910. RHINOLOPHUS FERRUM-EQUINUM INSULANUS, G. E. H. Barrett-Hamilton,
Ann. and Mag. Nat. Hist., March, 292; described from Cheddar, Somersetshire,
England (type in British Museum) ; Trouessart, 1910.

Le Fer-d-cheval of the French; ade grosse Hufetsennase of the
Germans.

Synonymy :—The synonymy of our two British Horseshoe Bats
was very much confused until the end of the eighteenth century, when
they were first separated by J. F. Gmelin, and later by Bechstein. Kuhl
thought them one species, and although Daubenton was well aware of
the distinctions between them, there is nothing in his descriptions to
indicate that he had formed any more distinct opinion. (See also under
R. hipposideros.)

Distribution :—X. ferrum-equinum ranges from sea-level to 7000 feet
in the Himalayas; from the south of England through central Europe,
the Mediterranean region—exclusive of Egypt—and the Himalayas to
south China and Japan. Its habitat includes at least the islands
Cyprus, Sicily, and Minorca. It is, with P. pzpzstrellus, the commonest
bat of Normandy (de Kerville, Maturaliste, 15th October 1891, 239);
and in the west, south-west, and centre of France it is also abundant,
but less so in the east (Rollinat and Trouessart, J/ém. Soc. Zool. de
France, x., 1897, 114). It is common in Guernsey (Bunting).

In Britain it is confined to the south, mainly the south-west of
England and the south and west of Wales, and is not known from
Scotland or Ireland (see below). It was discovered by Latham, who
found it in the powder mills of Dartford) Kent (Pennant, Arztish
Zoology, ed. iv., 129, pl. xiv., 1776), and later was encountered by Mon-
tagu in considerable numbers in company with X. Aipposzderos, in Kent’s
Hole, the well-known cavern near Torquay, Devon. More recent records
have connected it with Hampshire, the Isle of Wight, Gloucester,
Somerset, and Cornwall, in some parts of which counties it occurs
abundantly. So far as is known, it has never been reported from
Sussex (neglecting the indefinite record of one discovered on the sail
of a fishing-boat on Brighton beach, see Ruskin Butterfield); and it is
quite unknown to Laver in Essex (Bell’s statement, repeated in Cassell’s
Natural History, i., 283, and by Millais, that it occurs at Colchester,
being evidently an error, based, as Laver informs me, on. a statement
made by the botanist Curtis to Yarrell). Kelsall (Zoo/ogzst, 1884, 483)
states that one was shot on the Berkshire side of the river at Oxford
(lat. 51°, 46’) about 1875; 15 miles further south Noble took two in
his cavern in the same county, near Henley-on-Thames, on 14th
March 1909 (see Cocks, /ourn. cit., 1909, 154); the first-mentioned

1 Not, as stated erroneously, at Dartmouth, Devon (see Rowe).
EZ

230 RHINOLOPHIDAE—RHINOLOPHUS

Berkshire locality is almost the most northerly recorded for England,
but must just give place to Whitchurch (lat. 51°, 51’), near Ross, on the
Gloucester border of Hereford, whence, as Forrest informs me, Rosse
Butterfield received six from a cave in November 1910. At almost the
same latitude as Henley, De Winton observed it (vwzde Millais) in the
Zoological Society’s Gardens, Regent’s Park, London. Next come two
Gloucester localities, viz. Westbury-on-Trym and the Cathedral at
Bristol (Rudge King and Charbonnier). Amongst the other localities
from which it has been taken or recorded within its range are :—In
Somerset, Clifton (Eagle Clarke), Hampton Road, near Bath, Wells
Cathedral, and Clevedon (Charbonnier), the Mendip Caves (particularly
abundant, fide Laver; also Lewis, Zoologzst, 1906, 69) and Taunton
(Millais); in Wiltshire, Great Cheverell (/owrn. cit, 1910, 307); in
Surrey, Godstone (rare, Bucknill and Murray); in Kent, Rochester
Cathedral and Margate (Bell), Maidstone and Dover (Millais), and
Canterbury (abundant, Borrer, Journ. cit., 1874, 4129); in Hampshire,
Portsmouth (Borrer, /oc. cz¢.), and Christchurch (Trevor-Battye and
Lascelles); in Wight, Sandown, Farringford, Bonchurch, and the
Undercliff (the commonest large bat of the latter, see More, Bury,
Millais, Wadham); in Dorset, Tomson Manor (Salter, Zoologzst, 1865,
9835); in Devon, Teignmouth (Jordan, /Journ. czt., 1843, 75), Torquay
(De Hiigel, Journ. czt., 1869, 1768), Hooe, Plymouth, and Plympton
(Rowe) ; in Cornwall, several localities (see Clark); a Yorkshire record
in the Journ. cit., 1884, 483, Editor’s note, is an error (see Roebuck,
Journ. cit., 1885, 24).

In Wales the bat has a haunt at the Mumbles, near Swansea,
Glamorgan (F7e/d, 1st Jan. 1881, 24), and has been found by Tracy in
the old Wogan Cavern, near Pembroke Castle, Pembroke (Kelsall,
Zoologist, 1887, 89); Grabham also took a specimen from a disused mine
near Penmaenpool, Merioneth (Caton Haigh, Zoologist, 1896, 433);
and Forrest received another, shot by Rawlings on 5th February 1896,
at Barmouth, in the same county (both in lat. 52° 30’).

M‘Coy suggested that a bat of “great size” captured in Co. West-
meath, Ireland, prior to 1845, and characterised by the possession
of “a large-pointed appendage on its nose,” was of this species, but the
evidence of its identity was entirely hearsay, and it may well have been
an example of some exotic form. M‘Coy appears to have mentioned
the occurrence in a paper read before the Dublin Natural History
Society, on 12th Feb. 1845, reported in Saunders’s News-letter of the
same date, and quoted by J. R. Kinahan before the same Society on Ist
April 1859 (see Dublin Was. Hist. Review, vi., 383, 1859).

The almost entirely south-western distribution of 2. ferrum-equinum
(south of 52°) in England, together with its extension northwards on
the Welsh coast to a point nearly a degree north of its English range,

THE GREATER HORSESHOE BAT 231

is one of the most striking facts in the distribution of British
mammals.

Distribution in time :—Bones of this bat from Kent’s Hole, near
Torquay, appear to be of very different ages, some resembling in con-
dition those of mammoth, hyzna, and rhinoceros found with them,
while others are quite recent (see Owen, British Fossil Mammals,
etc, 15-16).

Breeding season, and number of young:—Very little information
on these points has been published by British naturalists. In Germany
the young are stated to be two, sometimes only one (Kuhl), but Rollinat
and Trouessart, whose interesting observations (Comptes rendus Soc.
Grol. (Paris), 26th January 1895, 53-54, and 6th July 1895, 534-536; also
Mém. Soc. Zool. de trance, x., 1897, 114-134) have been strangely over-
looked, find that in France the latter number is never exceeded.

As in the Vespertilionide, copulation takes place in the autumn (but
see under RX. hipposideros), the earliest date at which spermatozoa have
been found in the uterus of a female being 5th October. After copula-
tion the spermatozoa become imbedded in a “bouchon” or body formed
of mucus hardened after secretion within the vagina. In October the
females are extremely fat, the mammary glands show no trace of milk,
the ovaries are small and pale rose-coloured, the uterus normal with the
exception of a slight enlargement of the right horn, but the vagina full
of a thick white substance, rich in spermatozoa. By November this has
hardened to form the “ bouchon.”

Ovulation and fertilisation take place in March, or more often,
early in April (Duval, Journ. de 7 Anat. et de la Phystol., xxxi., March-
April 1895, 93-160), the exact date depending on the conclusion of the
hibernatory sleep. A female kept alive by Coward for nine weeks was
found to contain a small embryo on 28th February, but hibernation had
been interrupted during the whole period of captivity.

The fertilised ovum invariably takes up its position in the right
horn of the uterus. Subsequently the “bouchon,” now of no further
use, is expelled from the vagina, tearing the vulva in its passage.

By the end of April the right horn of the uterus containing the
embryo is as large as a pea or a very small hazel nut, and the first
young one was observed clinging to its mother on the 20th June.
The period of gestation is, therefore, ten to twelve weeks. By 3rd
July most of the young are born, but late births take place up to
about the 20th; the mothers have the pectoral mammary glands fully
developed, but the nipples are shorter than the false ones of the pubis:
the uterus is in atrophy, but the right horn still forms a large pocket.

Young of the first year, of both sexes, although associating with the
breeding females, prove on dissection to be not sufficiently developed to
breed; the young females pair in their second autumn; the internal

232 RHINOLOPHIDAE—RHINOLOPHUS

organs are very similar to those of the older females, but the right horn
of the uterus, although slightly longer, is no larger than the left, and the
false nipples are much smaller. One of these second-year females had
not yet paired on 27th October. The “bouchon” is never formed in the
females of the first year, and comparatively late in those of the second
year.

Contrary to the views of Carl Vogt (Association francaise pour
Pavancement des Sciences, x.; Compte rendu, 655, Algiers, 1881),
Rollinat and Trouessart find no evidence that the young females pair
in the spring of their second year ; they are sure that they do not do so
until their second autumn ; some, perhaps, not till the third.

The males undergo a season of rut, their organs being well
developed from September to May. The urethral gland, a structure
peculiar to these bats and of doubtful function, is then of enormous
size, although in the non-breeding season inconspicuous. Since the
organs remain functional not only during autumn—the pairing season—
but also throughout the winter and a part of the spring, it seems
natural to suppose that copulation takes place during the latter periods.
This suggestion, advanced by Robin (Bull. de la Soc. Phil. de Parts (7),
v., 26th March 1881, 88), and at first supported by Rollinat and Troues-
sart, was later rejected by them on the ground of the existence of so great
a number of virgin females just under a year old in April and May.
The enormous reserve of apparently wasted spermatozoa is in this
case a puzzling factor, but the authors suggest that it may be
reabsorbed and utilised in accordance with the Brown-Séquard theory
as an internal secretion. The presence of the “bouchon” in fertilised
females must prevent subsequent copulation in winter and spring.

Young males, although possessing spermatozoa in their second
autumn, probably do not breed until their third, when they are a little
over two years old.

In the urethra of many males there occurs a sort of soft “ bouchon,”
the function and origin of which is uncertain. It is possible that in
bats, as in some rodents,! the secretion of the prostate gland has power
to coagulate the mucous secretion of the vagina, and the beginning of
coagulation may take place in the urethra of the male.

Description :—The general appearance of this bat is typical of its
genus. In size it about equals the Noctule or Serotine.

The head (Plate XVIII., Fig. 4) is long, the occiput large and
rounded, the muzzle very tumid and furnished with long, stiff hairs.
The mouth opens straight and wide. The lower lip has a single groove
and occasionally traces of two others, one on either side.

The ear (Fig. 2, No. 10, p. 7) is rather large, broad at the base, and

1L. Camus and E. Gley, Comptes rendus Soc. Biol. (Paris), 18th July 1896,
787.

THE GREATER HORSESHOE ‘BAYT 233

when laid forward reaches well beyond the nostril ; the apex is abruptly
and distinctly pointed, and folded a little outwards ; the outer margin,
which is concave just below the tip, then slightly convex, is notched at
its base and produced along the side of the face towards the corner of the
mouth, forming in front of the auditory opening a rounded lobe which
appears to be capable of closing the ear; from the outer margin extend
ten or twelve sulci, which run tranversely to the middle of the ear and
are crossed at right angles by a conspicuous blood-vessel; the inner
margin is convex throughout, the lower two-thirds folded inwards so as
to form a kind of flap.

In the nose-leaf the horseshoe is broad, parallel to the muzzle, but not
nearly covering it, emarginate in front, and formed of three concentric
elevations, of which the thickened inner one forms the walls of a
depression in which are situated the nostrils. The sella, which is small
and fiddle-shaped, commences anteriorly as a prominent process, its
exposed surface being broadest anteriorly, and presenting a deep cup,
divided equally by a low vertical septum; about its middle this process
is somewhat contracted laterally, but posteriorly it again expands to
nearly its former breadth and terminates in a short but conspicuous
horn-like structure—the connecting process—rather acutely pointed
at either end of its greatest length, which lies parallel to that of the
body. At its base this horn is connected with the tongue-like lancet.
This latter is distinctly longer than broad, and, from a breadth anteriorly
almost equal to that of the horseshoe, tapers up the forehead to a
point. Anteriorly a mesial ridge and two series of transverse and
more or less horizontal septa divide the lancet into six irregularly
paired cells, the posterior pair being much the least prominent.

The thumbs and feet are moderately stout, the latter armed with
strong claws.

The wing (Plate IX., Fig. 3) arises from a point slightly above
the ankle; the small interfemoral membrane projects slightly in the
middle posteriorly; the tip of the tail is free. The antebrachial
membrane is ample.

The second phalanx of the third digit is very long, and at least
equals one and a half times the length of its metacarpal, which is
considerably reduced in length, and the same proportion also holds
good in the fourth digit. The fifth is slightly but decidedly the longest
of the three. The proportions of the three are represented by the
following figures :—

Forearm. ae eee a shane
1000 644 724 743

The false nipples are very long, transversely wrinkled, and flattened ;
their colour is yellowish or brownish,

234 RHINOLOPHIDA—RHINOLOPHUS

The fur is thick and soft, and in its texture suggestive of wool
rather than fur; it thickly clothes the whole face except the nose-leaf ;
fur, ears, and nose-leaf uniting to almost bury the eyes. On the hinder
surface of the ear the fur ceases at a point about one-third from the
base, but a few small hairs are sprinkled on the outer margin, and on
the inner a more conspicuous band reaches nearly to the apex.
The reflected portion of the anterior margin is somewhat thickly
covered with hair, which becomes thinner on the interior surface. On
the upper surface of the wing the fur extends approximately to a line
drawn from a point about one-third of the distance from the head of
the humerus to the middle of the femur; a band of hair sometimes
runs along the antebrachial membrane nearly to the elbow. The fur
does not extend directly on to the interfemoral membrane, which how-
ever is sparsely furred to some extent on either side of the tail. On
the under surface of the wing the distribution of the fur is somewhat
. similar, but there is no antebrachial band, a sprinkling extends from
the body almost to the elbow and knee, and the interfemoral membrane
is much more sparsely haired.

The nose-leaf is well supplied with bristles; in front there runs also
a prominent single fringe, and a double row along the upper lip.

The colour in adults is uniform above, some shade between “mars
brown” and “ Prout’s brown,” the bases of the hairs “ecru drab”; the
underside is “wood brown” washed with “ecru drab” on the throat
and neck ; there is no distinct line of demarcation. The wing, ear, and
foot are deep brown; the false nipples light yellowish or brownish.

Immature specimens are conspicuously greyer, the general colour of
the upper side (due to the hair-tips) being “drab” washed with brown
from the shoulders backwards, where the dark tips are longer and
almost conceal the whitish “ecru drab” bases. Beneath, the throat and
neck are “ecru drab,” the belly very light greyish drab. The young
females of the first year, although almost as large as the adults, may
also be distinguished by the absence of the false nipples, which do not
appear until the second autumn, and even then are not of the full size
(Rollinat and Trouessart).

The juvenile pelage persists at least until early February (Coward),
when the epiphyses of the metacarpals have undergone ossification,
although the teeth are still unworn. Exactly how and when the change
to the mature pelage is affected is unknown.

In the young at birth the eyes are closed and the underside is naked
and purple. There is a covering of short ash-coloured hairs on the upper
surface, which grow rapidly and assume a dusky colour. The mem-
branes are clear grey; the ears turned backwards; the nose-leaf as in
the adult. The eyes do not open until after ten days (Rollinat and
Trouessart).

THE GREATER HORSESHOE BAT 235

In the skull (Fig. 25, No. 1) the palatal bridge is long, very nearly
a third of the length of the maxillar tooth-row, a little more or less, but
never so short asa quarter. The auditory bullz are small, and the basi-
occipital is not specially narrowed between them.

The anterior upper and central lower premolars are either wanting
or minute, and, if present, are squeezed out of the tooth-row externally.
The upper canine and posterior premolar
overlap, the latter being external. The
central and posterior lower premolars are
in contact.

Variation :— Andersen finds a tendency
to three grooves on the lower lip, and in
the lancet to hastate, sometimes almost to
cuneate, shape.

Geographical variation manifests itself
in size, especially in that of the forearm, 1
the teeth and nose-leaf, and in the length
of the tail, Andersen (see above, p. 226)
recognises six sub-species, three eastern
and three western. The former are Tem-
minck’s zzppon of Japan and south China,
in which the size is moderate, the horseshoe
broad,and the teethrather small; Hodgson’s
tragatus, of the Himalayas, with large size
and broad horseshoe; and Andersen’s
regulus of the north-western Himalayas,
with narrow horseshoe and highly de-
veloped dentition. In all these the size
is comparatively large, the tail short, and
the two extra mental grooves often present.
The three western sub-species are compara-
tively small, with longer tails, narrower
skulls, and as a rule only one mental
groove. Andersen’s proximus of Gilgit is
short-tailed as compared with the typical
form. The latter ranges through southern
and central Europe, exclusive of the Fig. 6. DrdeeAu OF Anawee
Iberian Peninsula, to Transcaspia and the MENT OF TEETH IN Rhinolophus
Euphrates Valley, and has the forearm = ferrum-eqguinum.
usually 57 mm. and upwards. Cabrera’s (1) Upper and (2) Lower Jaw.
obscurus, of the Iberian Peninsula, the
Balearic Islands, and Algeria, is said to differ only in having the forearm
rarely exceeding 57 mm.; since there are doubts as to the validity of
that variety, it is necessary to distinguish the British form, which has

236 RHINOLOPHIDA:—RHINOLOPHUS

undoubtedly a shorter forearm than the type, although with a skull of
about equal size. It may be known as X. ferrum-equinum insulanus
(Ann. Mag. Nat. Hist., March, 1910, 292). Its forearm is even shorter
than that of obscurus, with which form it cannot be phylogenetically
identical, even if apparently indistinguishable.

DIMENSIONS IN MILLIMETRES :—

a pir = C 3S
~? nm D =] Se . A
Sle aa ee) a ea,
2 > & ba COE stared | ned ealieSeah areees ot ates a eae
oc 8 Pos = fa se Bh olverete fos a Go| Re
ra ~ aes s © $a o 3 n = S os
s $ See a Ee Ors too tale en g 3s | ee
uo] © 4 a8 S: ialp ss ica g = S 2 2
e oo op A Zz 5 fo) 2 o 7
g 4 2 tp 5 * = re S a g
q & orn = a i
See G
Male, Cheddar, Somer-

set (T. A. Coward)

5th Jan. 1907 . 66 23 15x8°75 | 38 24°5 | 10 54°5 | 6 85 36°75 | 39
Male,do. . 61 22 14:25 x 8°5 82°5 | 25 11 53°5 | 6 82 34 38°5
Male, do., 7th “Jan.

Eagle Clarke) . . | 68°5 | 22 Ss 33 8 - BEE! Be ap Pb w= 1/880

Average of males | ; Pare : Pa Re ee : : : ;
(approximate) f 68°2 | 22 14-7 x81 32 | 24°5 | 11:2 53°6 |6°1]| 83°4 | 85°4 | 38°9 | 334°6
Female, Cheddar, Som-

erset (T. A. povare):

5th Jan. 1907 . 64 22 14°5x8 38 24°5 | 10°5 55 6 84°5 | 35°5 | 39°56
Female, do., pe Jan. |

1907 . A 69 22 138 33 26 13 54 iE 88 37 42 340
Female, do. . : . | 63 22 15x80 184 25) aldsbie 14-2 Teo) Si buono iad 855
Female, do. . 5 +68: 21°5 14x8 82 25 13 54° | 7 84 37 41 350
Female, do. . 5 ail (65 23 14x8 34 26 11 ay al erg 85 34 39 355

Average of females | 90: j ‘ ¥ : <eill es : : .
(approximate) | 62°S| 221) 141xs 33-2 | 26°83 |11°8 | 54:5 | 6:9] 85:8 | 86-2 | 40-5 | 350

* Andersen gives forearm of thirteen specimens of both sexes as averaging 55:4, with extremes
of 58 and 53°8.

The female is, as in many other bats, slightly the larger sex (com-
pare also the weights on next page).

The dimensions of a newly-born young male were as follow, 27 zuches :
—head and body, 1-56; tail, -62; ear, -37; longest digit, -93; forearm,
-87; lower leg, -68; expanse, 4-56 (Whitaker in MS.). Young of ten
days age have an expanse of 190 to 210 mm. (Rollinat and Trouessart).

Proportionate lengths :—Foot, with claws, about -45 to -46 of lower
leg ; fifth metacarpal, about 1-09 to 1-11 of third ; lower leg, about -45 to
-46 of forearm, about -38 of head and body, and about -76 of tail.

Skull :—Greatest length, 24; basal length in middle line, 19:1 to
19-4; palatal length in middle line, 8; from posterior border of 7° to

THE GREATER HORSESHOE. BAT “2377

anterior border of canine, 8-8; greatest breadth at’ zygoma, 12-2;
breadth at constriction, 2-6.
Weight in grammes :—Fourteen males averaged 20-4, the extremes
being 23-6 and 17-3: one female weighed 28-1 (Charbonnier in MS.).
Distinguishing characters :—RX. ferrum-equinum cannot be confused
with any other British bat. The nose-leaf marks its genus, and the
length of the forearm, always over 50 mm., its species.

The two Horseshoe Bats were, by the earlier naturalists,
regarded as mere varieties of a single species, and the credit
of properly distinguishing them in this country belongs to
Montagu, who was also the first to discover the smaller species
as a member of the British fauna.

The larger bat was first made known as a European mammal
by Daubenton, and as a British species by Latham, the latter of
whom supplied Pennant with an account of it from specimens
taken in the saltpetre houses of the Dartford powder mills in Kent.

All the members of the singular family to which the present
species belongs are distinguished by the possession of a compli-
cated cutaneous development upon the nose, and it is to the
shape of the anterior portion of this organ that the Horseshoe
Bats owe their name. It is not easy to explain the precise use
of this very remarkable structure, but it seems unnecessary to
adopt any of the highly ingenious theories which have been
advanced to account for its presence. It is quite consistent
with probability, and indeed almost indisputable, that the
membranous expansions which are the property of the leaf-
nosed or large-eared bats, are in some way instrumental in the
execution of those marvellous passages through narrow or
intricate places, of which these creatures are past-masters, and
for the performance of which their eyes can be of little service.
No one who compares the motions of a Horseshoe with those
of a typical bat can fail to be struck by the marked contrast
between them. Active and alert as all bats undoubtedly are,
even when liberated in a room to which they are complete
strangers, yet the majority make repeated mistakes, and, as
has been more than once remarked, will fly against the glass
of a window, or strike it with their wings. A Horseshoe, on
the contrary, if liberated in a room, and with its attention
not otherwise occupied, as when in pursuit of its prey, avoids

238 RHINOLOPHIDA—RHINOLOPHUS

with perfect ease all sorts of objects, and, being able to instantly
detect the presence even of an almost invisible obstacle such
as glass, will examine a window pane, as if in search of some
means of escape, yet without so much as a touch of a wing.
When it is considered that the glass would appear to be—as
regards the sense of vision—equally perceptible to the one bat
as to the other, an experiment of this kind becomes particularly
interesting, tending, as it does, to show that a high develop-
ment of the cutaneous system is, in the Horseshoes, accom-
panied by a correspondingly acute perception of the environment.

The habit of flying about low down in a room, and amongst
the legs of tables and chairs, is so characteristic of these bats
that it immediately impresses itself upon the observer. It is
not unlikely that it has some close connection with the flight-
less insects, beetles and spiders, which, as Mr T. A. Coward
has shown, forma part of their diet. Not (as mentioned above)
only do these bats show great alertness in avoiding obstacles to
their flight, even in a place to which they are strangers, but
when at rest they display a remarkable power of perceiving an
approaching danger, even when it is behind them, or otherwise at
such a distance as to be out of the range of their necessarily
limited vision. In this connection, Mr Charles Oldham says :—

“Even when sunk in their winter sleep, they appreciate
aman’s approach. The eyes are, of course, then shrouded by
the wings and the sense of danger must be conveyed to them
either by hearing, smell, or, as seems to be most probable, by
the exercise of their extraordinary tactile sense which enables
them to actually feel the approaching danger. I have often,
whilst still some paces from it, watched a sleeping bat of this
species raise and lower its body by flexing the legs and twist
it from side to side with a jerky, spasmodic action. The
enveloping wings are meanwhile slowly relaxed and unfolded ;
then suddenly the bat drops from its foothold and flits away
into the recesses of the cave.”

It is remarkable that, although thus alert and active to
such a surprising degree, the senses upon which a Greater
Horseshoe relies for its information in regard to the outer
world are not those of sight or smell. Mr Bruce F. Cummings’

1 Zoologist, 1907; 292.

(

THE GREATER HORSESHOE BAT 239

found that, whereas the fumes of camphor or ammonia and the
glare of a strong acetylene lamp passed unnoticed, the rattle of
a cup and saucer, or the passing of a finger gently over_his
own forehead, at a distance of nine inches, put a bat of this
species at once on the guz weve.

It is probable, then, that, apart from the generally sensitive
condition of the wings and membranes, the sense of hearing is
the most important aid in the capture of their prey by these
bats. Mr Coward found that a large beetle was imperceptible
to his captives at a distance of only two inches, and so
long as it remained quiet, attracted no attention unless it
was actually touched. But directly it buzzed the result was
very different. A bat’s attention was at once attracted and
the beetle pursued and captured.

If the flight is characteristic, the method of alighting is
equally so. In the words of Mr Oldham,’ the ordinary pro-
cedure of other bats, when alighting on a vertical surface,
such as a wall, a picture-frame, or a curtain, is “by pitching
head uppermost. They then shuffle round instantly, and hang
by their feet, in a convenient posture for another flight.
But the Lesser Horseshoe Bat? is infinitely more adroit.
Immediately before it reaches the wall, or the object on which
it desires to rest, it turns a somersault, timing its action with
such nicety that it clutches the object with its feet, and is at
once in the attitude for taking flight again. The bat by per-
forming this manceuvre can suspend itself, in the act of alight-
ing, from a horizontal surface such as a cave roof, as readily
as from a vertical wall. Two individuals which I had under
observation used after each flight to disappear beneath a couch,
where I always found them suspended from the webbing which
supported the seat. By lying on the floor with my head under
the couch, and then disturbing the bats, I was able, on their
return, to witness this feat of aerial gymnastics, which is
perhaps unequalled among the higher animals.” Natterer’s
Bat and the Barbastelle alone in addition, so far as’ is known,
attempt the somersault, and that feebly, and with indifferent

1 Mem. and Proc. Manchester Lit. and Philosoph. Soc., x\ix., 2, 1-11, pl. and 4 figs.,
31st March 1905, but the peculiarity seems to have been noticed by de Jurine at
Geneva before 1800 (see Zoologist, 1850, 2815).

2 These remarks apply equally to the present species.

240 RHINOLOPHIDAE—RHINOLOPHUS

success. So accurate is this bat’s aim, and so sure its grip,
that it can find sufficient inequalities for a hold on objects from
which other species would inevitably slip off if they attempted
to settle upon them. Mr Coward, for instance, has observed
one alighting upon a smooth brass rod, and on the almost
imperceptible edge of wall-paper, at its junction with a cornice.

Unlike other species, neither of the Horseshoes is capable
of walking on a flat surface. Yet in captivity they have no
objection to alight on the flat. On such occasions their only
mode of forward progression is by a series of awkward leaps of
an inch or two in length: these are made with the wings half
spread, and the legs extended helplessly backwards, and end in
a thud as the bat falls prone on the ground. If left to them-
selves, they prefer to grope behind them with their feet, and if
a vertical surface be at hand, will climb it readily, ascending
backwards with or without the assistance of the thumbs. But
they can rise quite easily from a flat surface, leaping into
flight with astounding activity, no matter how confined the
surroundings. Mr Oldham has seen a Lesser Horseshoe take
wing from the bottom of a box three inches deep and not more
than five inches wide.

When hanging at rest in their natural attitude of suspension
by the feet, these bats are very beautiful objects. The wings
are draped around them in a manner not found in vespertilionid
bats, but rather resembling that of certain exotic fruit bats,
and so neat and ordered that it almost recalls the folding of
a flower or leaf-bud. Their appearance as a whole has been
likened to that of a butterfly-pupa by Mr R. Newstead, who
was the first to photograph one in this attitude. The legs
are held perfectly straight, and, with the exception of the upper
portion of the head, the tips of the ears’ and a narrow dorsal
space, the animal is completely enshrouded in its flying mem-
branes. Sometimes the head also is covered, but a narrow
space between the forearms is always exposed, this arrangement
being brought about by the upper arms being laid diagonally

' Zoologist, 1897, 537-538, and plate ii. See also, Oldham, of. cz¢. supra, the
present account being adapted from these two writers.

2 In captive Horseshoes the ears may be visible, but if in really good health they
are completely hidden in the wings, being slightly bent so as to fit into the natural
pockets formed by the membranes (Coward).

THE GREATER HORSESHOE’ BAT 241

across the back, the elbows meeting at the lumbar region,
where they touch the tip of the reflexed tail: the forearms,
bent sharply at the elbows, are laid lengthwise along the back
so that the wrists flank the ears at éither side, while the fingers
with their connecting membranes are folded over the ventral
surface of the body. The tail, instead of being bent forwards
is reflexed over the back,’ particularly so in the lesser species, and
closely overlaps the wrinkled wings between the fifth fingers and
the legs. In captivity the exposed portion of the back is usually
much greater, for the wings hang every hour more loosely as
the frail creature, which seldom survives capture, grows weaker.

As a bat hangs suspended when awake, it presents a
good deal of animation, changing from one foot to the other,
as it cleans itself, or twists and swings apparently to observe
anyone who approaches. The head is full of lively action, and,
-as remarked by Jonathan Couch many years ago, both ears
and nose-leaf appear to be very sensitive and mobile.

The Horseshoes are more partial to caves, particularly
those of limestone districts, for their places of retreat, than
are any other British species. But they affect also dark old
buildings, lofts and roofs of dwelling-houses, as found by R. F.
Tomes at Ragley, near Alcester. Their resorts, even when
they themselves are hidden safely in the secure recesses of some
crack or crevice, are usually betrayed by their excrement.
This has in some cases accumulated so deeply on the floor
below as to suggest an immemorial tenancy, an inference
borne out in the case of the famous Devonshire Cavern, Kent's
Hole, by the approximation in condition there of bones of the
present species with those of the Mammoth, Spotted Hyena,
and Woolly Rhinoceros, as if the bats had resorted to this
particular cavern at least from the Pleistocene era.

The two species of Horseshoe occur together in their
favourite caverns, but in Britain the present species had been
but little studied until Mr Coward made a special expedition
to the Cheddar caves of Somerset, the result of which was to
add many facts to our knowledge.’

1 A fact observed by J. Couch before 1853 (Zoologis¢, 1853, 3941).
2 See Proc. Zool. Soc. (London), ist August 1907, 312-324; also Mem. and Proc.
Manchester Lit. and Philosoph. Soc., 52, xi., 1-12, pl. and figs. 1-4, 21st April 1908.
X

242 RHINOLOPHIDAZE—RHINOLOPHUS

In the Cheddar caves and galleries, which are often of
considerable extent, the bats hang suspended from the walls or
roof, or creep into the fissures and crevices. They rest in two
distinct conditions, either singly, or more frequently in groups,
which may number as many as forty or fifty individuals. Even
when resting in company, however, the individuals of each
group, unless they happen to occupy a fissure or other confined
position, do not as a general rule “bunch” together, but hang
each at a small but definite distance from its neighbour.

Occasionally, however, especially in the unfamiliar surround-
ings of captivity, they show their social instinct by alighting
upon a comrade, in which case they cling tightly to the first part
that their feet may happen to clutch. Once Mr Coward saw
a party hanging together from a spur of rock like a swarm of
bees. They were all awake, and had evidently assumed this
formation under normal conditions on the way out for their
evening flight. This “bunch” rapidly dissolved when subjected
to the light of a lamp.

Mr Coward’s observations as a whole corroborate those of
Monsieur H. Gadeau de Kerville, who writes of the present
species in Normandy, where it is, with the Pipistrelle, one of the
commonest bats, a colony having been found to include no less
than one hundred and eighty individuals.

M. Kerville has published an interesting photograph taken
by flash-light, of a sleeping colony composed of about eighty
individuals.! He finds, however, that, even in the rather cold
climate of Normandy, the sleeping colonies wake up at times
and disperse to other portions of their cavern. But he is
unable to say whether all the individuals, isolated and grouped,
alter their positions, and whether the colony, when re-formed
in a new position, is composed of the totality of individuals of
which it had previously consisted, or of a greater number. He
is inclined to believe, without laying stress on the point, that
the re-formed colonies consist of smaller numbers.

Other French writers, Messieurs R. Rollinat and E. L.
Trouessart,’ writing from the Department of Indre, find this
bat (as is frequently the case in England also) in summer

| Naturaliste, 15th October 1891, 239 and plate (seen in reprint).
2 Mém. Soc. Zool. de France, 1897, x., 114 and 121.

THRE GREATER HORSESHOE BAY 243

deserting the caves in which it has spent the winter, and
installing itself in small parties in churches and deserted mills.
After an hibernation uncertain and liable to interruption by
any unusually warm days, it issues forth on mild March days,
and frequents, with low and clumsy flight, with momentary
accelerations, the corners of woods and parks, the sides of high
hedges, or the courses of winding streams and rivers.

Similarly, in England the bats probably do not hibernate,
at least in any strict sense of the word, but are active, and feed
whenever the weather is mild, and even in time of frost may
shift their positions within the caves, both when disturbed and
on their own initiative; in winter they prefer a position at
some distance from the entrance. On these points Mr Coward
writes that not only did they change their position, but
that colonies which he visited on several occasions gradually
diminished in numbers and finally disappeared, having appar-
ently been disturbed by his presence, though the bats themselves
were not actually molested; one colony consisted of forty on
29th December; on the 31st, when visited after sundown,
it contained only eight, and two of these flew out of their
own accord. On 3rd January the spot was unoccupied, and
no bats made use of this particular position during the four
following days on which he was able to watch them. Another
colony, consisting of two companies of twelve and eight respec-
tively, was two days later reduced to ten, and on the following
day to four. A third colony of forty or fifty was similarly reduced
on a second visit. Mr Coward found that the bats, at the end
of December and beginning of January, were not hibernating ;
they woke without artificial stimulus in the caves, took wing, and
actually left the caves of their own accord, apparently to feed. .
Messrs Coward and Oldham watched bats emerge from a hole
in one cave, and pass out between 4.40 P.M. and 5.20 P.M. on
5th January, and Mr Cummings observed the same thing at
Barnstaple, Devonshire, in December and January.

Mr Coward’s notes, although prolonged a week further
by Mr Cummings’s! experience near Barnstaple, extend only
to 16th January. Detailed records for the succeeding months
are not yet available, but Couch’s account of a Greater

! Zoologist, 1907, 288-294.

244 RHINOLOPHIDA:—RHINOLOPHUS

Horseshoe taken on the wing in Cornwall’ on rith January,
and Mr F. C. Rawlings’s specimen shot at Barmouth on 5th
February, deserve mention. ‘The internal temperature of the
caves visited both by Mr Coward and Mr Cummings seemed
to be fairly even, varying between 50° and 52° Fahrenheit,
and it is believed to be about the same throughout the
year.

Mr J. G. Millais aptly describes this species as sailing and
fluttering with visibly broad wings and delicate butterfly flight,
but seldom rising to any height in the air. His statement,
however, that it appears at a rather late hour in the evening
will possibly need correction in view of Mr Coward’s observa-
tion that it began to fly at 4.40 P.M. in mid-winter at
Cheddar.

There can now, I think, be little doubt that the Horse-
shoes do not, like most other bats, consume their prey while on
the wing, but habitually alight to eat it, conveying it for this
purpose to certain favourite dining-places within the shelter of
the caves. These, even when the diners are absent, are
betrayed by the débris of wings, elytra, and other fragments,
as well as by the heaps of excrement which fall to the ground
during and after a meal. The extent of these refuse-heaps
indicates that the bats have strong preferences for certain spots,
to which they return time after time. This fact was first
mentioned by Mr A. H. Macpherson,” but has been insisted
upon by Mr Coward. The latter naturalist finds that the
ordinary food of the larger species consists of big beetles and
moths, and the condition of the refuse-heaps leads him to
infer that the beetles form the bulk of the winter repasts, while
in summer beetles and moths are consumed in about equal
numbers. Besides these, the remains of flies of more than one
size and species have been identified from the refuse-heaps,
and the presence therein of portions of certain flightless beetles,
hibernating moths, and of a large cave-haunting spider,® led him

' Zoologist, 1853, 3941. [bid., 1887, 262.

* The following insects are also mentioned by Coward as forming part of the
diet :—the large beetles MZelolontha vulgaris of Fabricius, Geotrupes spiniger of
Marsh, and G. stercorarius of Linnzus ; a staphylinid beetle of the genus Quedius ;

the flightless beetle Mebria brevicollis of Fabricius, and a Pterostichus; a geode-
phagous beetle, perhaps of the genus Amara, and, perhaps, a Dytiscus,; the moths

THE GREATER HORSESHOE BAT 245

/ from the first to suspect that this bat may habitually take some

at least of its prey while at rest.

The habits of the Greater Horseshoe, as observed by Mr
Coward, in captivity, were very instructive. ‘On one occasion
a bat dropped near a beetle which had buzzed, for undoubtedly
the buzzing of a beetle at once attracted the bat’s attention ;
the bat moved its head to and fro, the lower edge of the horse-
shoe touching the floor. The beetle walked a few inches away,
and then again attempted to fly; instantly the bat followed it
in a series of little jumps, really short flights of a few inches,
and after two or three jumps reached and fell upon the beetle,
which it at once thrust into its interbrachial membrane.
Directly it had secured the beetle it rose from the floor, flew
to a customary perch and, there hanging, consumed it. In the
cage the method was similar; the bat dropped on to the floor
of the cage, lying with extended wings, and either feeling or
smelling round—at least that is what the action suggested—
until it found a beetle; directly one was secured, it sprang up,
turned in the air, and clutched the bar of wood, only twenty
inches above it, with its feet. The beetle was then pushed into
the wing as usual, and the head and perhaps other fragments
dropped. This, then, is evidently the way in which flightless
beetles and spiders are caught, and possibly coprophagous
beetles may be thus picked up when they are crawling over
dung.

“This is, however, not the only way in which the Greater
Horseshoe secures its food; it can and does catch insects on
the wing. G. typheus is a beetle which flies during mild
weather in winter, and when I released a dozen beetles in my
room in the evening, two or three would quickly attempt to fly.
It was when this occurred that I felt certain that the bats hunt
and locate their prey mainly by means of their acute hearing.
The deep booming buzz of the flying beetle at once roused the

Scotosia dubitata of Linneus, Gonoptera libatrix of Linnzus, 77iphena orbona of
Fabricius, 7. fronuba of Linnzeus, and Xylophasia polyodon of Linnzeus. Several
Diptera, including Muscide apparently of the genus Lucz/éa; a wasp-like Hymen-
opteron, and caddis-flies. Numerous pellets of excrement were carefully examined for
Coward by R. Newstead, to whose identification many of these names are due.
No doubt many other insects might be added, and the moths 77iphena fimbria and
Agrotis saucia were identified by J. E. Harting for Macpherson (oc. cé¢.).

246 RHINOLOPHIDAZ—RHINOLOPHUS

Horseshoes to activity, even when, as was often the case after »
eating two or three beetles, their heads were drooping, and
they were relapsing into sleep. Usually the bat left its foot-
hold immediately the beetle began to buzz, and as these beetles
are not always quick in getting on to the wing, the bat
frequently skimmed over and missed its prey. But when the
beetle had risen two or three inches from the ground, it was
doomed ; the bat came down like a falcon stooping, and with
marvellous precision caught the flying beetle in its jaws, and
carried it off to some place where it could pitch and devour it.
For several weeks this performance was repeated on an average
two or three times each night, and though on a few occasions
the beetle got well into the air before it was captured, by far the
greater number were secured before they had risen many inches
from the ground.”

In feeding, the Horseshoes differ from the typical bats
in one marked particular. The interfemoral membrane being
too small for use as a pouch, they usually requisition the
posterior part of the wing to enable them to hold an insect
of troublesome proportions. When dealing with a beetle, Mr
Coward’s bats always fluttered their wings rapidly, as if almost
worrying it with the rapid movements of the head. These move-
ments recalled the vibration of an insect’s wings rather than
the ordinary struggle of a bat when held in the hand. The
process was first observed by Mr Coward in his captive
specimens. A large beetle was thrust into the natural bag,
the claws of the corresponding leg “were usually released from
their hold, and the whole wing brought suddenly forward, by
simultaneous stroke of arm and leg, to meet the head. The
beetle was practically beaten against the membrane by rapid
movement of the bat’s head, assisted by the forward stroke of
the wing. This wing-action, suggestive of the uses of a hand,
has no exact parallel in the apparently similar use of the inter-
femoral pouch by vespertilionid bats. The beetle was moved by
the bat against the membrane, for its position in the mouth had
frequently to be shifted before the bat could devour the abdomen
and reject the head, and sometimes the action of head and wing
together actually pushed the beetle further into the mouth.
When the beetle was first seized the wings of the bat were

(¢

THE GREATER HORSESHOE BAT 247

only slightly unfolded, held free but with the membrane partially
hiding the body, and when the bat took a beetle from the hand
it beat rapidly with both arms but did not grasp with the
thumb. Directly the beetle was in the bat’s jaws the wings
were further opened, and hung quite loosely whilst the beetle
was being devoured.

“After a few seconds the head was withdrawn from the
wing and the beetle masticated ; the rejected portions fell, and
the bat, generally suspended by one leg, swung from side to
side. This swinging round was even more remarkable when
both feet were attached to some hold; the animal could then
turn almost completely round, crossing its legs, without altering
the position of its feet. When the beetle was finished, the bat
usually bent forward, and two or three times touched the object
from which it was suspended lightly with its lips; this was
especially noticeable when a bat had been feeding when hanging
from my hand. Frequently, also, one leg was brought forward,
and the teeth scratched or the lips combed by the claws, prob-
ably to get rid of some particles of beetle which were sticking to
the teeth or lips.

‘When the beetle was quite finished, and the subsequent
performances had been gone through, the bat hung, bending its
whole body forward, turning from side to side, and moving its
head, ears, and nose-leaf with great rapidity ; it appeared to be
looking for food, but perhaps searching for prey would be a
more correct way of expressing it.”

At first the number of beetles eaten per bat was from five
to eight each night, but later, when they were regularly feeding
themselves, they took as many as ten or twelve, and even occa-
sionally sixteen ina night. The quantity eaten did not depend
upon the number left in the cage, for frequently beetles were
untouched in the morning; this may, however, have been due
to inability on the part of the bats to find and secure them all.

“The Horseshoe drinks by lapping with the tongue. It is
a thirsty animal, and we can only suppose, from its behaviour
in captivity, that it obtains water in its natural state ; possibly,
like other bats, it hovers over pools of water and laps whilst on
the wing.”

Mr Coward's captives afforded him an opportunity of ascer-

248 RHINOLOPHIDAZ—RHINOLOPHUS Fy.

taining the position of the curious reduced interfemoral »

membrane and tail. During the whole process of pouching ,
and eating a beetle the posterior portion of the tail remained in
the characteristic recurved position.

“This reflexed tail (writes Mr Coward), or to be more
exact, portion of the tail, is constantly in this curious position.
In flight the anterior portion of the interfemoral membrane is
stretched between the slightly flexed legs ; the end of the tail is
upturned ;' and when the bat is scrambling or climbing, the tail
is held in the same position; when at rest the tail is flat upon
the back if the wings are half-open, or lies partially concealed
by the forearms if the animal is closely wrapped in its wings.”

The sounds made by this bat are variable. Mr Coward
compares its voice to a sparrow-like chirp or chattering,
while Mr Millais describes it as a somewhat subdued squeak,
louder in the young, which, when clinging to their mothers or
following them about when nearly full grown, keep up a con-
stant high-pitched double cry. Two which he kept in his own
house maintained this cry, apparently without ceasing, through-
out the day and night.

It seems probable that, though not without exception, the
sexes keep their own company. This was Mr Coward’s
experience, and it is also possible that, as suggested for
the Lesser Horseshoe, there is an actual disparity in their
numbers, the males being more plentiful. In France, according
to Messrs Rollinat and Trouessart, the pregnant females unite
to form more or less numerous bands in April, and do not
separate until they have reared their young. They are fre-
quently accompanied by a few non-breeding young of both
sexes, and even, very often, by adult males.

Messrs Rollinat and Trouessart have published an interest-
ing account of the breeding habits.” They find that the
pairing season is in autumn, but that fertilisation, and conse-
quently the true period of gestation, does not commence until
the resumption of the period of activity in spring. This varies
with the vigour of the individual and the situation of its winter

' Cummings corroborates this point, oc. c7Z., 291.
® Loc. cit. ; also, Comptes rendus Soc. Biol., 26th January and 6th July 1895, 53-54,
and 534-536 (reprints).

THE GREATER HORSESHOE BAT 249

retreat, so that the date of birth of the single’ young is uncertain.
The first, however, appear about the 2oth June, after a period
of gestation lasting ten or eleven weeks ; the last are probably
born before the end of July.’

The young are carried about by their mothers, attached to
the false nipples by their teeth. When they are hungry they
transfer their hold to the true nipples, and on the milk derived
thence they are nourished for a period, estimated from the
analogy of the Mouse-eared Bat, of about two months. They
grow rapidly, and by the middle of September are nearly as
big as their parents.

The attitude of the young one, clinging as it does by its teeth
to the false nipples of its mother, differs in a marked particular
from that of the typical bats. Its wings lie close to its
mother’s abdomen, its feet cling one to each of her flanks, its
tail, with the tip recurved as in the adult, stretches near her
neck. The result is that, as the mother hangs head down-
wards, the little one’s head is uppermost, a position unique
amongst British bats.

Mr Coward's observations have done much towards dispell-
ing the deep ignorance hitherto prevailing in regard to the
habits of one of the finest and most remarkable of British bats.
He was, in fact, the first English naturalist who succeeded
in keeping it alive in captivity, in one case for a period of
nine weeks.

There are some curious myths circulating amongst conti-
nental naturalists, some of whom have stated that the Horse-
shoes sometimes suck the blood of sleeping birds and mammals.
Fatio even quotes without disapproval Kolenati’s statement
that the present species roams by night with criminal intent
amongst birds’-nests, or haunts for sinister ends the rocky
mountain refuges of the Chamois.

The large size, broad wings, and curious butterfly-like flight
should make this species easily recognisable.

' Note, however, that in Germany Kuhl found two, sometimes one (see above,

p. 231).
* Arthur Whitaker had one born in captivity on 14th July.

250 RHINOLOPHIDAZ—RHINOLOPHUS

THE LESSER HORSESHOE BAT.

RHINOLOPHUS HIPPOSIDEROS (Bechstein).
RHINOLOPHUS HIPPOSIDEROS MINUTUS (Montagu).

1760. LE FER-A-CHEVAL, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon’s
Histoire Naturelle, viii., 131-133, pl. xx., figs. 1 and 2; also, Mém. de l’Acad. Roy.
des Sct. 377, 382, pl. xv., fig. 4, 1759, published 1765 ; described from France.

1774. VESPERTILIO FERRUM-EQUINUM (in part), J. C. D. von Schreber, Déze
Sdugthiere, i., pl. \xii., two lower figures, 174, 190; evidently naming Daubenton’s
Le Fer-a-cheval; Montagu, Transactions Linnean Soc. (London), ix., 162, 1808 ;
Bingley ; Clermont.

1776. VESPERTILIO EQUINUS (in part), P. L. S. Miiller, Vatursystems Supplements
und Register Band, 20.

1785. VESPERTILIO UNGULA (in part), P. Boddaert, Elenchus Animalium, \., 7%;
renaming Schreber’s V. ferrum-eguinum.

1788, VESPERTILIO FERRUM-EQUINUM, f. minor, J. F. Gmelin, Systema Naturae, i., 50.

1792. VESPERTILIO FERRUM EQUINUM MINOR, Robert Kerr, Animal Kingdom, 99
(not V. molossus minor of Kerr, op. ctt., 97); described from France.

1797. NOCTILIO HIPPOSIDEROS, J. M. Bechstein, Der Zoologe, I., i.-Vill., 65.

1798. VESPERTILIO HIPPOCREPIS (in part), F. von P. Schrank, Fauna Boica, i., 64.

1800, VESPERTILIO HIPPOSIDEROS, J. M. Bechstein, in Tomas Pennani’s Allgemeine
Uebersicht der vierfiissigen Thiere, ii.. 629 footnote, see also 615 and 736;
Donovan.

1803. RHINOLOPHUS MINOR, Etienne Geoffroy, Catalogue Mammifcres Mus. @ Fist.
Nat. (Paris), 57; described from Paris.

1804. VESPERTILIO (? RHINOLOPHUS) HIPPOCREPIS, Johannes Hermann, Observa-
tiones Zoologicae, 18.

1808. VESPERTILIO MINUTUS, George Montagu, Zvans. Linnean Soc. (London), ix.,
163, etc.; described from Devonshire, England; Pennant, ed. of 1812; 183,415,
pl. xiv., figs. 3 and 4.

1813. RHINOLOPHUS BI-HASTATUS, Isidore Geoffroy, Ann. du Mus. a’Hist. Nat,
xx., 259; renaming Daubenton’s Le petit Fer-a-cheval.

1816. PHYLLORHINA MINUTA, W. E. Leach, Systematic Catalogue of the Specimens
of The Indigenous Mammalia and Birds that are Preserved in The British Museum,
etc. (London), 1 ; named, without description, from Torquay, Devon.

1817. RHINOLOPHUS HIPPOSIDEROS, W. E. Leach, Zoological Miscellany, iii., 2, sp. 2,
pl. 121, e¢ hlurimorum auctorum.

1829. RHINOLOPHUS BIFER, Jakob Kaup, System der Europiische Thierwelt, i. 104 ;
without description, hence a omen nudum.

1862-63. RHINOLOPHUS HIPPOSIDEROS, var. TYPUS, ALPINUS, et PALLIDUS (in part),
Carl Koch, Jahrbiicher des Vereins fiir Naturkunde im Herzogthum (Nassau), 530-
531; described vaguely and “scarcely determinable” (Andersen).

1885. RHINOLOPHUS BIHASTATUS, var. KISNYIRESIENSIS, Daday Jeno, Ovrvos-
Természettudomdnyt Ertesito, etc., x. (3), 274; described from Kis-Nyires, Trans-
sylvania.

THE LESSER HORSESHOE BAT 251

1887. RHINOLOPHUS HIPPOSIDEROS var. TROGLOPHILUS, Daday Jeno, Ertekezések
A Természettudominyok Korébol (Budapest), xvi., Nos. 7, 8, pl., figs. 1 and 2, 1886 ;
described from Kis-Nyires, Transsylvania.

1891. RHINOLOPHUS HIPPOSIDERUS, Blandford ; Flower and Lyddeker ; Johnston.

1905. RHINOLOPHUS EURYALE HELVETICA, K. Bretscher, Vierteljahrsschrift der

Naturforschenden Gesellschaft (Ziirich), xlix., 256, 1904; thus identified by
Andersen.

1905. RHINOLOPHUS HIPPOSIDERUS MINUTUS, Knud Andersen, Proc. Zool. Soc.
(London), 17th October, 142; Trouessart (1910).

Synonymy :—The Lesser Horseshoe Bat was first recognised as
distinct on paper by J. F. Gmelin in 1788, and was formally named
by Bechstein in 1797. It was discovered in Britain by Montagu, who
wrote an elaborate paper describing its specific characters. The name
which he proposed for it—Vespertilio minutus—is antedated as applied
to the species as a whole by Bechstein’s hzpposzderos; having been
based, however, upon English specimens, it is available for the British
race as differentiated: by Andersen (see Ann. and Mag. Nat. Hist,
Dec. 1905, 648-662; and Proc. Zool. Soc. (London), 17th October
1905, 142).

A curious instance of the possibilities resulting from strict adher-
ence to rules of nomenclature suggests itself by Gmelin’s use of the
words major and minor. He clearly did not intend to apply them sub-
specifically as such, but, writing in Latin, he used the words descriptively.
Nevertheless, strict adherence to rule might conceivably disregard his
intentions, and, considering only the form of what he wrote, assign
these names to the two bats which he had in view. In that case
Rhinolophus major would become a synonym of R. ferrum-equinum, but
hipposideros would give place to mznxor. I find no rule governing this
instance laid down in any code of nomenclature, and, until such rule be
made, I shall not venture to adopt an alteration so inconvenient.

Distribution :—R. hzpposideros ranges from Ireland to Gilgit, and
from the Baltic to Sennaar, Africa. It ascends the warm valleys of the
Himalayas to at least 6000 feet in summer. It is divided by Andersen
into three closely allied sub-species, viz., Heuglin’s szzzmus, inhabit-
ing the Mediterranean region, Egypt excepted, with the Balearics, Malta,
Corsica, and Crete, to Sennaar and Keran; hippostderus (szc) proper, of
central Europe north of the Balkans and Alps with Cyprus, through north-
west Persia and Armenia to the extreme north-west Himalayas; and
Montagu’s mznutus, of the British Islands. The distribution in France,
as given by Rollinat and Trouessart (J7ém. Soc. Zool. de France, x., 125,
1897), may fitly be compared with the British. Although inhabiting
the whole country it becomes rarer in the north-east, but is very
common in the south-east, south, centre, and west.

The British distribution of R. hzpposideros repeats and amplifies the

252 RHINOLOPHIDA:-—RHINOLOPHUS

features of that of R. ferrum-equinum. Thus, while quite unknown in
east Anglia and rare in the Midlands, it is common in many parts of
the south of England from Kent westwards, is widely distributed in
the west and in Wales, and may be -said with certainty to range as
far north as Ripon in Yorkshire. The eastern boundary of its
English range is thus roughly crescentic, the tips of the crescent
resting on Kent and Yorkshire. Furthermore, the’ bat is found in
Ireland, but here again, although stated to be the “Common Bat” of
parts of the west, where it occurs from Kerry north to Galway, it has
never been detected in the east, nor, one doubtful record excepted, even
in the centre.

Tracing out this crescent, H. A. Macpherson knew of a Hertford
specimen, and cites Sevenoaks in Kent as one of its habitats (Zoolo-
gist, 1887, 152); while Millais has seen others from Maidstone and
Chiselhurst, and, with Ogilvie-Grant, has frequently observed it in
Kensington Gardens, London, whence a specimen was identified by
Harting; Tomes found it at Godstone, Surrey (Bucknill and Murray);
but no records are forthcoming from Sussex ; and Hart knew of only one
Hampshire specimen (Trevor Battye and Lascelles); while another
reported to Kelsall as having been taken at Niton in Wight many years
ago (Zoologist, 1887, 90) would appear to be the solitary record for that
island. The former records are additional to the information collected
by Kelsall in 1887 (doc. cit.), at which time the species had no known
habitats in the south-east of England. Its absence from Essex, Suffolk,
Norfolk, Cambridge, Lincoln, Bedford, Rutland, and Huntingdon can
hardly be due to oversight in the face of the presence in these counties
of a long succession of naturalists, including Laver, Rope, Caton Haigh,
and Cordeaux. The first-named in particular has, as he positively
assures me, in a lifetime devoted to the study of the local fauna, never
seen nor even heard of a local specimen, so that Millais’s statement
that he has “seen specimens from Colchester” would seem to need
confirmation.

In the south-west the species was discovered by Montagu in Wilt-
shire in two localities (7rans. Linnean Soc., ix., 1808, 162-171); and after-
wards (with R. ferrum-equinum) in Kent’s Hole, an extensive limestone
cavern near Torquay in Devon, and it has since been encountered
frequently within the county borders (see Jordan, Zoologist, 1843, 75 ;
Borrer, Journ. cit., 1874, 4129; Lilford, /ourn. cit., 1887, 63; Hollis,
Journ. cit. 1907, 110-111). Further records up to 1887 have been
summarised by Kelsall (doc. cz¢.), showing that it occurs, often abund-
antly, in Cornwall (see also Couch, Journ. czt., 1853, 3941, and Millais),
sometimes indeed, almost to the exclusion of all others (Clark); in
Somerset, where the Mendip Caves may be mentioned as a specially
favoured haunt (see also Lewis, Journ. czt., 1906, 69, and Millais); and in

— eee.

XVI.

PLATE

BARBASTELLE BATS.

THE LESSER HORSESHOE BAT 253

Dorset. North of 51° 30’ north latitude its numbers become uncertain,
and, although stated by Tomes to be not rare at Cirencester, Char-
bonnier and Lloyd Morgan report it as scarce at Bristol, and seldom
obtained north of the Avon. I know of no records either for Oxford
or Buckingham, and of one only for Berkshire (Cocks, Journ. cit., 1906,
186); so that the bat is practically unknown from Cirencester eastwards
to the North Sea.

The species is, however, by no means entirely southern in its
distribution, since it appears again across the Bristol Channel in
Monmouth, where Donovan found it at Raglan Castle in 1802, and
thence it is widely distributed to the extreme north of Wales and in the
north-east to Ripon in Yorkshire, but is rare or absent from the central
English counties south of the Wash. The details are worth noting :—It
has been found by Storrie at Bridgend in Glamorgan (Kelsall); by
Tracy frequently at Stackpoole (Kelsall); by Proger (specimen sent to
Forrest) at St David’s in Pembroke; by Lingwood at Sufton (Aum.
and Mag. Nat. Hist., May 1840, 185); by Wallis near Ross (Zyrans.
Newcastle-on-Tyne Nat. Hist. Soc., xi, 243, 1894); and by Hewitt,
who sent a specimen from Ledbury; all in Hereford. It has been
noticed sparingly in many parts of Merioneth by Caton Haigh,
who believes it to be generally distributed in North Wales
(Zoologist, 1887, 152). Further north, Oldham has detected it in
Carnarvon (Journ. czt., 1903, 430); and Newstead, Caton Haigh, Oldham
and Coward in Denbigh and Flint (/ourn. cit., 1887, 152; 1896, 255;
1897, 537-538; 1906, 70-71). Towards the east there is only one
known Shropshire specimen, that in the Worcester Museum; it flew
into Steele Elliott’s house at Dowles on the Worcester county border on
6th July 1904 (Journ. ctt., 1905, 308); Coward, however, found it in the
Ceiriog Valley in Denbigh, four miles from the Shropshire border, in
1900. Tomes rates it as by no means rare in Worcester, and local
rather than rare in Warwick; Millais has examined a single Stafford
specimen, and Hardy one from Edwinstowe, Nottingham (Kelsall),
while Cheshire has one very doubtful record, dating from 1834
(Coward and Oldham). In Derby it is found in fair numbers, chiefly
in limestone caverns and old lead workings at Matlock, and in the
surrounding country even as far as the Peak district. It appears
to be absent from the Trent valley and the basin of the lower Dove
(Jourdain). In Yorkshire it was discovered by James Ingleby,
from whom, in January 1876, Laver received specimens taken near
Eavestone, Ripon (Roebuck, Zoologist, 1882, 186); Storey has also
reported it from near Pateley Bridge in Nidderdale (see Fve/d, 12th
April 1884, 499; Vaturalist, 1886, 339; Zoologist, 1895, 65), but a
statement that it is the “prevailing species” in a locality so near the
northern extremity of its range has not been corroborated, and would

Ye 2

254 RHINOLOPHID/Z—RHINOLOPHUS

seem to be an error. Millais states that it has been recorded from
Durham and Northumberland, but it is evident that Ripon must, until
further details be forthcoming, be regarded as the most northerly limit
of its British range. The record by A. R. Young of its occurrence at
Crookston Castle, Renfrew, Scotland (ew Stat. Acc. Renf., 1845, vii.,
162), has not been confirmed (see Alston).

In Ireland this species was first discovered by William King in
Galway in June 1858, and by Foot in Clare on 1oth March 1859.
The former obtained only one specimen (Proc. Dublin Univ. Zool.
and Bot. Ass., 15th April 1859, in Dublin Wat. Hist. Review, vi. 522-
525); but the latter adduced strong evidence to prove that it is
the common bat of the district around Ennis (Proc. Dublin Nat. Hist.
Soc, ist April 1859, in Dublin Mat. Hist. Review, vi. 379-381).
These localities have since been confirmed for Clare by J. R. Kinahan
(Zoologist, 1861, 7617-7624), and Scharff (/v7sk Naturalist, 1902, 175),
and for Galway by More, by whom two specimens, taken at Coole
Park, near Gort, were presented to the Dublin Museum. The bat
has since been reported from Kerry by Hardy, who encountered
a large colony at Muckross Abbey in July 1885 (Kelsall, Joc. céz,,
92-93), and later from the same district by Forrest (zx /z¢. of 3rd October
1905).

As noticed under the preceding species, there is an old record, dating
from 1845, of the occurrence of a leaf-nosed bat of uncertain species in
Co. Westmeath: this, if substantiated, would antedate all other accounts
of Horseshoe Bats in Ireland.

Distribution in time:—The remains of this bat have not been
recognised in the fossil condition.

The breeding season, period of gestation, and number of young
are, according to Rollinat and Trouessart, similar to those of R. ferrum-
equinum (see under that species); except that, owing to later emergence
from hibernation, the young are not usually born so early, a fact which
corresponds with Whitaker’s dissection of females containing embryos
nearly ready for birth on 14th and 24th July (Waturalst, March
1907, 74, and zz “Uiz.). Proger, however, saw young ones born in
South Wales about 20th June (zz /¢.). He has also once observed
spring courtship (see Proc. Cardiff Nat. Soc., March 1905, reprint, 5,
1906), but there is no evidence of consummation.

As in R. ferrum-equinum, the males and females found in autumn
may be divided into three classes, according to their ages, as shown
by their genital organs, up to the third autumn. The authors conclude
that, although well-developed in this respect, neither sex pairs
until the third autumn, when they are nearly two and a half years old ;

1 Jameson, Jrish Naturalist, 1897, 36, gives the dates of occurrences differently,
but, as I think, incorrectly.

;
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a ee ee ee ee ee a

THE LESSER HORSESHOE BAT 255

females are found still virgin until their third October, in spite of the
fact that adult males have the organs in a seemingly functional con-
dition until May.

Description :—In general appearance this bat resembles the pre-
ceding species, but its size is very much less, and its colour darker,
both in fur and membranes.

The ear (Fig. 2, No. 11, p. 7) is rather more deeply sinuate on the
outer margin, the transverse sulci are less apparent (disappearing in
dry skins), and the basal lobe is larger in proportion.

In the nose-leaf the horseshoe is less closely applied to the head,
the concentric rings, especially the median, are more conspicuous,
and the sella is cuneate, not fiddle-shaped, and has its summit
pointed.

In the wing (Plate XVII., Fig. 3) the second phalanges of digits
three and four are less than one and a half times the length of the meta-
carpals. The fourth metacarpal is slightly the longest of the three.

The general colour of adults is, above,
brownish “drab,” with variations either
towards “wood-brown” or “Prout’s
brown,” the bases of the hairs being

everywhere “ecru drab”; the underside
is “ecru drab,” with variations towards
“drab-grey.” I

The wing, ear, and foot are coloured as
in R. ferrum-equinum.

As in the last species, young indi-
viduals are as large as their parents
by their first September, but are greyer,
being very nearly “mouse-grey” above,
with the bases of the hairs everywhere
“drab-grey.” The upper side is clothed
with downy hair at birth (Whitaker).

In the skull (Fig. 26) the auditory
bullz are so large that the basi-occipital is
very much narrowed between them so as
to become a mere bridge of bone. Some-
times the bullae are nearly in contact. Fie 6 een Bet eee

The anterior upper premolar, though Rhinolophus hipposideros.
small, is in the tooth-row, the upper canine (1) Upper and (2) Lower Jaw.
and posterior premolar being well sepa-
rated. In the lower jaw the anterior and posterior premolars are
usually almost or quite in contact, the central external, but there is
some variation in this respect.

The individual variation of this species has not been studied in any

256 RHINOLOPHIDAZE—RHINOLOPHUS a

detail. Kelsall (Zoologzst, 1887, 93) mentions a white one, but without
giving details, and two continental specimens are noticed by Cantoni.

Geographical variation manifests itself chiefly in size, the length of
the forearm being 39 to 41-7 and 34-7 to 38, and of the skull 16 and
14-5 to 15-5 mm, in the typical sub-species and in mznémus respectively.
Minutus agrees in skull with the type, in forearm with mzxzmus. The
differences are slight, but constant for a series.

DIMENSIONS IN MILLIMETRES :—

= =]
ye E z = = =
aa Bf 2 ° = : ke
Sl eel eee oy S25) os os) je 9 eS
2 » Fh bog wg Prsn| s 3 3 3 Ss or
ie 2 ae eI Hi ASE 3 = a = H Sa
= 2 Sas S 2 538 2 i 5 3 15
BS oe R = ls) a to =) = ne
3 a et) O° ss bre | a 3 8 @
S 5b o's Hig S 3 = © 3
= am = = | a a Fs
E S As o| = =
<) me S
Three males (H. J. Char-) { a \ { 8a°4 \ { 23 26 \
bonnier) i Oneal! edleG ex 37°3 St 24 28
caPethe hl’ \} is If Ul 35-3 \ eas | 26
Thirty specimens) Max. as 15°5 | 10-11°8 | 27 18°5 8-7 | 39 a6 24°9 | 28-2
measured by K.
Andersen. f Min. ae 14°2 G=T6li i) ZB TEs eel) 86:84] i ae 22°8 | 24°7
Female, Devonshire (1),
measured in dried skin
at British Museum. - 39 15°5 5C 26 8 88 | 36 4 51 23°5 | 25 241
Female, do. (2), do. . ° 38 15°5 Ag 28 10 | 10°8 | 38 4 53 23 26 250

Nore :—Embryos within the mother reach an expanse of 100 to 110, and by July the young attain to 190.

Proportionate lengths :—TI oot, without claws, about -50 of lower leg ;
fifth metacarpal, about 1-1 of third; lower leg, about -46 of forearm and
about -71 of tail.

Skull :—Greatest length, 16; basal length in middle line, 13-3 to
13-5; palatal length in middle line, 5-0; from posterior border of 7° to
anterior border of canine, 5-7; greatest breadth at zygoma, 8; posterior
breadth, 6-8.

The weight of three is given by Couch (Zoologist, 1853, 3941), as
77,74, and 73 grains=5, 4:8, and 4:7 grammes.

Distinguishing characters:—The length of the forearm, always
under 40 mm. in British, and only slightly exceeding 40 mm. in
continental specimens, at once distinguishes this species from A.
Serrum-equinum.

The Lesser Horseshoe is in appearance and habits to a
large extent a small edition of the larger species. It is, how-
ever, more widely distributed, and was consequently the better
known of the two up to the date of Mr T. A. Coward’s essays

DHE SLESSER: HORSESHOE BAT 257

on its congener. It appears to be in every way a more fragile
and delicate bat, and is to be found in greatest abundance in
districts where the presence of caves affords it an equable
temperature during the cold days of winter.

It is difficult to compare its flight with that of its larger
congener. Although apparently slow in its movements, it 1s,
perhaps, less so than the larger species. It vibrates its wings
more frequently, but is not so graceful, and does not glide so
often. Yet it is very active, since Mr Arthur Whitaker, who
has effected the capture with his net of seven other species
while out of doors and at full liberty, writes me that its
wonderful agility completely defeated him for half an hour in
the confined space of a room.

The behaviour of a Lesser Horseshoe Bat, under the novel
conditions of confinement in a strange room, has been well
described by the late R. F. Tomes, who remarked that it dis-
played in its search for a means of exit an ability which was
quite extraordinary. It literally flew into every part of the room,
going behind and under everything, even under a bookcase stand-
ing against a wall, although there was scarcely a space of three
inches between it and the floor. Some bookshelves in a recess
especially attracted its attention, and after examining them
diligently, it darted into a vacancy occasioned by the removal
of an octavo volume, and out again into the open room, with-
out having been seen to touch anything even with the tips
of its wings. But it was most interesting to observe it when
making an examination of the window, searching every pane
over and over, inch by inch, until it might properly be said that
no portion of the glass remained unexamined. While doing
this, the wings were kept in a vibratory state, the face of the
animal being directly in front of the glass, and very near to it,
as if looking out of the window. The general manners of the
creature, when thus engaged, conveyed in a slight degree the
idea of a hawk-moth when hovering in front of a flower.
From the behaviour of the individual, and its peculiar mode
of flight, it was difficult to repress the idea that it was either
feeling its way about, like a blind person, or feeding for an
opening by which to escape. But, at the same time, its
shyness when approached gave Tomes the strong but, as it

258 RHINOLOPHIDAEK—RHINOLOPHUS

now appears, questionable idea that its sense of sight
was by no means deficient. In its habits the Lesser is not
known to differ from the Greater Horseshoe, but its smaller
size must necessarily affect its food, since it is not large enough
to seize the big beetles on which the latter so largely sub-
sists, but it is evidently powerful enough to capture moths.
Several naturalists have found that, in captivity at any rate, it
will eat the cave-spider and one or both of the cave-haunting
moths ;* and the fact that Mr Coward’s captive specimens
more than once alighted on a table to eat mealworms lying on
it, taken in conjunction with that writer’s observations on the
larger species, makes it appear almost certain that both Horse-
shoes may habitually feed when at rest.

This bat is one of the commonest if not the ‘‘common bat”
of some parts of the west of Ireland, and one of the first printed
accounts of a Horseshoe bat-cave was penned in that country by
F. J. Foot, in 1859.” This was supplemented by a further
exploration of the County Clare caves, undertaken by Foot and
J. R. Kinahan in 1861. The caves examined were all in the
neighbourhood of woods or plantations, and were the winter
habitat of certain spiders, moths, and gnats. The bats did not
seem to be particular as to either the height from the ground or
the part of the cave where they hung. They were found
suspended at all elevations, from those out of reach of a man to
within two inches of the ground. Although most frequently
tenanting the dark inner recesses, they were also encountered at
rest in broad daylight at the entrance. They were distributed
either singly or in companies, not, however, thickly crowded
together, and it was shown that their lethargy, at least in March,
was not so profound as to prevent them from shifting their
quarters. Foot supposed that their movements must be due to
alterations in the moisture of the cave, causing them to retreat
to the driest parts of it, and Kinahan thought that they might
feed on their insect companions. It is remarkable that of fifty-
four bats, carefully and separately examined, all but four were

! Scotosia dubitata and Gonoptera libatrix.
2 “Proc. Dublin Nat. Hist. Soc.,” printed in Dublin Vaz. Hist. Review, vi., 379-381,
1859.

3 Zoologist, 1861, 7617-7624.

;
|
)
|
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~~ Ta

THE LESSER HORSESHOE BAT 250

males, and of the four females two were found on 1st April at a
place where on 8th March there had been four males, while an-
other was hanging by herself in a separate part of one of the
caves. Similarly, of twenty specimens examined by Foot in 1859,
only one was a female. In corroboration, Mr Charles Oldham
writes me that of fourteen bats taken by Mr Coward and himself
from the Cefn Cave, near Denbigh, on 4th March, twelve were
males and two females. Further observation will no doubt
show whether the numbers of the sexes are unequal,! or whether
Foot and Kinahan’s experiences merely indicated that the adult
females had departed to bring forth their young apart from the
males. On the other hand, Messrs Coward and Oldham inform
me that they have seen males and females hanging together in
March, April, and on 2nd December. The idea that the males
outnumber the females, although at first sight improbable, has
gained in importance from Mr C. B. Moffat’s? suggestion—
based partly on a study of birds and insects in Ireland—that
fertility may decrease and females may become scarce as the
outskirts of the geographical range of a species is reached. It
is quite in keeping with the known disagreement between the
observations of continental and of British naturalists as regards
the number of young in bats.

Another point suggested by the work of Foot and Kinahan
is that, like other species, the bats are not necessarily to be
found in the same retreats in summer as in winter. One cave,
that of Balliallia, was entered twice in March of two different
years and once in August, and, although tenanted by bats
in the former, was empty in the latter month. There is,
however, probably no definite rule about this habit, and the
reverse was found to be the case at Cheddar, Somerset, by
Mr Coward.

Foot and Kinahan may be said to have demonstrated with
considerable accuracy the general features and characteristics
of the cave-life of these bats; in fact, the only point in regard
to which later observers have shown them to be in error is in
the supposed necessity for woods and plantations in the neigh-
bourhood of the bat-caves. Both Mr Coward and Mr Oldham

? It is suggested by E. Hollis for Devonshire, see Zoologist, 1907, III.
* Irish Naturalist, 1907, 140-144.

260 RHINOLOPHIDAEK—RHINOLOPHUS

have supplied me with independent information that they
have found the bats inhabiting caves opening on to bare
hillsides.

At Cefn, on 18th November, with a cave temperature of 46°
Fahrenheit, Mr Coward! found the bats so wakeful that two
were on the wing, and these were joined by three others on his
disturbing them. The excrement on the ground beneath them
seemed to indicate that they had fed after retiring to the
cave, especially as the remains of the cave-haunting spider”
showed that this species had formed part of their diet. Other
observations to the same effect have been made by Messrs
Newstead and Oldham, who have attempted to ascertain
the exact duration of the winter retirement. On this point
Kinahan in Clare and Mr Coward in Denbighshire found
the caves untenanted in July and August, but the bats
were at home in the Irish caves on 1st April, and in the
Welsh ones on the 4th of the same month.* As regards the
autumn, Mr Oldham,* who has had specimens taken on 24th
September from a cave in Carnarvonshire, suggests that this
haunt may have been occupied throughout the summer months.
In any case it is clear that the bats frequent the caves where
they hibernate at least from some time in November to some
time in April, and that during that time they frequently shift
their positions within the caves, but whether or to what extent
they ever leave them is uncertain. ‘Even during their period
of activity,” writes Mr Oldham, ‘from spring to autumn the
diurnal sleep of bats is profound. The heart beats feebly,
respiration is diminished, and the temperature falls. The
creature, if taken in hand, feels cold and lifeless, and some
time elapses before, with sneezes and spasmodic twitchings of
the limbs, it emerges from its lethargy, an epitome of restless
activity and pulsating life. The phenomena of normal sleep
are intensified during hibernation; but, even if other proof
were lacking that the winter sleep is broken, it is highly im-
probable that the vital functions would be so far in suspense
that food taken immediately prior to the Bat’s retirement in
autumn would remain unassimilated in the stomach, or as

' See article on Greater Horseshoe, supra. 2 Meta menard.
3 In Kent’s Hole, Devonshire, on the 8th (Oldham). * Zoologist, 1903, 430.

PLATE XIX.

GREATER HORSESHOE BATs.

THE LESSER HORSESHOE BAT 261

fecal matter in the intestines, until the resumption of an
active existence in spring.”

As in the case of the previous species, the observations of
Messrs R. Rollinat and E. L. Trouessart in the Department
of Indre, France, throw much light on the breeding habits.
They find, for instance, that during the period of gestation
and of rearing the young, the breeding females of these delicate
creatures—the most delicate, as the authors think, of all
European bats—forsake the companies of powerful Greater
and Euryale! Horseshoes with which they have spent the
winter. They are often accompanied by males of their own
kind, but, apart from these, the only strangers allowed to asso-
ciate with them are a few Notch-eared Bats.”

Unfortunately, little is known of these important matters
for this country. Mr T. W. Proger*® had one born in captivity
on 20th June in South Wales, and several females which
he examined were about to become mothers at that date.
On the other hand, Mr Whitaker* found fully-developed
embryos in dead females from Somerset on 14th and 24th
July. The breeding season is, therefore, a long one, but in
other respects is probably similar to that of the larger species.
In France, Messrs Rollinat and Trouessart find that this is
the case, but the winter retreats are deserted at a slightly
later date, so that the period of gestation and the date of
birth of the young are delayed accordingly.

In France the first young one was found with its mother on
13th July, but others had evidently been born at some time
previously. The mothers, if frightened, will deposit their
young in a safe place and attempt to escape without them.
By 25th August lactation seems to have finished, and the
young are nearly as big as their mothers, who are now in poor
condition. Soon afterwards the nursing parties disperse to
the caverns, where the winter is spent; in September they
regain their condition, and in October they pair again.

These authors remark on the frequent presence at all
seasons of very thin individuals, in contradiction to the fact
that both sexes of the Greater Horseshoe are always very fat.

1 R. euryale of Blasius. 2 Myotis emarginatus.
MRE 4 Naturalist, March 1907, 74, etc.

262 RHINOLOPHIDA—RHINOLOPHUS

They attribute this to the fact that the Lesser Horseshoe is
delicate, and does not possess very vigorous powers of flight ;
they have satisfied themselves that the least wind interferes
enormously with its evolutions, so that it frequently remains
at home and fasts at times when other species are able to catch
their food.

Messrs Rollinat and Trouessart made the interesting
observation that when a number of mothers and their young
are placed together after capture, the young travelled from one
to the other, and were received by all. This fact made the
authors think that, unlike other bats, such as the Mouse-eared,
the females of this species may possibly assist each other in
rearing the young.

So far no one has succeeded in keeping Lesser Horseshoes
alive in confinement for more than a short time. They rarely pro-
gress sufficiently in domesticity as to consent to eat, and, asa
rule, pine away within three or four days’ time. Writing to Mr
J. G. Millais of one which he succeeded in keeping alive for
four days, Mr Coward remarked that it ‘took readily to meal-
worms, after the juice of one had been smeared on its face ;
the first day it would eat only half a mealworm, but the next
day it took six fair-sized-worms and a small caddis-fly. It
would not touch this fly until it had eaten the grubs. This
bat found great difficulty in masticating the hard skin of the
worms, its teeth being small and its jaws feeble, but it took
to them well and asked for more, biting my fingers and even
taking a bit of my jacket in its mouth. It drank water from
the end of a camel’s-hair brush, and also took a little milk.”
When resting on the table the tail was held above the back,
and when it was struggling with a large mealworm there
was no movement of the tail perceptible.

When frightened or attacked, the Lesser Horseshoe Bat
squeaks loudly, and its voice has been divided by Mr Coward
into two cries, the one a short, sharp ‘“‘chap” or “chip,” and
the other a “chatter,” each of much lower pitch than those of
most bats. Mr Oldham has likened the former cry, which is
uttered both while on the wing and when at rest, and which he
renders as ‘‘tchek, tchek,” to a diminutive of the alarm note
of the greater spotted woodpecker.

THE LESSER, HORSESHOE BAT 263

Although easy to recognise when captured, the identifica-
tion of this bat when on the wing is more difficult; but here
again its size, fluttering flight, with frequent gliding intervals,
and its light colour, are a combination of characters peculiar to
the species.

iy
Yee
1 Fp

ee.

PLATE XX.

LEssER HORSESHOE BaTs. (Natural size.)

0
wine ae
healed: he
’ 4 * st

win Vii

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