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PART L— October. Price 2s, ecf. net. 

A HISTORY OF 

BRITISH MAMMALS 



BY 

GERALD E. H. BARRETT-HAMILTON 

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



W/r// TWENTY-SEVEN FULL-PAGE PLATES IN COLOUR, FIFTY-FOUR IN 

BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND 

FIFTY SMALLER ILLUSTRATIONS 

DRAWN BV 

EDWARD A. WILSON 

B.A., M.B. (Cantab.) 




^\S'\o'1 



GURNEY AND JACKSON 
10 PATERNOSTER ROW, LONDON, E.G. 

1910 



In Pt&ntaratioPi 



A NEW AND REVISED EDITION OP 
YARRELL, NEWTON, AND SAUNDERS' 

History of British Birds 

EDITED BY 

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

Keeper ol the Natural History Department. The Royal Scottish Museum; Member of the 

British Association Committee on the Migration ol Birds as Observed on the British 

and Irish Coasts; Correspondlnj: Fellow of the American Ornithologist*' Union; 

Correspondirender Mitglied de» Ornithologlschen Vereins in WIen ; 

Membre Honoraire du Bureau Central Ornithologique Hongrois ; 

Member ol the British Ornithologists' Union, etc. 

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF BACH SPECIES 
SPECIALLY EXECUTED BY 

MISS LILIAN MEDLAND 



THE 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 



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 IN 

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 



GURNEY AND JACKSON 

10 PATERNOSTER ROW, LONDON, E.G. 

1910 



Temporary Title Pagc.^ 



OEG 2 1910 '^' 



FOREWORD. 

MR OLDFIELD 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 larofe 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." 



PRELIMINARY ANNOUNCEMENT. 

PpOR many years Tomes and Alston's edition of Bell's 
History of British Qtiadr^tpeds 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 Zoology. 
— 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 Design of the Work. — 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 

a 2 



VI 



PRELIMINARY ANNOUNCEMENT 



the chancre of view between 1874 (the date of Bell's last edition) 
and 19 10. 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. 

Division 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. 

Habits and Life History. — The habits will be fully 
traced, thus making the popular side of the subject very 
complete. The literary and historical aspect is n(3t 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. 

Technical 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 Paper. — 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 of Publication. — 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 i8th 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.— Flying Mammals. 



Order CHIROPTERA, or BATS. 



Vespertilionid/E, or Typical Insect-eating Bats. 



Genus Nyctalus. 
Noctule Bat. 
Leisler's Bat. 

Genus Pipistrellus. 
Common Bat, or Pipistrelle. 

Genus Vespertilio. 
Serotine Bat. 
[Parti-coloured Bat (non-British).] 

Genus Myotis. 
Daubenton's, or the Water Bat. 
[Rough-legged Water Bat (non- 
British).] 



Genus Myotis — continued. 
Whiskered Bat. 
Bechstein's Bat. 
Natterer's Bat. 

[Notch-eared Bat (non-British).] 
[Mouse-eared Bat (non-British).] 

Genus Plccotus. 
Long-eared Bat. 

Genus Barbastclla. 
Barbastelle Bat. 



[The Hoary Bat (non-British).] 



Rhinolophid/E, or Horsesiioe-nosed Bats. 



Genus Rhinolophus. 
Greater Horseshoe Bat. 



Genus RJiinolophus — continued. 
Lesser Horseshoe Bat. 



CONTENTS 



VOLUME 11. — Land Mammals. 



Order INSECTIVORA. 



Genus Erimiceus. 
Hedgehog. 



Genus Talpa. 



Mole. 



Genus Sonw 
Common Shrew. 
Pygmy Shrew. 

Genus Neoviys. 
Water Shrew. 



Order RODENTIA. 



Genus Lepus. 

Brown Hare. 

Scottish Varying or Blue Hare. 

Irish Hare. 

Genus Oryctolagiis. 
Rabbit. 

Genus Sciurus. 
British Squirrel. 

Genus Muscardinus. 
Dormouse. 

Genus Micromys. 
Harvest Mouse. 



Genus Apodciniis. 

Long-tailed Field Mouse. 

Hebridean Field Mouse. 

Fair Island Field Mouse. 

De Winton's Yellow-necked Field 
Mouse. 

St Kilda Field Mouse. 

Genus Mus. 
House Mouse. 
St Kilda Mouse. 
Faroe Island Mouse. 

Genus Epiuiys. 

Black Rat. 
Brown Rat. 



CONTENTS 



XI 



Volume II. — Land Mammals — continued. 



Order ^OVi^^T\ K— continued. 



Genus Ai"vicola. 
Water Vole. 

Genus Microtus, 
Short-tailed Field Vole. 
Hebridean Vole, 
Orkney Vole. 



Genus Microtiis — continued, 

Sanday Island Vole. 
Westray Island Vole. 

Genus Evotoinys. 
Red or Bank Vole. 
Skomer Vole. 



Order UNGULATA. 

Genus Bos. j Genus Cervtis — continued. 

[White Park Cattle : domesticated,] [Fallow Deer : introduced.] 



Genus Cervus. 



Red Deer. 



Genus Capreolus. 



Roe Deer. 



Genus Felis. 
Wild Cat. 



Order CARNIVORA. 

Sub-order Flssipedia, or Land Carnivora 

Genus Mustela. 
Weasel. 

Common Stoat. 
Jura Island Stoat. 
Irish Stoat. 

Genus Putorius. 
Polecat. 

Genus Martes. 

Pine Marten. 



Genus Vulpes. 
Fox. 

Genus Meles. 
Badger. 

Genus Lutra. 
Otter. 



Xll 



CONTENTS 



VOLUME III. — Aquatic Mammals. 



Order CARNIVORA. 



Sub-order Pinnifedia, or Seals. 



Genus PJioca. 

Common Seal. 
Ringed Seal. 
Greenland Seal. 

Genus Cystophora. 
Hooded Seal. 



Genus Halichcerus. 
Great Grey Seal. 

Genus Erignatus. 
Bearded Seal. 

Genus OdobcBnus. 
Atlantic Walrus. 



Order CETACEA, or WHALES. 
Sub-order Mystacoceti, or Whalebone Whales. 



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

Genus Megnptera. 
Hump-backed Whale. 



Genus Balcenoptera. 
Common Rorqual. 
Sibbald's Rorqual. 
Lesser Rorqual. 
Rudolphi's Rorqual. 



Sub-order Odontoceti, or Toothed Whales. 



Genus PJiyseter. 
Sperm Whale, or Cachalot. 

Genus Hyperoodon. 
Common Bottle-nosed Whale. 



Genus Ziphins. 
Cuvier's Whale. 

Genus Mesoplodon. 
Sowerby's Beaked Whale. 



CONTENTS 



XIII 



Volume 111. — Aquatic Mammals — continued. 



Order CETACEA, or ^YiMJ^'^— continued. 



Sub-order Odontoceti, or Toothed Whales — continued. 



Genus Monodon. 
Narwhal. 

Genus Delphinapterus. 
White Whale. 

Genus Orcinus. 
Killer. 

Genus Pseudorca. 
False Killer. 

Genus Grampus. 
Risso's Grampus. 



Genus Globicephala. 
Ca'ing or Pilot Whale. 

Genus Pkoccena. 
Common Porpoise. 

Genus Delphinus. 
Common Dolphin. 

Genus Tursiops. 
Bottle-nosed Dolphin, 

Genus LagcnorJiynchus. 

White-sided Dolphin. 
White-beaked Dolphin. 



Extinct Mammals. 



APPENDIX. 

I Domestic Mammals. 



LIST OF FULL-PAGE PLATES IN 
COLOUR 



Noctule Bat. 

Daubenton's Bat. 

Bechstein's Bat. 

Natterer's Bat. 

Common Shrew. 

Pygmy Shrew. 

Water Shrew. 

Pine Marten. 

Stoat in summer. 

Weasel and Stoat, to show 
pelage. 

Stoat and Weasel in winter. 

Scottish and Irish Hares. 

British Squirrel. 



Seasonal changes in British 
Squirrel. 

Dormouse. 

British P'ield Mice. 

British House Mice and allies. 

British Rats. 

Orkney Vole. 

British Voles (i.). 

British Voles (ii.). 

Red Deer. 

Roe Deer. 

Common Seal. 

Bottle-nosed Whale. 

Killer. 

Common Dolphin. 



LIST OF FULL-PAGE PLATES IN 
BLACK AND WHITE 



Noctule Bat. 

Leisler's Bat, 

Pipistrelle Bat. 

Serotine Bat. 

Whiskered Bat. 

Long-eared Bat. 

Barbastelle Bat. 

Greater Horseshoe Bat. 

Lesser Horseshoe Bat, 

Hedgehog. 

Mole. 

Brown Hare. 

Scottish Hare. 

Rabbit. 

Harvest Mouse. 

Long-tailed Field Mouse. 

Water Vole. 

Short-tailed Field Vole. 

Red Deer. 

With 250 Line 



Roebuck. 

Polecat. 

Wild Cat. 

Fox. 

Badger. 

Otter. 

Grey Seal. 

Ringed Seal. 

Harp Seal. 

Bearded Seal. 

Hooded Seal. 

Walrus. 

Hump-backed Whale. 

Sperm Whale. 

Porpoise. 

Ca'ing Whale. 

Twenty Plates of Whales (semi- 
diagrammatic). 

Drawings, including Maps. 



CONTENTS OF PART I. 

PAOES 

General Introduction to Bats . . . .1-8 

(To be completed in forty-eight pages.') 

Vespertilionid^ ....... 49 

Genus Nyctalus . . . . • .52 

The Noctule, or Great Bat ..... 58 

Leisler's Bat ........ 83 



ILLUSTRATIONS 

Dormouse. (Coloured.) 

Noctule Bats. 

Heads of — (i) Nyctalus noctiila, (2) Nyctalus Icisleri, and (3) Vespettilio 
serotinus, 

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

Wings of — (i) Nyctalus leisleri, (2) Pipistrellus pipistrellus, and (3) Myotis 
daubentoni. 



CHIROPTERA. 

BATS. 
Sub-order MICROCHIROPTERA. 

INSECT-EATING BATS. 



CHIROPTERA. 

BATS. 
Sub-order MICROCHIROPTERA. 

INSECT-EATING BATS.i 

The twelve British species are all included in the great sub- 
order of insect-eating bats or Microchiroptera. They fall into 
two families — the Vesper HlionidcF, or typical simple-faced, and 
the Rhino lop hi dcB, 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 Pinax [\66o), and Ray, in 
his Synopsis (1693), made no attempt to diagnose species, but 
Albin, in his Natural History of Birds'^ (1740), gave a place 
to three — the "small common sort," the Double-eared (namely, 

* For many references to the literature of these animals, see N. H. Alcock's 
paper on the vascular system of bats in Froc. 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 igo8, 135-193). 

- iii., 95. 

3 



4 INTRODUCTION 

the Long-eared), and a flying fox, the latter, of course, exotic. 
Pennant in his earHer editions (1766 and 1768) recognised only 
two, which he called the Short-eared and the Long-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 Quadrupediuu (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 Vespej-tilio altivolans. In his 
eleventh letter, dated 9th September 1767, he had noticed the 
common Vespertilio murinus ^ and the Vespertilio mcribus, 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 ^^nd 
Pennant in 181 2 were able to enumerate only half the species 
now catalogued. Montagu's name mimitus, 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 

* Called "the little bat" in letter xxvi. of 8th Dec. 1769. 

2 "A List of the Species of Vespertilionidce found in Great Britain," Zoological 
Journal, ii., 108-110, 1826. 

^ Myotis myosotis (Bechstein). 



CHIROPTERA 5 

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 Nycialus, the 
high-flying swifts of their kind, to the head of their tribe, with 
Myotis as the most primitive vespertilionid genus, and the 
family Rhino lop hi da; 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 ScotophiltLS 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, Pipistix litis, 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 Nyctalus hitherto associated 
with the fruit bats. Thus Nyctahis replaces Pterygistes 
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. : — Nyctalus with two species ; Pipistrellus with 
one ; Vespertilio with one ; Myotis with four ; Barbastella with 
one ; and Plecottis with one. The Rhino lop hides, with one 
genus [Rhino lop htis) 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. i) will be 
found useful. 

In all British bats except the Rhinolo- 

phidce 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 

^'^•VA?rEAr°'^ ear. At the base of the outer or opposite 

a^ tragus; margin, and especially conspicuous in the 

^, antitragus ; RJiinolophidcB, ariscs a lobe known as the 

<:, anterior margin : .•. '-t->i i i • r i 

«^. posterior margin. antitmgus. The shapc 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. In a series of 




CHIROPTERA 











Fig. 2. — Ears of British Bats, except Plecotus auHtus (diagrammatic). 



1. Nyctalus noctula. 

2. N. leisleri. 

3. Pipistrellus pipistrellus. 

4. Vespertilio serotinus. 



5. Myotis dauhentoni. 

6. M. mystacinus. 

7. M. bechsteini, 

8. M, nattereri. 



9. Barbastella barbasiellus. 

10. Rhinolophus ferrum-eqmnum. 

11. R. hipposideros. 



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, 




Wing of a Bat (diagrammatic). 



Membranes : — am, antebrachial ; im, lateral ; im, interfemoral ; dp, dactylopatagium ; 
pc/, post-calcarial lobe. Skeleton ; — /i, humerus ; r, radius ; «, ulnar ; u\ wrist ; di, 2, 3, 4, 5, 
digits I, 2, 3, 4, 5 ; mi, 2, 3, 4, 5, metacarpals i, 2, 3, 4, 5 ; / femur ; //>, tibia ; /fi, fibula ; 
/, 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 

' See Andersen, Ann. and Mag. Nat. Hist., Nov. 1906, 372-373. 



PART IL— November. Price 2s. Sd. net, 

A HISTORY OF 

BRITISH MAMMALS 



BY 

GERALD E. H. BARRETT-HAMILTON 

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



Wrri/ 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.S. (Cantab.) 




GURNEY AND JACKSON 
10 PATERNOSTER ROW, LONDON, E.G. 

1910 



%\sizi 



fn Preparation 



A NEW AND REVISED EDITION OP 
TARRELL, NEWTON, AND SAUNDERS' 

History of British Birds 



EDITED BY 

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

Keeper of the Natural Hixtory Department, The Royal Scottish Museum; Member of the 

British Association Committee on the Misrration of Birds as Observed on the British 

and Irish Coasts; Corresponding Fellow of the American Ornithologists' Union; 

Correspondirender Mitglied des Ornitholosischen Vereins in Wien ; 

Membre Honoraire du Bureau Central Ornitbologique Hongrois ; 

Member of the British Ornithologists' Union, etc. 

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES 
SPECIALLY EXECUTED BY 

MISS LILIAN MEDLAND 



THE 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 



CONTENTS OF PART II. 

PAGES 

General Introduction to Bats .... 9-16 

( To he completed in forty-eight pages.) 

Vespertilionid^ — 

Genus Nyctalus — 

Leisler's Bat ....... 89 

Genus Pipistrclhis . . . . . .100 

The Common Bat, Pipistrelle or Flitter-Mouse . . 103 

Genu.s Vespertilio . . . . . .127 



ILLUSTRATIONS 

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

Portions of Finger Skeletons of Bats (diagrammatic), enlarged to show 
Stages in Ossification of the Joints. 

Fig. I. — Typical Molar Teeth of an Insectivorous Bat (after Miller). 
Fig. 2. — Young Pipistrelle. 

Leisler's Bats. 

Pipistrelle Bats. 



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. 




II. 


I ( Vespertilionidce) 


nil {Rhinolophidce) 


III. 


3 n 


2 


IV. 


3 „ 


2 


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 suffi- 
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 RhinolophidcF. 

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. 

B 



lo 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 9th 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 as a 
feature of high diagnostic value. 

The actual dimensions of the various species are shown 

' This is not a universal characteristic of the CJiiroptera^ 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:— 












i 



"3 


^ -.1 
— 


1 


5:3 


0. 
t-< . 
cS>-i 

CD 


75 

1 




Nyctalus noctula 
5 to 7 males 
7 to 8 females . 


77-5 

77 


50 
51-5 


4;t-5 
[49] 


17-5 
17-5 


12 

12 


91-5 
92-5 


48-25 
50 


38 
40 


[360] 
[375] 


N. Uisleri 

10 of both sexes 


[596] 


42 


[41] 


17 


8 


[72-5] 


38-5 


31-5 


[300] 


Pipistrellus pipistrdhis 
9 to 12 males 
5 to 11 females . 


42 
43-35 


29-5 
30-28 


31-85 
30-5 


[10-5] 
11-36 


5-65 
7-14 


50 
50 


25 
25 


25 
25 


[-208] 
213 


Vespertilio serotinus 

7 males .... 
11 of both sexes 


73 

74-2 


52-8 
52-1 


52-0 
52-3 


20-8 
20-8 


16 -5 


86-5 
86-6 


47 
47 


43-5 
44 


368 
300 


My Otis daubentoni 
19 of both sexes 


45-5 


80-5 


33-5 


16-4 


9-1 


59-8 


32-2 


30-2 


223-8 


M. mystacinus 

6 males .... 


[46] 


34 


[82] 


15-5 


[7-5] 


[53-0] 


[29] 


[28] 


[2'28] 


M. bechsteini 

5 


[55] 


41 


[35] 


19-5 


(?) 8-10 


64 


33-5 


33 


[250] 


Jlf. natter eri 

15 females .... 


45-1 


39-4 


37-15 


16'5 


8 


65-6 


33-1 


33 


[274] 


Plecotiis oMrif j(s 

10 males .... 
17 females .... 


48-2 
45-7 


37-5 
38-8 


41-(3 

42-2 


17-3 
18-3 


S-8 
8-8 


62-2 
65 


31-3 
32 


30 1 
30-9 J 


[255] 


Barbastella barbastellus 
6 males .... 


[48] 


37-6 


43-4 


18-6 


6-7 


66-4 


34-8 


32-5 1 


[250 
-260] 


Rhinolophus ferrum-cquinum 
11 males .... 
5 females .... 


63-2 
05-8 


53-0 
54-5 


32 
33-2 


24-5 
25-3 


11-2 
11-8 


83-4 
85-8 


36-4 
36-2 


38-9 
40-5 


334-6 
[360] 


R. hipposideros 


[38] 


37-5 


25-2 


17-5 


S 


[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 .02 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. 


Nyctahis noctuJa 


•13 


.6 


,, leisJeri 


.14 


•7 


Pipistrelliis pipistrelliis 


.14 


•7 


Vespertilio serotinus 


.14 


.7 


Myotis daiibentoiii 


.16 


.8 


„ mystacimis 


.149 


•7 


,, hcchsteini 


.16 


•7 


,, natter eri . 


.14 


.8 


Plecotus auritus . 


.14 


•7 


Barhastella barhastelhis 


.14 


•7 


Rh ino loph iis ferru m -eq it in ii m 


1 I^ 16 


.8 


,, hipposideros 


1 -'O •^'-' 





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 Nyctahis 
noctula and Pipistrellus the lower leg is shortest. The foot is 
longest in Myotis daubentoni and JV. noctula, and shortest in 
Barbastella. The expanse is greatest in Nyctahis, Pipist^'elluSy 
Vespertilio, and Myotis nattere^d in the order named, and 
least in M. daubentoni. A narrow wing is indicated in 
Nyctalus, and less so in Vespertilio and Barbastella, by the 
elongated third metacarpal, considerably longer in each case 
than the fifth. The converse is the case in Pipistrelhis and 
M. nattereri, in which the two metacarpals are about equal. 
Barbastella 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 :— 





S 


'S 








'to 
'■3 

bo 

a 



"3 


"3 


p. 


Nyctalus nor.tula 

7 males .... 

N. IHsleri 
Both sexes 

Piinstrelhis pipistrellus 

12 males .... 
11 females .... 

Vespertilio serotintis 
10, both sexes . 

Myotis dauhentoni 

13 males . . . } 
6 females . . . / 

M. viystacinus 

6 males .... 

Af. bechsteini 

5 

M. nattereri 

15 females .... 

Plccotus aurit^is 

10 males .... 
17 females .... 

Barbastella barbastdlus 
6 males .... 


100 

100 

100 
100 

100 

100 

100 

100 

100 

100 
100 

100 


99 
[97] 

108 
100 

100 
91 

[S5] 

95 

111 
109 

114 


35 

40 

[35] 
37 

39 
44 

45 

47 

42 

46 
47 

49 


24 

19 

19 
23 

20 

25 

[22] 

[19] 

20 

23 
21 

18 


183 

[172] 

169 
165 

166 

163 

[157] 
154 
106 

166 

167 

176 


96 

91 

84 
82 

90 

88 

[S5] 
81 
84 

83 

82 

92 


76 

75 

84 
82 

84 

83 

[82] 
80 
84 

80 1 

79 / 

86 


[720] 

[714] 

[705] 
703 

690 
613 

[605] 
[695] 

[680] 

[678] 


Average ") 
(excluding M. bechsteini) f 




101 


42 


21 


168 


87 


81 


088 


Rhinolophiis ferriim-equinum 
11 males .... 
5 females .... 

R. hipposideros 


100 
100 

100 


60 
61 

67 


40 
46 

47 


21 

22 

21 


155 
157 

[139] 


66 

67 

68 


72 
74 

71 


624 

[642] 

[653] 



The few fio-ures available for Af. bechsteini indicate an 
exceptional bat ; short-tailed, but with long lower leg and broad 
short wing. 

T\\& Rhino lop hides 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 W^ter 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 Rhino lop hid(£, 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, op. cit. 
infra, p. 22. 



CHIROPTERA 15 

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 Myotis it is — 

.2-2 i-i . 2-2 

mi , mc , mpm =22, 

3-3 I- I ^ 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 Myotis to 
thirty-two, as in Vespertilio 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 
formulae, 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 Rhino lop hi dcE, 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 Myotis possesses 
the lowest type of dentition, the full formula in this case being — 

pm ^ — ^, m ^ — ^ = 38. 



3-3 i-i 3-3 3-3 



1 For further details see L. F. E. Rousseau, "Mem. Zool. et anatomique sur la 
Chauve-Souris commune," etc., in Guerin's Mag. de Zool., 1839, pi. 7 ; also Wilhelm 
Leche in Lund's U?nversitets Ars-skrift, xii., 1-47, 1876, pis. i and 2 (analysed in 
Archiv filr Naturgeschichie, i., 1877, 353-364), and xiv., 1-37, 1878, pis. i and 2. 



i6 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 

upper premolars, with pm _ ^ and a total of thirty-six teeth. 

In PipistTelliLS, the next highest, a pair of lower premolars has 
vanished (both remaining pairs being quite small), and the 

formula is pm — ^^, 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 Vespertilio the former 
tooth has entirely disappeared, reducing the total to thirty-two, 
all of which are functional. In Rhinolophtis the number is 

again thirty-two, but in this case the premolars are , with 

the anterior upper and central under often rudimentary : the 

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 i and premolar i 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} 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. Hist.., April 1908, 346-348. 



HISTORY OF BRITISH BIRDS— continued. 

histories of very many others, which were formerly little known, 
have 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- 
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 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 
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 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- 
known bird artist. 



10 PATERNOSTER ROW, LONDON, E.G. 



In th e Press, to be published shortly 

STUDIES IN BIRD- MIGRATION 

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

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 Illustrations and Maps 



WITH 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 
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. 

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 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), 
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. 



10 PATERNOSTER ROW, LONDON, E.G. 



Oliver and Boyd, Printers, Edinburjrh. 



PART iJL—December, Price 2s, 6d. net. 

A HISTORY OF 

BRITISH MAMMALS 



BY 

GERALD E. H. BARRETT-HAMILTON 

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



W/TH 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. CCantas.) 




GURNEY AND JACKSON 

10 PATERNOSTER ROW, LONDON, E.C. 

1910 



\/lvf>^74-4- 



In Preparation 



A NEW AND REVISED EDITION OP 
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 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 



THE publication of Yarrell's "History of British Birds" was 
commenced in 1837 ^^^ 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 



CONTENTS OF PART III 



General Introduction to Bats 

{To he completed in forty-eight pages.') 



17-24 



VeSPERTILIONID/E — 




Genus Vesper tilio .... 


129 


The Serotine ..... 


• 130 


The Parti-coloured Bat 


• 139 


Genus Myotis ..... 


140 


Daubenton's, or the Water Bat 


143 


[The Rough-legged Water Bat] 


157 


The Whiskered Bat . 


. 158 



ILLUSTRATIONS 



Full-page {Coloitrcd aiid Black and White). 

Natterer's Bat. {Colotired.) 

Wings of — (i) Nyctalits nocttila ; (2) Vcspertilio serotinus ; (3) Rhino- 
lophus ferniin-equiniivi. 

Serotine Bat. 

Heads of — (i) Myotis mystacinns ; (2) Myotis danhcntoni ; (3) Afyotis 
natter eri ; (4) Pipistrellus pipistrellus. 

Whiskered Bat. 

Figures in Text. 

Diagram of Arrangement of Teeth of Vespertilio serotinus. 
Front View of Incisors and Canines of Vespertilio serotinus. 
Diagram of Arrangement of Teeth in Genus Myotis. 
Front View of Incisors and Canines oi Myotis daiibentoni. 
Front View of Incisors and Canines of Myotis mystacinus. 



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. i). 

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. i). 

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 to a 
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 

^ For further details see Herluf Winge, "Jordfundne og nulevende Flagermus 
{Chiroptera), fra Lagoa Santa, Minas Geraes, Brasilien," from E. Mtiseo Lundii, vol. 
ii., paper i, 1893. 

C 



i8 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 Scotophilus 
ornahis (Blyth). An African false vampire [Lavia 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 Hipposiderus 
fulvus assumes a vivid coloration during the breeding season. 
In Scotophihis kuklii, 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 

* E.g., Thomas' Rhinolophus denti of Kuruman, South Africa {Ann. Mag. Nat. 
Hist.., May 1904, 386) ; and his Vespeftilio matschiei of Aden {An7i. Mag. Nat. 
Hist., Nov. 1905, 573. 

- E.g., Miniopterus dasythrix of Temminck and M. 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.^ 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, op. 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 occupy 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 

* Daubenton's, Natterer's, Bechstein's (once), and Lesser Horseshoe (once). 
2 Bull. Soc. Amis. Set. 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, 7ioctula, 
pipistrellus, datibentoni, and ambitus are regularly met with 
in all five, but two more, mystacinus ^ and iiattereri,^ differ only 
in their apparent absence from Scotland. Two, leisleri 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-equimt7n is southern and south- 
western, while serotums is distinctly south-eastern. Lastly, 
two, murimis and nilssoni, are found only in Skandinavia and 
Denmark, while two more, myosotis and emarginattis, 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 

' I exclude from the Skandinavian fauna, for the purposes of this article, Pipi- 
strellus nathusii and Myotis dasycneme; the status of either is doubtful. 
2 With two Scotch records. ^ 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. 
Bechsteini and nathusii 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 

^ 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 {pp. 
cit., 155, 466, 486, 549), or catch shrimps, fish {op. cii., 397), small birds or mice 
(A. M. Primrose, also E. Gleadovv, y<9«r«. Bombay Nat. Hist. Soc, xvii., 1021-1022). 

^ Some important allusions to habits are scattered through Dobson's Catalogue of 
Chiroptera. P. H. Gos?,&'s Naturalist in famaica, 185 1, contains original observations 
on the bats of that island. His allusions to the crepuscular habits (p. J 62) 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 
Vespertilionidc^ 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 

' Dobson's Catalogue of Chiroptera, 160. - 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, 

^ 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 WoUey, Zoologist, 1846, 1204. 

3 G. J. Talbot, Field, loth Oct. 1903, 635 ; F. J. Montgomery, Field, 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, 453^ : 
in British Columbia the large rainbow trout leaps at and probably catches them — 
John Macoun, quoted by E. Thompson Seton, ii., 1181. 

* Zoologist, 1887, 426-427. ^ J. H. Wilmore, Zoologist, 1886, 242. ^ Op. cit. 



PART IV,— February, Price 2s. 6d, net. 

A HISTORY OF 

BRITISH MAMMALS 



BY 

GERALD E. H. BARRETT-HAMILTON 

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



Wrr// TWENTY-SEVEN FULL.PAGE PLATES IN COLOUR, FIFTY-FOUR IN 

BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND 

FIFTY SMALLER ILLUSTRATIONS 

DRAWN BV 

EDWARD A. WILSON 

B.A., M.B. (Cantab.) 




% 



(,Sb^ 



GURNEY -AND JACKSON 

PATERNOSTER ROW, LONDON, 5.C. 
1911 



In Preparation 



A NEW AND REVISED EDITION OF 
TARRELL, NEWTON, AND SAUNDERS' 

History of British Birds 



EDITBD BY 

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 Ornitholoxischen Vereins in Wien ; 

Membre Honoraire du Bureau Central Ornithologique Hongrois ; 

Member of the British Ornithologrists' Union, etc. 

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES 
SPECIALLY EXECUTED BY 

MISS LILIAN MEDLAND 



THE 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 



CONTENTS OF PART IV. 



General Introduction to Bats 

(To be completed in forty-eight pages.) 



PAGES 

25-32 



Vespertilionid.e — 




Genus Myotis — 




The Whiskered Bat . 


169 


Bechstein's Bat 


. 172 


Natterer's Bat 


. 178 


The Notch-eared Bat 


189 


The Mouse-eared Bat 


190 


Genus Plecottts .... 


192 


The Long-eared Bat 


194 



ILLUSTRATIONS 



Full-page (^Coloured and Black and White). 

Noctule Bats. (^Coloured.) 

Wings of — (i) Myotis mystaciniis ; (2) M. becJisteini ; (3) M. nattereri. 

Heads of — (i) Myotis bechsteini ; (2 and 3) Barbastella barbastellus 
(front and side views) ; (4) Plecotics aiiritits. 

Long-eared Bats. 

Figures in Text. 

Side View of Skulls and Teeth of — (i) Nyctalus Icisleri ; (2) Myotis 
becJisteini ; (3) M. nattereri. 

Front View of Incisors and Canines of Myotis bechsteini. 

Front View of Incisors and Canines oi Myotis ?iattereri 

Diagram of Arrangement of Teeth in Genus Plecotus. 

Front View of Incisors and Canines of Plccotiis auritiis. 



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- 

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

^ A remarkable instance is related by A. J. Dasent, Field, 7th Sept. 1889, 350. 

^ E.g., amongst others, Nyctalus azoreutn of the Azorean oak woods ; Rhyn- 
chonycteris naso of Amazonia and Guiana (E. A. Goeldi, Ibis, 1904, 518) ; Laviafrons 
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. cit., 140). 

D 



26 INTRODUCTION 

gregate, each as a 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 far as 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 vespertiKonid 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 manoeuvre 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 

^ Platymops viacmilani of Thomas, Ann. 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 Trans. 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 as a 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. It is 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 

^ 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, op. cit.^ 635. ^ F. Norton, Midland Naturalist^ 149, 1883. 

3 C. Nicolle and C. Comte state that of forty-seven examples of Pipistrellus 
kuhlit, marked and liberated at a distance of two kilometres from their retreat, six 
were eventually recaptured where originally taken.— Cf^w//". rend. Soc. Biol., Paris, Ix., 
738-739, 1906. 



28 INTRODUCTION 

settlinof down to the water surface. Others alter their beats 
but Httle 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, 
Loner-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 Op. cit. infray p. 32. ^ As noticed by Gilbert White. 

3 J. G. Millais; C. M. Smith, Field, 20th July 1889, 97 ; Lord Lilford, Zoologist, 
1887, 66 {M. mystacinus). * Op. cit. infra, p. 30. 





^'^. 





t C-'t^IJaorv. 



.^-^ 



NoCTUi.ii Dais. (ALuui '.. n.ituial size.) 



CHIROPTERA 29 

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 1 6th 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 Anitnals, ii., 1909, 1161. 

- Nycteris { = Lasiurus) semota. 

^ R. M. Barrington, The 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 etres organises, ii., 179, edition Jean Senebier, Geneva, 
1807. ^ Vesper tilio nilssoni. 



CHIROPTERA 31 

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 lyino- 
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 
1 89 1 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, 

^ Pipistrellus abramus. - P. nathusii. ^ Rhinolophus euryale. 

* Lakeland, 1892, i. ^ Infra, p. 223. 

^ B. Benecke, Zoologischer Anzeiger, ii., 1879, 304-305 ifiipistrellus and auritus) : 
Eimer, op. cit., 425-426 {ttoctula and pipistrellus): E. van Beneden and C. Julin, 
Archives de Biologie, i., 1880, 551-571, pls. xxii. and xxiii. {daubentoni, dasycneme, 
myosoiis, mystacinus, nattereri, emarginatus) : O. Van Stricht, Anatomischer Anzeiger, 
xix., SuppL, 1901, 208-210 [noctula, pipistrellus, myosotis): Fries, Ueber die Fort- 
pflanzung der enheimischen Chiropteren, Nachrichten, K. Gesellsch. der Wissensch, 
Univ. Gottingen, 1879, 295-298, {noctula, pipistrellus, nathusii, abramus, serotinus. 



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. DuvaP 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, mysfacinus, nattereri, bechsteini, auritus, barbastellus, hipposideros) : Carl 
Vogt, Association frangaise pour ravancement des Sciences, sess. lo, Alger, 1881, 
Conipte rendu, 655-662 : thus including all British species except leisleri. 

^ Etudes sur P Embryologie des Chiroptcres, i, Paris, 1879. For a description of 
copulation, which takes place while the animals are at rest, not in flight, see Hahn, 
op. ciL, 161; Victor Fatio, "Vertebres de la Suisse," i., 23, 1869; Duval, "Sur 
I'accouplement des Chauves-Souris," in Comptes rendus Soc. Biol., Paris, 23rd Feb. 
1895, 135-136- 

■^ "Sur la reproduction des Chauves-Souris," Mem. Soc. Zool. de France, ix., 214- 
240, 1896, 3 My Otis myosotis. 



PART v.— March, Price 2s, 6d. net, 

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 IN 

BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND 

FIFTY SMALLER ILL USTRA TIONS 

DRAWN BY 

EDWARD A. WILSON 

B.A., M.B. (Cantab.) 




^ 11 1 L 



GURxNEY AND JACKSON 

PATERNOSTER ROW, LONDON, E.G. 

1911 



In Pireparation 



A NEW AND REVISED EDITION OP 
YARRELL, NEWTON, AND SAUNDERS' 

History of British Birds 



EDITED BY 

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

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 ; 

Mcmbre 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 



THE 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 Pa^o 3 of Cover 



CONTENTS OF PART V. 

PAGE8 

General Introduction to Bats .... 33-40 

( To be completed in forty-eight pages.") 

Vespertilionid.e — 

Genus Plecottis ....... 209 

Gqwms Barbastella . . . . . .210 

The Barbastelle . . . . . .212 

The Hoary Bat ...... 222 

Rhinolophid.e — 

Genus Rhinolophus . . . . . .225 

The Greater Horseshoe Bat .... 228 



ILLUSTRATIONS 

Full- PAGE (Co/oured and Black and White). 
Squirrel. {Coloured.) 

Wings of — (i) Plecotus aiiritus ; (2) Barbastella barbastellus ; (3) Rhino- 
loplms hipposideros. 

Heads of — (i, 2, and 3) Rhinolophus hipposideros ; (4) Rhitiolophus 
ferrum-equinuni. 

Figures in Text. 

Diagram of Arrangement of Teeth in Genus Barbastella — (i) Upper 
and (2) Lower Jaw. 

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

Diagram of Nose-leaf of Rhinolophus. 

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

Diagram of Arrangement of Teeth in Rhinolophus ferruni-equinuin — 
(i) 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 9th June (a minimum of 
fifty-eight days if they emerged from hibernation on ist 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. 

^ Duval, " Etudes sur I'Embryologie des Cheiropteres," in Journ. de fanat. el de 
la Physiol., xxxi., 93-160, March to April 1895 ; but the authors once found a Serotine 
with an embryo in the left horn. 

E 



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 mammae 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- Sequard 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. 

' Op. cit. supra ^ P* j^- 



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 wino-. 

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 

' 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. 



36 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 i8th 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 eniarginatits. 

•■' E.g., Osburn, in the case of a captive Monophyllus, op. ciL, 84. 




Squirrel. (About I natural size.) 



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 
ingfuinal resfion. 

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 Nyctahts and Pipistrellus, 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 Barbastella alone I have no information. 

- See W. H. Hudson for Molossus bonariensis in The Naturalist in La Plata, 10 1- 
104 : Chapman and Hall, Limited, London, 1892 The Red Bat of N. America has 
four mammae, and one weighing 1 1 grammes has been taken alive nursing four 
young, weighing collectively 127 grammes (M. W. Lyon, jun., Proc. U.S. Nat. Miis., 
xxvi., 425-426, 1903). 

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

^ P. hesperus (H. Allen) ; P. abramus of Java has also been found with two 
embryos (see G. M. Allen, Bull. Mus. Comparative Zool., Harvard College, Hi., 3, 
July 1908, 45) ; and P. subflavus of N. America with three (Hahn, op. cit., 162). 

'" "The Problems of an Island Fauna," Irish 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 

' 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 1 10 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 {Set. Proc. Roy. Dublin Soc, IV., 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 hunsr 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 

' Article on Horseshoes, infra. 

- Zoologist^ 1897,46 ; T. P. Bartlett, /ir?//r«. r//., 1844,613 ; A. C. ?i\x(\X}[\, Journ. cit., 
1854, 4245. 

^ Act iv., sc. I. 



PART VL—lst May, Price 2s. Sd, net. 

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 IN 

BLACK AND WHITE, AND UPWARDS OF TWO HUNDRED AND 

FIFTY SMALLER ILLUSTRATIONS 

DRAWN BY 

EDWARD A. WILSON 

B.A., M.B. (Cantab.) 




GURNEY AND JACKSON 

PATERNOSTER ROW, LONDON, E.G. 

191 1 



In Preparation 



A NEW AND REVISED EDITION OP 
YARRELL, NEWTON, AND SAUNDERS' 

History of British Birds 



EDITED BV 

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

Keeper of the Natural History Department, The Royal Scottish Museumi; 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 Wlen ; 

Membre Honoraire du Bureau Central Ornithologique Hongrols ; 

Member of the British Ornithologists' Union, etc. 

ILLUSTRATED BY ORIGINAL COLOURED PLATES OF EACH SPECIES 
SPECIALLY EXECUTED BY 

MISS LILIAN MEDLAND 



THE publication of Yarrell's "History of British Birds" was 
commenced in 1837 ^^^ 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. M^ 

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

[Continued on Page 3 of Covet 



CONTENTS OF PART VI. 



PAOEB 



General Introduction to Bats {completioii) . . 41-48 

Rhinolophid^ — 

Genus Rhinolophus — 

The Lesser Horseshoe Bat ..... 250 

Note. — The Section on Bats being now complete, will — with a General 
Introductio7i to the whole work, Titles, Indexes, etc., to be published 
later— form Volwne I. 

VOLUME II. — Land Mammals 
Order INSECTIVORA 

TALPID.E ........ 2 

Genus Talpa ....... 2 

The Common Mole, Moldwarp or Want ... 5 

The English local names have been revised in part by Mr IV. W. 
Skeat, M.A. {assisted by Professor W. IV. Skeat), and in part by 
Mr C. M. Drennan, M.A. Lond., late Scholar Emm. Coll. Cajnb. ; 
the Celtic and Gaelic names by Dr E. S. Quiggin, M.A., Ph.D., 
Fellow a7id Lecturer in Moderti 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 
assistance has been rendered by Mr M. C. A. Hinton regarding 
extinct Mammals. 



ILLUSTRATIONS 

Full-page {Black and White). 

Barbastelle Bats {on coated paper). 

Greater Horseshoe Bats {do.'). 
Lesser Horseshoe Bats 

Figures in Text. 

Diagram o^T^ethoi Rhmolophus hipposideros — (i) Upper and (2) Lower 

Jaw. 
Side View of Teeth of Talpa Europcea. 
Encampment of Mole, 
Mole Excavations (18 Diagrams). 



MAY 27 1911 



CHIROPTERA 41 

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 : (i) 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 

^ Animal Life a7id Intelligence, 1890-91, 247. 

- 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 {op. cit., supra, vide p. 3). 

^ Messrs Rollinat and Trouessart, Comptes rendiis Soc. Biol., Paris, 23rd June 
1904, 1-4 (reprint). 

Y 



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 Nyctalus 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 Vespertilio and 
Myotis the ear and tragus are larger, but the glands are less 
extensively developed. Barbastella and PlecotuSy on the other 
hand, present a very remarkable extreme, both of ear and 
glandular development. Lastly, in Rhmolophus we see a simple 
ear without a tragus, but with well-developed antitragus, large 

' In the families VespertilionidcE and Rhinolophid(Z 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, Bull. Mus. Comparative Zoology (Harvard, 
U.S.A.), X., 3, 160, 1882-83. 

^ Spallanzani, De Jurine, Spadone, etc. See Edward Newman, Field., 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. Schobl, Archiv fiir Mikrosk. Anat., Bd. vii., i, pi. 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 {op. cit., 176). 



CHIROPTERA 43 

auditory bulla, and a special nasal organ of sense, the "horse- 
shoe." In all species the whiskers or vibrissse 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 probablycon- 
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, op. cit., 191. 



44 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 nigrht and fifteen hundred in the nit^ht 
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 

^ Gegenbaur's Morphologisches Jahrbiich^ xxxvii., i., 70-119, 1907, with plates. 
2 Vide Newman, Fields 27th Dec. 1873, 656. 

^ 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 
weicrht 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 Chiroptcra^ 83. 

- 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 : — 

A KEY TO BRITISH BATS. 

I. EXTERNAL CHARACTERS: 
I. Without Nose-Leaf : — 

1. Ears separated — 

(A) External ear-border terminating under the angle of the mouth — 

(a) Post-calcarial lobe prominent — 

(a') Tragus broadest at apex . NYCTALUS p. 52 

(a) Size larger, forearm | ^_ ^^^^^^^^^ ^ . p. 58 

rarely under 49 mm. .J r ^ 

(/3) Size smaller, forearm 1 ,, ^^.^^^^,. ^ . p. 83 

rarely over 44 mm. .J r ^ 

(b) rragusbroadestatcentre I pipistrellus J P' '°2 

(b) Post-calcanallobemsignificant I serotinus J P- ^30 

(B) External ear-border terminating^ i/fi^/nTrc ^ , .r^ 

under the base of the tragus .J r -r 

(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 \ M. daubentoni . p. 143 
ankle to tail . . J 
(/i) Calcar extending half- | ^j t^,i,,,,, . -^ 

way from ankle to tail J -^ v j 

(b') Ears long ; when laid for- \ 

ward extending far be- \ M. bechsteini . . p. 172 
yond nose-tip . . j 

(b) Posterior margin of inter- ^ ]/ fi. ,• ►.o 

femoral membrane fringed . j "^ ' ' • P* / 

2. Ears united at base — 

(A) Ears very long, much longer than f PLECOTUS \ 

head (about 34-38 mm.) . . ( auritus j P* ^"^ 

(B) Ears of moderate length (about ] BARBASTELLA\ 

13-16 mm.) . . . .\ barbastcllus / P* ^^^ 

II. With Nose-Leaf ..... RHINOLOPHUS p. 225 

1. Size larger, forearm 50-55 mm. . . R. ferrnm-eqninum p. 228 

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



BARBASTELLA\ 

barbastelhis i P' -^ 



- N. noctula . . p. 58 



CHIROPTERA 47 

II. TEETH AND SKULLS: 
I. Nasal Cartilages well developed : — 

1 . Four functional teeth in tOOth-row beliind upper canine — 

(A) No rudimentary cheek-teeth .( VESPERTILIO \ 

■^ y 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 

equal in height to third ; upper I KTyCTAT f/^ 

surface of rostrum convex, and ^cyo 

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 ,:r , . , . 

■section; greatest length f^-^^'^^^^'^ • • P- 83 

of skull about 15 mm. . j 

2. Five functional teeth in tooth-roW behind upper canine — 

(A) Five teeth in tooth-row behind f PIPISTRELLUS \ 

lower canine . . . . \ pipistrellus j P' ^^^ 

(B) Six teeth in tooth-row behind / PLECOTUS \ 

lower canine . . . . \ anritns ) ^' ^^"^ 

3. Six functional teeth in tOOth-row behind tipper canine — 

(A) Upper />/«. II. smaller, its tip about 

reaching cingulum oi pni. I. : — 

(a) Size larger, greatest length of \ ,. dauhentoni n 14^ 

skull about 15 mm. . . j ^'^- ^^^^oentom . p. 143 

(b) Size smaller, greatest length of 1 ,^ . . o 

skull about 13 mm. j M. mystacinus . p. 158 

(B) Upper pm. II. larger, its tip 

reaching well above cingulum 
of pm. I. : — 

(A) Upper /m II. about equal- | ^. 

hngpm. I. .... j r / 

(B) Upper/;;.. II. distinctly smaller j ^^ ^,,j,,teini . . p. 1 72 

than/;;z. 1 j v t 

IL Nasal Cartilages and Upper Incisors \ pfjrAjQTQpjjrj^ 

RUDIMENTARY j P* 5 

1. Greatest length of skull about 24 mm. . R. ferrum-equimim p. 228 

2. Greatest length of skull about 16 mm. . R. hipposideros . p. 250 



VESPERTILIONIDiE. 

TYPICAL INSECl-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 premaxillae, 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 Vespertilionidae 
followed in the present volume differs widely from that of Bell. 
The changes may be indicated by placing the two systems side 
by side : — 



BelVs system o/" 1874 : — System of present work : — 

ScOTOPHiLUS NOCTULA becomes Nyctalus NOCTULA. 

N. LEISLERl. 

PiPISTRELLUS PIPISTRELLUS. 



S. Leisleri 

s. pipistrellus 

s. serotinus 

Vespertilio Bechsteinii 

V. Nattereri 

V. Daubentonii 

V. mvstacinus 



Vespertilio serotinus. 
Myotis bechsteini. 
M. nattereri. 

M. DAUBENTONI. 
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, 
ScotophUus was based by Leach {Trans. Linnean Soc, London, 
xiii. 71-72, 1822) on 5". ktthlii, a species quite different from 
any of those mentioned above. It was, therefore, employed in 
error by Bell. Vesperugo of Keyserling and Blasius, a substi- 
tute for ScotophUus, has been shown by Mr G. S. Miller to be 
inadmissible in any sense in zoology (see under genus Nyctalus). 

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 crenus of two such dissimilar animals as serotinus and 
nodula. If the separation of noctula and leisleri from true 
pipistrellus be refused, the generic name Nyctalus must, by the 
laws of priority, be applied to all three ; but this question will 
be treated below. The transference of the name Vespertilio 
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 i^Ann. 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 Linnseus's genus Vespertilio (Systema 
NaturcB, I., x., 31-32, 1758), included seven species — vampyrus, 
spectrum^ perspicillatus, spasma, leporinus, auritus, and murinus, 
only two of which, auritus and niuri^tus, 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 Plecotus by Geoffroy in 18 18 
{^Description des Mainmiferes . . . en Egypte, 112). Thus murinus 
is left as the type of the genus Vespertilio. 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 
Stcecica, in the first of which Linnseus mentions only one bat, 
the " Laderlapp," " Fladermus," or " Nattblacka." This he calls 
*' Vespertilio caudatus, naso oreque simplici''' (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 auritus naso oreque simplici, auriculis duplicatis 
capite majoribus' (pp. 1-2, 1761). These had already received 
binomial names, Vespertilio murinus and V. auritus respec- 
tively, in the tenth edition of the Systema Nature, where the 
following diagnosis of V. murinus is given : ''V. caudatus, naso 
oreque simplici, auriculis capite minoribus'' (p. 32, 1758). In 
the second edition of the Fauna Suecica the teeth of V. 



VESPERTILIONID^ 51 

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

It thus appears that the Vespertilio murinus of Linnaeus, 
the type of the genus Vespertilio, is a common Skandinavian 
bat with ears shorter than the head, and with the dental 

formula — 

.2-2^ i-i , i-i ^-3 

I — , c , pm m - — - = -12. 

3-3 I-I 2-2 7^-2, 

The only known Skandinavian bats which combine these 
characters are the members of the group to which V. serotinus 
belongs, and which are commonly known as Vesperus in Europe 
and Adelonycteris in America, but to which Mr Oldfield 
Thomas applied Rafinesque's name, Eptesicus {Proc. Zool. Soc, 
London, 1896, 791, ist April 1897). 

The identification of the species murinus amongst the 
Skandinavian members of the genus Vespertilio, although a 
matter of considerable difficulty, does not affect the use of the 
generic name. Nilsson {Skandinavisk Fauna, Ddggdjuren, 17- 
20, Ed. I., 1847) decided that it must have been the bat to which 
Natterer later gave the name discolor. He therefore placed the 
latter in the synonymy of V. murinus of Linnaeus, and reinstated 
Bechstein's name myosotis (correcting it to myotis), for the Vesper- 
tilio murinus of Schreber. Nilsson did not recognise Vesperugo 
as distinct from VespeiHilio. Hence he said nothing in regard 
to the tenability of the generic names. Ten years later, Blasius 
(Fatcna Deutschlands, S'dttgethiere, 74, 1857), though admit- 
ting that the Vespertilio murinus of Linnaeus 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 Vespertilio of Linnaeus 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 

* 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. 
^ In Linnaeus's statement the figures 4 and 6 are evidently transposed. 



52 VESPERTILIONID^— NYCTALUS 

made to include any of the Linnsean species. Lilljeborg {Sveriges 
och I\Fo7'ges Ryggradsdjur, i., 124-126, 144, 1874) followed 
Blasius, insisting that it is impossible to determine whether 
Linnaus's bat is the species afterwards called Vespertilio 
discolor by Natterer, or that called Vespertilio nilssoni 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 Vespertilio to a group 
containing no species known to Linnseus, 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 Linnseus's 
Vespertilio murimis 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, Exxursions in Madeira^ etc., 36 ; based on N. 

verrucosus of Bowdich, antedating Pterygistes madeir(Z of Barrett- Hamilton, An7i. 

and Mag. Nat. Hist., Jan. 1906, 99. 
1829. Pterygistes, Jakob Kaup, System der Europdischen Thierwelt, i., 99, 100 ; 

based on Vespertilio '•'• proterus et leisleri." 
1839. Vesperugo, a. Graf von Keyserling and J. H. Blasius, Wiegmann's Archiv 

fur Naturgeschichte, i., 312 (part) ; based on Vespertilio serotinus of Schreber and 

eleven other species (see below). 
1842. NOCTULINIA, J. E. Gray, Ann. and Mag. Nat. Hist., Dec, 258 ; based on "A^. 

proterus and N.fulvus." 
1856. Panugo, F. a. Kolenati, Allgeineine deutsche Naturhist. Zeitung (Dresden), 

Neue Folge ii., 131 and 172 ; based on leisleri of Kuhl, and noctula, 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 Vesperugo of Keyserling and Blasius, as originally 
defined, is inadmissible. It included twelve species : serotinus, 
discolor, nilssoni, savii, leucippe, aristippe, noctula, leisleri, 
kuhlii, albolimbatus, nathusiiy and pipistrellus. These were 




Dormouse, (s natural size.) 



NYCTALUS 53 

arranged in two subgenera — Vesperugo including the thirty-four- 
toothed species, and Vespe^'-us those with thirty-two teeth. The 
subgenus Vesperus is exactly equivalent to the restricted genus 
Vespertilio of Linnaeus, to the genera Eptesicus of Rafinesque 
and CnephcBus 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 nodula and leisleri, 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 noctttla-leisleri 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 {Irish 
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 Noctulinia was 
merely characterised as " with the feet quite free, the wing 
being only attached to the ankle ; they are otherwise like 
Scotophihts,'' and consequently contained more species than 
Nyctalus, as used here ; it was adopted by Blyth and Jerdon 
(see under synonymy of N. nocttcla). 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 Pterygistes and Bowdich's Nyctalus (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 {Zoologist, 1898, 100), then 
reluctantly accepted {Ann. and Mag. Nat. Hist., July 1901, 
34), and now seems likely to meet with universal approval. 

The genus Nyctalus includes two British species {N. noc- 
tula and N. leisleri). These are remarkable for their powerful, 
swift-like flight, and the fact that, unlike the Pipistrelle, which 

G 2 



54 VESPERTILIONID.E— 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 N. noctula — viz., in Japan, N. lasiopterus (Schreber), with the 
forearm measuring 60 or more mm. ; central and south-eastern 
Europe,^ N. maximus (Fatio), a mysterious giant of undetermined 
status (see Mr Miller, Proc. Biol. Soc, Washington, 13th June 
1900, 156), with the forearm of 68 mm. and remarkable skull ; 
Sumatra, N. sumatrana (Cuvier^) ; China, N. plancyi (Gerbe) ; 
the Himalayas, N. labiata (Hodgson) ; and Mozambique, 
N. macuanus (Peters) ; the last four very little known. Allied 
to A^. leisleri are N. stenopterus (Dobson) of Borneo and 
Malaysia, a very distinct dusky bat with the forearm of about 
2i"j mm. ; my A^. montamcs, of the Himalayas, and N. ve^^rucosus 
(Bowdich), antedating my N. madeircEy of Madeira, of size 
similar to leisleri, but with distinct crania ; and N. 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 "^'ingQ, Jordf undue og 
nulevende Flagermaus (Ckiropterd) fra Lagoa Santa, Minas 
Geraes, Brasilien, in E. Museo Lttndii, ii., i, Kopenhaven, 1893) 
are Pipistrelliis and Barbastellus, described below : Chalinobus, 
of the Australian and Ethiopian regions, with fleshy lobule to 
the lower lip ; Scotophilus, of the Ethiopian, Oriental, and 
Australian regions, with only thirty teeth ; Harpy iocephalus, 
of the Oriental region, with tubular nostrils ; the long-eared 
Otonypteris, from N.E. Africa and the Himalayas ; Nyctophilus, 
with thirty teeth and small nose-leaf, from Australia ; Lasiurtcs, 
of North and South America, with four mammse and hairy 
interfemoral membrane ; and Antrozous, 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, Bull. 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 
the ear (Plate IV., Fig. i, 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. i 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. i, p. S6 : Plate IX., Fig. i, p. 126). 

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



56 



VESPERTILIONID^— NYCTALUS 






Fig. 4. — Side View (diagrammatic and enlarged) OF Skulls AND TeeTH OF 

I. Nye talus noctula. 2. Vesper tilio serotinus. 

3. Rhinolophus ferrum-equinum. 



NYCTALUS 



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 bullae moderately large (Fig. 4, No. i, p. 56). 

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



3-3 



I - I 



pm 



2-2 



3-3* 



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





Fig. 5. — Diagram of Arrangement of Teeth in Genus Nyctalus. 
(l) 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 Chiroptera, 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 
Rhinolophidse are evidently the most highly organised of 
insectivorous Bats." 



58 VESPERTILIONID.E— NYCTALUS 

THE NOCTULE, OR GREAT BAT 

NYCTA L US NOCTUL A (Schreber). 

1760. La Noctule, L. J. M. Daubenton in E. L. le Clerc, Comte de Buffon's Histcire 
Naiurelle, viii., 128-129, I35-I37j P'- xviii., fig. i ; also Man. de PAcad. Roy. des 
Set., 380, pi. ii. (r5), fig. i, 1759, published 1765 ; described from France. 

1775. Vespertilio noctula, J. C. D. von Schreber, Die Sdugthiere, i., pi. lii., 
166-167, evidently naming Daubenton's La Noctule ; Bingley ; Pennant ; Donovan ; 
Fleming ; Jenyns ; Bell (ed. i) ; Clermont ; Newman. 

1776. Vespertilio lardarius, P. L. S. Miiller, Natursystems Suppletnents und 
Register Band, 15, naming Schreber's Speckmaus, i.e. V. NOCTULA, as above. 

1776. Great Bat, Thomas Pennant, British Zoology (ed. 4), i., 128, pi. xiii., No. 38. 
1789. Vespertilio altivolans, Gilbert White, Selborne, letter xxxvi. to Thomas 
Pennant, September 1771 ; described from Selborne, England. 

1789. Vespertilio magnus, John Berkenhout, Synopsis of the Natural History of 
Great Britain and Ireland, ii., naming Pennant's Great Bat. 

1806. Vespertilio serotinus, Isidore Geofiroy, Ann. du Mus. d'Hist. Nat., viii., 
194, pi. 46, fig. I ; transference of name in error. 

[1812. Vespertilio auriculatus, John Walker, Essays o?i Natural History, 
472 ; described from Edinburgh, Scotland, but only doubtfully referable to 
N. noctula.'] 

18 16. Vespertilio major, W. E. Leach, Systematic Catalogue of the Specimens of the 
Indigenous Mammalia and Birds that are Preserved ift the British Museum, etc. 
(London), 5 ; renaming V. noctula, but without description. 

1 819. Vespertilio proterus, Heinrich Kuhl, Ann. der Wetterauische Gesellschaft 
fiir die gesamjnte 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, Ornis (Jena), iii., 26 ; described 
from Jena, Germany. 

1829. Pterygistes proterus, Jakob Kaup, System der Europdischen Thierwelt, 

i., TOO. 

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 filr Naturgeschichte, i., 317 ; Blasius ; Fatio ; Dobson ; Harting, Zoologist, 
1887, pi. iii., 167-171 ; Blanford ; Woodward and Sherborn ; Lydekker ; Millais, 
pi. 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, Mammals 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, van minima, Victor Fatio, Vertebres 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, North 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 ; Mebely ; Cabrera ; Millais, 61. 

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

La Nodule of the French ; die fruhfliegende Fledermaus of the 
Germans. 

NoctJile, from the French noctule, diminutive from the Latin nox, 
genitive nodis, i.e., " 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 
counties. Ree rot, i.e., rere or rear rat (see " rere mouse" under 
PiPlSTRELLE) of Gloucester (Newstead). Ystlum (pronounced " slim ") 
Fawr, i.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, (i), 185 1, 37 ; Rodd, Zoologist, 1891, 347), but rare in Wight 
{MoxQ, Journ. cit., 1894, 148 ; Wadham), N. nodnla 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 



6o VESPERTILIONID^— 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, Zoologist, 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, Harting, on the authority of Lee, records 
its occurrence at Kendal in Westmorland {^Zoologist, 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 {Statistical Account of the Parishes of Applegarth and 
Sibbaldbie, 1835, 175), and Fleming identified with it John Walker's 
Vespertilio auriculatus} Of late years. Service {Ann. 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 Nyctalus. 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 {Ann. Scott. Nat. Hist., 19 10, 52-53). 

So far as is known, A^. noctula 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. leisleri" {Catalogue, 215), but in 1889, writing to Barrington of 
some further Irish specimens, he thought that, if certain characters 

^ Unfortunately Walker's description of V. auriculatus, except that it clearly does 
not apply to a Rhinolophus, 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, serotinus, which is 
not known to occur in Scotland, than to N. noctula. Alston evidently considered the 
description inexact, or he would not have connected it with Myotis 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 duplicatae, capite multo minores. Exterior major ovata obtusa. Interior 
minor brevior lanceolata. Palmae palmato-alatae maximas, pollice unguiculato. Plantas 
pentadactylae fissae, digitis pilosis unguiculatis. Cauda geniculata, 6 articulis. 
Membrana juxta caudam, margine ciliato. . . ." 



THE NOCTULE, OR GREAT BAT 6i 

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 " {sic). The matter had not been settled at the 
time of Dobson's death, but de Winton compared these Irish with English 
individuals of N. leisleri, and found no difference between them (Alcock, 
Irish Naturalist, 1899, i/O-i). 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 Ban well, 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 



VESPERTILIONID^— 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 N. leislcri, the coat is pale and faded in early summer, 
and deepest and richest 
in early autumn just 
before hibernation. Old 
males are said to possess 
the brightest golden fur. 

The skull is typical of 
the genus. It is character- 
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, but 7nuch broader in 
transverse section at the 
base : its crown is hol- 
lowed out for the recep- 
tion of the tip of the 
lower canine. The lower 
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 





Fig. 6. — Front View of Incisors and Canines of 
Nyctalus noctula (enlarged and diagrammatic.) 



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 viaximus. 

DIMENSIONS IN MILLIMETRES:— 



Males. 




>. 


C 


.a 

■a 




M 


1 

Si 






'So 


iq 


> 




03 * 




















■a 


cj 








c 

cS 
1 

w 




c4 

£ 

Si 

2 

H 


s 





■a 
S 





t3 

a 


1 

a 



t4 


s 


21 "1 
® a 




No. of Items 
Maximum . 


7 


5 


6 


7 






7 




5 


5 


5 


3 


SO 


18 


7 


63 


- 17-18 




^1 'I 

- 50 - 

. 49 j 










r 375 


Average 


77-5 


17 


h 


49-5 


> 


7-S 


87-96 


46-5-50 


36-5-39 -5 


1 


Minimum . 


76 


16 


4 


43 













l 353 


Females. 


No. of Items 
Maximum . 


8 


6 


4 


8 






9 




7 


7 


7 


3 


82 


18 




56 


1 ■ 




r ^* 1 










[ 387 
i 362 


Average 


77 






49 


'^ 17-18 


11-12 


..3 

I 50 j 


7-8 


91.94 


48-52 


38-42 


Minimum . 


73 


16 




41 













Norgate (Zoologist, 1906, 26) gives the average of many as 14J inches, and the extremes as varying 

from 14i 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 m'^ 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 3012 



64 VESPERTILIONID^— 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 A^. leisleri, 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 Natti7'-al History 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 Vespertilio altivolans. 

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 be 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 

' British Zoology^ London ed. of 1812, i, 179. ^ Zoologist^ I905> 427-4'29. 

^ Fields 7th March 1874, 218. * Stanley Lewis, Zoologist, 1906, 69. 

H 



66 VESPERTILIONID^— 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 the wing 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 

* Zoologist, 1874, 4153. ^ Thomas Ford, Field, loth September 1898, 470. 
^ 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 manoeuvres 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 lono" without noticinor 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. 2 ibid.^ 1901, 53. ^ Ibid., 1901, 70. 



68 VESPERTILIONID.E— 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 

* See, for instance, Oxley Grabham, Zoologist^ 1899, 131. 

2 Trans. Caradoc and Severn Valley Field Club, 1900 (Feb. 1901), 243. 

3 In MSS. * Zoologist, 1845, 953. 



PLATE in. 




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 flicrht. 

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 

' Zoologist^ 1889, 258. 

^ Letters xxii., xxvi., and xxxvi., to Thomas Pennant, dated 2nd January 1769, 
8th December 1769, and September 1771. 
2 Fields 23rd April 1892, 585. 

H 2 



70 VESPERTILIONID^— 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 i8th, 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 ist 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 

* Zoologist, 1904, 455- ^ Ibid., 1885, 344. 

^ E.g., Forrest, Steele Elliott, Henry Laver (specimen obtained on the loth), 
Millais. 

* Zoologist, 1891, 167. ^ Ibid., 1890, 217 ; 1891, 305. " Ibid., 1905, 427-429, 



THE 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, 

^ Zoologist^ iQOij 51-59- 



f 

72 VESPERTILIONID^— 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 



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." 

' Zoologist^ 1901, 153. - Irish Naturalist, 1905, 105-106. ^ Zoologist, 1905, 307. 



74 VESPERTILIONID^— 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 

* 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 75 

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.^ 

^ Zoologist^ 1887, 170. 2 Supra, p. 32. ^ Naturalist, 1905, 325-330. 

Mn MSS. ^ Supra, p. 33. 



je VESPERTILIONID.E— 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.^ 

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

2 F. R. Rodd, Zoologist, 1891, 347 ; Oldham, etc. 
^ Proc. Zool. Soc. (London), 1834, 129-132. 



PLATE IV. 






Heads (natural size) of 

I. Nyctalus nocUila. 2. Nyclalus letsleri. 

3. Vesper tilio serotinus. 



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 
raised 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 20th 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 i6th 
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 

^ Naturalist^ I905» 325-330. 



78 VESPERTILIONID^— 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 placenta. 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 30th 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 

^ 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 to a 
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 



8o VESPERTILIONID^— 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 Mr H. J. Charbonnier found 
within the body of a female killed in June a young one having 
an expanse of wing of 3J 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 i6th 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 i% ; 
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. 2)l\ 

* I.e.^ the true period from ovulation followed by fertilisation (not from copulation) 
to parturition. 

2 Naturalist^ March 1899, 7i- ^ Field^ 7th March 1874, 218. * Ibid. 

^ 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 8i 

The Great Bat has frequently been kept ahve 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 I 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 

' Zoologist^ 1 90 1 J 5I-59- 

I 



82 VESPERTILIONID^— 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 : — Mamestra persicarice, Leucania palle?ts, Hepialis humuli, 
H. sylvinus, H. hectus^ Ruinia cratcegata^ Urapteryx sambucata^ Odontopera bidentatn, 
Fidonia atomaria, F. piniaria, Xylophasia polyodon^ and Aviphidasis betularia; the 
list may be completed by the Poplar Hawk Moth, Smerinthus populi^ with one of 
which species Whitaker fed his captive Noctule {Naturalist^ I905) 325-330). 

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

^ As Euchelia jacob<£ce or Abraxas grossulariata. 

* 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 
slio-ht 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). 

18 10. Die RAUHFLUGLiCHE Fledermaus, T. P. Leisler, Magazinfiir die Neiiesten 

Entdeckungen in der Gesammten Naturkunde (Berlin), 156; described from 

Hanau, Germany. 
1819. Vespertilio LEISLERI, Heinrich Kuhl, Ann. der Wetterauische Gesellschaft 
fiir die gesammte Naturkunde., iv., 46, naming Leisler's Rauhfliigliche fledermaus ; 

Jenyns ; Bell (ed. i) ; Clermont ; Newman. 
1819. Vespertilio dasykarpos leisl. («V), Heinrich Kuhl, op. «/., 49, quoting 

Leisler's unpublished MSS. 
1829. Pterygistes LEISLERI, Jakob Kaup, System der Europdischen Thietwelt., i., 

100 ; C. B. Mofifat (doubtfully), Irish Naturalist, 1905, 104 ; Miller ; Thomas ; 

Johnston ; Mehely ; Cabrera ; Millais, 8 and 76. 

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

[?i839. Vespertilio pachygnathus michahelles {sic\ J. A. Wagner's ed. of 
J. C. D. von Schreber's Die Sdugthiere, Supplement i., pi. Iv. ^. There is no 
description, but Fitzinger states that Michahelles found this bat in Dalmatia : 
it appears to be a small Nyctalus^ 

1839. Vesperugo LEISLERI, A. Graf von Keyserling and J. H. Blasius, Wiegmann's 
Archiv fiir Naturgeschichte, \., 318; Blasius; Fatio ; Dobson ; Blanford ; Alcock ; 
Mofifat. 

1856. Panugo LEISLERI, F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitung 

(Dresden), Neue Folge, ii., 131, 172. 
1870. Noctulinia LEISLERI, L. J. Fitzinger, Sitzungsberichte der Kaiserlichen Akad. 

der Wissenschaften (Vienna), Ixii. (i), 218 ; Harrison Allen, Proc. U.S. National 

Museum, xvi., 30, footnote, 1893. 
1898. PiPlSTRELLUS LEISLERI, Oldfield Thomas, Zoologist, 100. 

Le Vesperien de Leisler of the French (Fatio), die rauharviige 
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 VESPERTILIONID.E— NYCTALUS 

(Winge), and probably the shores of the Mediterranean generally, and 
from Ireland probably to China. As in the case of N. noctula, 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 {Field, 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 GrSihhd.va, Naturalist, 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 {Zoologist, 
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 doubtful 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. leisleri" 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 N. noctula that 
I have no hesitation in connecting his specimens with that species. 
Possibly the band of hair on the arm, which occurs in N. noctula as well 
as in A^. leisleri, 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. 




O XI 

*: 6 

e P 

B ■" 

1) O 

O o 

<u >— ■ 




■^ (U 



^ 



PLATE VII. 






Wings (^ natural size) OF 



T. Nyctalus leisleri. 2. Pipistrellus pipistrellus. 

3. My Otis daubento7ii. 



LEISLER'S BAT 85 

should not occur in other parts of England, Wales, and even in Man 
and Scotland (see Service's observations under A^. tioctula). 

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 BlackstaffLane, 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 Natural History 
and Philosophical Society on 25th April 1860.^ 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. cit., 1874, 4236), 
Wicklow (Barrington, Journ. cit., 1875, 4532), Fermanagh (Barrington, 
Journ. cit., 1883, 116), Westmeath (Moffat, Irish Naturalist, 1897, 135), 
Cavan, Louth (Jameson, /w^rw. cit., 1897, 41-42), Londonderry (Alcock, 
Jourji. cit., 1899, 174), Carlow and Wexford (Barrett-Hamilton, /(C?z^r«. 
cit., 1900, 134; Moffat, 162; Pack Beresford, 1906, 194). Its range, 
although a few county records, e.g., 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. 

Tlie number of young, although stated to be usually two in Germany, 
is not known to exceed one in Britain (see Tomes, Zoologist, 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 VII., 
Fig. i), 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 



VESPERTILIONID^— 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 noctula-Yike 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 N. 7ioctula. 

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 four-teen examples (seven of either sex) of 
the allied N. azorejiiii 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 N. noctula, its struc- 
ture is weaker and more papery, 
and the crests, especially the 
sagittal, are absent or feebly de- 
veloped (Fig. 16, No. I, p. 174). 

The teeth are similar, but 
much smaller. The outer upper 

incisor is comparatively small, its cross-section at the base being about 
equal to, not about double, that of the inner incisor. The anterior upper 




Fig. 7. — Front View of Incisors and 
Canines of Nyctalns leisleri (enlarged and 
diagrammatic). 



LEISLER'S BAT 



87 



premolar is relatively larger, and the lower incisors, being smaller, are less 
crowded and do not usually overlap (Figs. 7, p. 86, and 16, No. i, 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 N. noctula^ 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 


:— 










>> 

■a 

.a 


.a 

■a 
g 






fci 




3 


i 


'3) 


73 


> 
1 




a 

a . 
am 




■a 


s 


^i 


'5 


0? 


^l 


c9 




2 


& 


c4 


X to 

V a 




1 




Ml 


H 





T3 

a 
t5 




.a 
B 

a 


§ 


il 


1 


1 
g 




Leach's specimen in 


























British Museum (No. 


























d. 63a) .... 














43 












Male, Mexboro', Yorks. 


























(H.J. Charbonnier, No. 


























90. 7. 6. 1. of British 


























Museum collection) 














43 












Male (in alcohol), Bray, 


























Ireland (R. M. Barring- 


























ton per J. E. Halting, 


























No. 90. 2. 14.1. of British 


























Museum collection) 


56 








19 


9 


42 






38 


32-5 




Male, Co. Clare, Ireland 


























(Dublin Museum) . 










16-5 


7 


42 






40-2 


32 




Male, Co. Dublin, Ireland 


























(Dublin Museum) . 










17 


S 


42-5 






40 


31 




Male, Co. Louth, Ireland 


























(Dublin Museum) . 










16 


8 


40 






37 


30-5 




Female (in alcohol), Ar- 


























magh, Ireland (R. M. 


























Barrington) . 


57-5 


14-5 


6 




18 


7-5 


42 


6 


71 


36 


30 




Female, Co. Armagh, Ire- 


























land (R. M. Barring- 


























ton per G. B. Dobson, 
























No. 89. 11. 12. 5. of 


























British Museum collec- 


























tion ; labelled " var. 


























hibernictis " in Dobson's 


























handwriting . 


61 


14 


4 




19 


9 


44 




74 


39 


S3 




Female, Belfast, Ireland 


























(R. Patterson per G. B. 


























H. Barrett - Hamilton, 


























No. 01. 3. 15. 1. of 


























British Museum collec- 


























tion) .... 


62-5 


17 


4 


41 






43 










*295 


Female, Co. Dublin, Ire- 
























(approx.) 


land (Dublin Museum) . 










18 


8 


44-5 






39 


33 




Female, Co. Dublin, Ire- 


























land (Dublin Museum) . 










17 


8 


43 






39-5 


31 




Male, immature, Co. 


























Cavan, Ireland (Dublin 


























Museum, in alcohol) 
Male, immature, Co. 










15-5 


8 


39 




62 


35 


28 




















(approx.) 








Dublin, Ireland (Dublin 


























Museum, in alcohol) 

Average of adults \ 
(approximate) . / 










15 
17 


7-5 


38-6 






34 


29 




59-5 


15 


4-.'i 




8 


42 




72-5 


38-5 


31-6 





llf inches in the flesh, ^e B. Patterson, corroborated independently by Whitaker, who gives the 
average wing-expanse of six Yorkshire females as 304 (Naturalist, 1907, 386). 



VESPERTILIONID^— 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), 1 890-1 891 : — 





E 


.Q 


Digit 
II. 


Digit III. 


Digit IV 




Digit V. 


tlO 
























rt 


s 


























s 




d 


c3 






C3 






05 













H 





1 


.5 









-3^ 


C3 


X 
a 
rt 


MC15 


& 

3 








t4 


cS 


"rt 


ce <A 


ce 


75 


% =s 


cS 


"^ 


rt t3 










<o 


"S 


Si 


;=i<M 


"® 


.c 


fi- 


"S 


A 


;?=<N 










% 


155 


Ph 


P-i 


s 


Ph 


S 


^ 


&H 




r 


41 


6-5 


38 


40 


16 


19 


38 


14 


7 


32 


8 


7 


16 




41 


6 


38 


40 


16 


19 


39 


15-5 


7 


33 


8 


6-5 


16 


Males . . .-; 






























41 


6 


39 


40 


16 


IS 


39 


16 


6-5 


33 


8 


7 


16 


V 


41-5 


5 


39 


40-5 


17 


20 


39 


16 


7 


33 


8 


7 


15 


j 


40 


6 


39 


37 


15-5 


18 


37 


14-5 


7 


32 


8 


6-5 


15 


Females 


41 


5-5 


38-5 


39 


16-5 


20 


39 


15 


7-5 


33 


8 


7-5 


17 


Average of 7 


41 


6 


39 


40 


15-5 


20-5 


39 


15-6 


7 


33 


8 


7 


16 


40-9 


5-7 


38-6 


39-5 


16 


19-2 


38-5 


15-2 


7 


32-7 


8 


6-9 


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, 1 1-25 to 12 ; 
palatal length in middle line, 5-5 to 6; from posterior border of vi^ 
to anterior border of canine, 6-5 to 6 ; ditto in lower jaw, 6 to 6-5 ; 
greatest breadth at zygoma, 10 to 10-5; posterior breadth, 9; breadth 
between orbits, 8 ; breadth at constriction, 5 (nearly). 

Weight: — 14 to 20 grammes (Moffat, IrisJi 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 N. noctula. 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 pdour, 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 VESPERTILIONID^— 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. Moffat/ The latter was 
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 

^ Zoologist, 1874, 4071-4074. - Irish Naturalist, 1897, 41- 

3 Ibid., 1899, 169-174, and map. "» Ibid., 1900, 235-240 ; 1905, 99-101. 

^ 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 {Naturalist, 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. 

^ Zoologist, 1883, 116. " Irish Naturalist, 1900, 134. 



LEISLER'S BAT -91 

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 in a 
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 m.ade 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 57^^°, 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 21st 
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 i6th 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 

^ Moffat, Irish Naiuralisf, 1897, 135. 



92 VESPERTILIONID^— NYCTALUS 

time of flight ; for I have never seen it when the day's viaxiinum 
was below 52|-°, and never nuiiieroiisly 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 loth 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. 1 1 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 

^ Irish Naturalist^ 1899, 169-174, and map. 



PLATE I. 



m.e,. 




jo^.a.e. 



Portions of Finger Skeletons of Bats (diagrammatic), enlarged 
TO show Stages in Ossification of the Joints. 

m. Metacarpal;///!, phalanx i;ph2, phalanx 2; ine, metacarpal epiphysis; p/i I e. first 
phalangeal epiphysis ; p/i 2 />, second phalangeal epiphysis. 

A. Young specimen with epiphyses quite distinct from phalanges and metacarpal. 

B. Older but still immature specimen, in which ossihcation, 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 T,2)y 3^> -4» ^^^ ^3 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 1 2th I again saw five come out: the first one 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. 

" I therefore aro^ued that since I could not detect the bats 
going m in the evening, I must endeavour to catch them as 
they came o?i^ in the morning. This would at least prove the 
fact, though not the time, of their retirement after dusk. So, 
on the nig^ht of Ausfust 12th, I fixed a net at midnig-ht 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 . . . 

^ " In explanation of these figures, I should state that at Ballyhyland (lat. 52° 31' N., 
long. 6° 43' W.) sunrise on August loth is 4 minutes later than at Dublin, and 
sunset I minute earlier." 

K 2 



94 VESPERTILIONID^— 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 longer period than "j^, or a shortei'- 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 I3|- minutes, the 
earliest i minute, the latest 23 minutes, and the mean between 
the extremes 1 2 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 Jirst 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 2>3 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. loth three (instead of four) emerged from the common 
den; on the iith, 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 



g6 VESPERTILIONID/E— 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 MoflTat 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 

* Irt'sA Naturalist, i899) 169-174, and map. 




^ 




vo 





to 




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-^ 



t-5 



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<u 


w 


B 




g 


5 


3 


X 


CD 




1 


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D 


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C/3 






f- 


— 


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LEISLER'S BAT 97 

the shrill screech, and from that time one hears the tinklinsf 
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 
I ith 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 1 6th 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 

^ Naturalist, 1907, 387, 415, etc. - Scatophaga stercoraria. 

^ Probably a mycetophilid, also an acalypterate muscid. 



98 VESPERTILIONID/E— 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 living 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. - Tipula oleracea. 

2 Hcematopota pluviaUs. * Bombus terresMs. ^ B. lapidarius. 

^ Mofifat, Irish Naturalist^ 1897, 135 ; Jameson, Irish 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 Harrington, 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 (ew 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 

' Zoologist, 1854, 4365. 2 JSTaturalist, 1907, 385-6. 

^ N. azoreum. ^ Zoologist, 1882, 16. 



100 VESPERTILIONID^— 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 Europdischen 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. kuhlii (Natterer). 

1839. Vesperugo, a. Graf von Keyserling and J. H. Blasius, Wiegmann's Archiv 
fiir Naturgeschichte, i., 312 (part) ; based on Vespertilio serotinus of Schreber and 

eleven other species (see under Nyctalus). 

1840. ROMICIUS, Edward Blyth, C\iv'\&x^s Animal Kingdom, 75 ; misprint for ROMICIA. 
1856. Hypsugo, F. a. Kolenati, Allgemeine deiitsche Naiurhist. Zeitung (Dresden), 

Neue Folge, ii., 131, 167-169 ; based on Vesperugo maurus of Blasius, and V. 

krascheninikowii of Eversmann. 
1856. Nannugo, F. a. Kolenati, op. cit, 131, 169-171 ; based on Vesperugo nathusii 

of Keyserling and Blasius, Vespertilio pipistrellus of Daubenton, and V. kuhlii 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 Nyctahis applies also to Pipistrellus, 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. i), as exemplified by that of 
P. ptpistrelhis, 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 



PLATE Vr. 




Leisler's Bats. (Approximately natural size.) 



PIPISTRELLUS 



lOI 



perceptible, and the auditory bullae are more prominent than 
in Myotis, less so than in Barbastella. 









Fig. 8. — Side View (diagrammatic and enlarged) OF Skulls and Teeth of 



1 . Pipistrellus pipistrellus. 

2. Myoiis d(Xubentoni. 

3. M. mystacmus. 



4. Plecotus aurilus. 

5. Barbastella barbasielbis. 

6. Rhiiwlopkus hipposideros. 



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



I02 VESPERTILIONID/E— 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). 





I 2 

Pig. (j. — Diagram of Arrangement of Teeth of Piphirellus pipntrellus. 
(l) 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, pipistrellus, 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 Naturaliste, 1879, 16, 125), 
and Sweden. But its reputed occurrences in Europe should 
no doubt be credited to P. nathusii (Keyserling and Blasius), 
with light-coloured posterior wing-border and long lower 
canine, described as of central Europe. Of other species, 
P. kuhlii (Natterer), a southern form, also taken at Gironde, 
and with a sub-species reaching Cape Colony (/*. kiihlii fuscaius 
of Thomas), has the posterior border of the wings white and 
the outer upper incisor minute ; P. savii (Bonaparte), a form of 
wide Pala;arctic range, with a representative — P. albolimbatus 
(Kuster) — in Sardinia, and shown by Forsyth '^\2!]ox {^Atti della 



COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 103 

Sor. Toscana diSci. Nat., 3, fasc. i, Z^j, Pisa, 1877) to be synony- 
mous with luaurus of Blasius and bonapaiii 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. subflavits (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, Pmax, 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 

Histoire Naturelle, viii., 129-130, 135, 137, pi. xix., fig. i ; also, Mem. de I'Acad. 

Roy. des ScL, 381, pi. i. (14), fig. 3, 1759, published 1765 ; described from France. 
1766. Short-eared Bat, Thomas Pennant, British Zoology, ed. i, 55. 
1775. Vespertilio pipistrellus, J. C. D. von Schreber, Die Siiugthiere, i., pi. liv., 

167, evidently naming Daubenton's La Pipistrelle ; Jenyns, Trans. Linnean Soc. 

(London), xvi., 163, read 3rd Feb. 1829, published 1833 ; Bell (ed. i) ; Clermont ; 

Flower and Lydekker. 
1789. Vespertilio murinus, Gilbert White, Selbome, letter xi. to Thomas Pennant, 

9th Sept. 1767, original edition, 32 ; Berkenhout ; Bingley ; Donovan; Fleming; 

not V. murinus of Linnaeus, 1758 = (probably) V. DISCOLOR of Natterer ; not 

V. MURINUS of most continental writers = MvOTis myosotis (Bechstein). 
1811. Vespertilio pipistrella, P. S. Pallas, Zoographia Rosso-Asiafica, i., 123, 

No. 48. 
1825. Vespertilio pygm/EUS, W. E. Leach, Zoological Journal, i., 560, pi. 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 Eicropiiischen Thierzuelt, 

i., 98; Miller, Ann. and Mag. Nat. Hist., Oct. 1897, 384; Thomas, Zoologist, 1898, 

100 ; Johnston ; Mehely ; Cabrera ; Millais, 79. 
1833. Vespertilio brack yotus, L. A. F. Baillon, Mem. de la Soc. Roy. d' Emula- 
tion d^ Abbeville, 50 ; an unrecognisable species, probably referable io pipistrellus; 

described from a specimen {collo super, tiigro) 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, Europaische Fauna, i., 9 ; 
described from Zurich, Switzerland (type in Zurich Museum). 

1840. "V. MELANOPTERUS et STENOTUS, Brehm," aiict. et op. cit.; described from 

Renthendorf, Thuringia, Germany, and Switzerland. 
1840. "V. PUSILLUS, Brehm," rtz/r/. et op. cit. 



I04 VESPERTILIONID^— PIPISTRELLUS 

1840. Vespertilio lacteus, C. J. Temminck, Mo7iographtes de Mammalogie, ii., 
245 (for date see Wiegmann's Archiv, 1841, 23) ; described from two nearly white, 
immature bats of doubtful origin in the Leiden Museum (see Dobson, Catalogue^ 
225). 

1841. Vespertilio vispistrellus, C. L., Prince Bonaparte, Fauna Italica, i., pi. xi., 
fig. I ; described from Italy. 

1842. Kerivoula GRISEUS, J. E. Gray, Ann. 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 Meridionale, i., 24, is thus placed 
by E. L. Trouessart, Bull, de la Soc. d'' Etude des Sci. Nat. de Nimcs, 7, i, 55, 1879 ; 
and, on the authority of Paul Gervais, identified with P. pipistrelltis though slightly 
smaller ; it was described from Nimes, France. But Gervais, Histoire Naturelle 
des Mammiferes, i., 216, 1854, cites V. nigrans next to pipistrellus, but as a distinct 
species, under the name of V. nigricans Gene : this is evidently not K 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, Sitsungsberichte der Kaiserlichen 
Akad. der Wissenschaften (Vienna), xxviii. (3), 246, and xxix. (10), 334, fig. 7. 

1 862- 1 863. Nannugo pipistrellus, a, van tvpus, p. 490 ; /3, van flavescens, 
p. 491 ; 7, var. NIGRICANS, p. 491 ; 5, var. LIMBATUS, p. 492 ; Carl Koch, 
Jahrbiccher des Vereins f'iir Naturkunde iin Herzogthuvi (Nassau), xviii., 487-500 ; 
described from Nassau, Germany. 

1870. Vesperugo MINUTISSIMUS, L. J. Fitzinger, Sitzungsbcrichte der Kaiserlichen 
Akad. der Wissenschaften (Vienna), Ixii. (i), 252. 

1874. SCOTOPHILUS PIPISTRELLUS, Thomas Bell, History 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 ; die Zivergflcder- 
maus of the Germans. 

Synonymy : — This presents no difficulty. Those who object to 
the Scomber scomber principle will probably find the combination 
Pipistrellus pygnicsus (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 Pai'ish, 1892, 137), 
from back,^ backie, bak, bakke, bauckie, baukie, or bawkie, an old form 
of bat, having variants in Denmark, Iceland (compare ledhrblaka, i.e., 
leather-flapper), Norway (compare Old Norse blaka, i.e., to flap), and 
Sweden (compare nattblacka, i.e., a bat), and still used in Scotland, often 
in combination as backie-bird, etc. (Alston) ; barnmouse, bastat, bathy- 

^ A word usually applied to ships, and probably meaning sailing or floating away 
(Wright). 

- Compare — 

" The laverock and the lark, The baukie and the bat, 
The heather-bleet, the miresnipe. How mony birds be that ?" 

— CnxUBKRS, Popular Rhymes of Scotland {&d.. 1870), 198. 



COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 105 

mouse (Forbes) ; bat-mouse, of Stafford (Adams, MS.) ; batty mouse, 
billy-bat {Plecotus, Wright) ; bit-bat (Forbes), blin'bat, i.e., blind-bat, of 
Kirkcudbright (Service, Zoologist, 1878, 427); blind-bat, of Dublin 
(Moffat, MS.) ; brere-mus (Forbes), (?) = reremouse ; chipper, i.e., 
chirruper, from its cry of "chip, chip" (Forbes); flickermouse (Ben 
Jonson), as in New Inn, iii., i — 

" Once a bat, and ever a bat ! a rare mouse, 
And a bird o' twilight ; . . . 
Come, I will see the flickermouse " ; 

flinder (Forbes), compare Dutch vlindcr, i.e., a butterfly ; flinder-, flinter-, 
i.e., butterfly-mouse (compare in Caxton's Reynard the /77,r (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 fledevmaus), of many counties, 
e.g., Yorkshire (Atkinson, op. cit.) and Essex (Laver), as in Ben Jonson's 
Alchemist, v., 2 — 

" My fine flitter-mouse, 
My bird o' the night ; " 

glaik of Lothian (Forbes), i.e., a transient glance or gleam ; haddabat 
(Miller and SkevichX^y, Fen land, 1878, 358); hatbat (Forbes); leathern 
mouse, leathern wings (Forbes) ; leather-winged bat of Wexford (Moffat, 
MS.); mouse bat, of Buckingham and Berkshire (Cocks, Zoologist, 1878, 
334); oagar-triunse, of Shetland (Forbes), (?) = -hiunse, -hinnse or 
-hunch, i.e., a frightful creature (Wright) ; pipistrelle, mostly a book- 
name, from French pipistrelle, from Italian pipistrello, vipistrello, vespis- 
trello, variants or diminutives of vcspertillo or vespertilio, from Latin 
vespertilio, 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 hreremus, from hre'ran, i.e., to move, shake, stir, and 
vij'ts, i.e., 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 ialtagox ialtog, with variants dialtog,fialtag ox 
taltag, perhaps from iall — leather, sometimes combined with leathair, 
i.e., leather ialtag ; craicneach (?) = craicneog, i.e., skinny, as in Man 
(see Kermode) ; callah or cal-luch, and feascarhich, mioltog leathair, 

L 



io6 VESPERTILIONIDvE— PIPISTRELLUS 

i.e., leather fly ; sciathan leathair, i.e., leather wings ; ystlum, as in Wales ; 
see under NOCTULE (Alston ; Forbes ; O'Connell ; Wilde, Pj'oc. Roy. 
Irish 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. subfiaviis 
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 Myotis 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. auritus 
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 Al. mystacinus, as Jenyns found to be the case to a 
great extent at Bath (see Harting, Zoologist, 1888, 164). In Shropshire 
also its numbers are less than those of P. auritus (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 M. mystacinus (Coward and Oldham), in Yorkshire probably about 
as numerous as that species or P. auritus (Roebuck quoted by Harting, 
loc. cit., 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 M. 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, Irish Naturalist, 1900, 233). 



COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 107 

It has probably found its way to every island of any extent, and 
has been identified from Orkney (Spence, Ann. Scott. Nat. Hist., 1909, 
47), Islay, Mull (Alston; Gilmour, Mag. cit., 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, Mag.cit., 1904, 125 ; also 
Millais), Jura(by myself), the Blaskets (Kane, Irish Naturalist, 1897, 88), 
and other Irish islands, and at such outlying lighthouses as the Fastnet, 
Rockabill, and Blackrock, Co. Mayo (Barrington, Migration 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, Zoologist, 1888, 24). 

Distribution in time : — Pleistocene remains from Ightham Fissure, 
Kent, have been provisionally referred to this species by E. T. Newton 
{Quarterly J ourn. 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. pipistrellus 
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 XL, 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 Nyctalus, 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 



io8 



VESPERTILIONID^— PIPISTRELLUS 




hairs are everywhere dusky, the dusky basal portion reaching a length 
of 7 mm. out of a total of lo 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 Nyctahis, since there seems 
to be a tendency to deeper tints in March and to comparative pallor 

in summer until August. There are cer- 
tainly no marked changes, and such as 
occur areobscured by individual variation. 
Two young ones (Plate II., Fig. 2, 
p. 16), born in the possession of Whitaker 
had at birth a few straggling hairs on the 
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- 
third day, when the animal died. 

The skull and teeth are typical of 
the genus (Figs. 8, No. i, p. loi ; 9, 
p. 102 ; and 10). 

Individual variation is very frequent, and tends in three directions, 
i.e., 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, Field, 
14th Sept. 1889, 408, and J. M., Joiirtt. ciL, 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 




Fig. 10. — Front View of Incisors 
AND Canines of Pipistrellus pipi- 
strellus (enlarged and diagrammatic). 



COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE 109 

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 
Diacropteriis ( Verhandlungcn der kaiserlich-k'dniglicJien aool.-bot. Gesell- 
schaft (Vienna), xii., 1862, 250), described by Zeittelles, from Kaschau, 
Hungary. Cabrera's sub-species inediterranetis, 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, Ann. and Mag. Nat. Hist., March 1909, 
258), and of a Ouetta specimen in the British Museum. 

DIMENSIONS IN MILLIMETRES 1 :— 



Males. 






































■s 






3 




_^ 













>M 




to 
c 


a 






.a 




;3 



^' 


t3 


> 


s 









S. 


to 




bo 




S 


.a 

4J 


'3 


"rt 


"3 


a. 




■a 




s 


*^ 


'.J. 




-& 




i i 


.t;> 


P* 


V 


K tao 




c 
















. C8 




ts 


C3 


® a 




(33 




C5 


bb 


H 





=2" 





T- 




=5 


i! 


1'? 




cS 




to 












3 













W 












S 






h5 


;^ 


S 





No. of Items . 

Maximum 


9 
44-5 






9 


4 


12 


12 










3 


J 






-> 31-85 
I. 29-2 


11-75 


6-75 


30 "1 










r212 

-. 208 
I 206 


Average . 


42 


10-00 


4-00 


10-5 


5-65 


29-5 '- 
27-5 j 


3-6-5 


50 


25 


25 


Minimum 


39 






10 


4 










Females. 


No. of Items . 
Maximum 


11 






11 


5 


5 


U 










6 


52 


/ 






C 33 
-,' 30-5 
i 29-21 


12 


8 


31-25 -^ 








r 218'5 
- 213 


Average . 


43-35 


10-00 


4-00 


11-36 


7-14 


30-28 ^ 
29 J 


8-5-5 


50 


25 


25 


Minimum 


34-5 






10-5 


6-75 










. 203 



1 For these I am partly indebted to Wliitaker. 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 (Zoo/ogist, 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, 

L 2 



I lo VESPERTILIONID^— 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 v^ 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 [loc. cit.) weighed 28, young and old, 
in July, and found them varying from 2-6 to 5-4 (40 to 83 grains) : 
Moffat {Irish 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, loth 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, Barbnstella, and 
RJiinolophus, leaving only Myotis, of which two species, bechsteini and 
nattereri, are distinctly larger ; daubentoni is less so, but has distinct 
coloration and markedly larger foot ; viystacinus 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. nathusii 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 — in fact 
the smallest British mammal — its w^eight 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 mterimts, not of Linnrcus = Myotis myosotis (Bechstein). 



COMMON BAT, PIPISTRELLE OR FLITTER-MOUSE in 

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. 



1 12 VESPERTILIONIDyE— 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. MlUals relates, a wall had to be pulled 
down before a large colony could be ejected, their squeaks 
and scratchlngs 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 Piplstrelle 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 

^ Zoologist^ 19065 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 blov/n 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. 

^ 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, 

3 Zoologist, 1853, 3936-3943. 



1 14 VESPERTILIONID^— PIPISTRELLUS 

More recently, Mr C. B. Moffat ^ has taken the trouble to 
compare the movements of bats, hedgehogs, and frogs from 
26th October 1901 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 43^^° 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 

' Irish Naturalist, 1904, 81-87. 

2 The corresponding figures as given by Jenyns are 40° 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 loth 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), ''Et in cibatu aUices gratissivii,'' 
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 

' Zoologist, 1887, 69. - Ibid., 1S87, 143. » Qp^ ^/^_^ js^.^ ^942. 

'' C. Plinii scciindi Naturalis Historiae, ed. of Joannes Harduinus, lib. x., cap. Ixi. 
(Ixxxi.), 454 (Paris), 1685 ("and for food gnats [are] very pleasant [to it]"). 
^ Zoologist, 1887, 293. 



1 16 VESPERTILIONID^— 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 2ist-22nd, 
1 899) 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, 

^ Catalogue of CMroptera, \.v\\., iooinoie, 1878. 
~ Irish 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 1 6th, 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 



1 18 VESPERTILIONID/E— 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 I 
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 Moft'at 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 

^ Naf. Hist Review (Dublin), 1854, 23-24. 



COMMON BAT, PIPISTRELLE OR FLITTER-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 i6th 
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. 



I20 VESPERTILIONID.^— 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-Inferieure," 
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 '' Miiiimai7i pro corpore vocem emithnity^ 
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 

> Bull. Soc. Amis Sci. Nat. (Rouen), 4th April 1901, reprint, 2. 
^ Field, 29th August 1903, 405. ■' Naturalist, 1907, ']']. 

■* Metamorphoses, lib. iv., 10. ■'' 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 nth 
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, loth, i8th, and 19th ; 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 9th 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 21st 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 in a 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- 

^ See his " Notes on the Breeding Habits of Bats," Naturalist, 1905, 325-330. 

- A statement by J. J. Briggs {Zoologist, 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. 

^ Irish Naturalist, 1896, 95. 

* Now in the Exeter Museum, and kindly submitted to me for examination by 
Edwin Hollis. 

M 



122 VESPERTILIONID^— 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 infantes " ^—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 

^ Loc. cit. supra, p. 115, "[The mother] flies embracing twin young." 
2 Proc. Zool. Soc. (London), nth 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 VESPERTILIONID^— 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 



PLATE II. 




y^c. yDc<3?- mcd. ecdj 



JL. 



B. 



a 



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

A. Crown view of upper molar. B. Crown view of lower molar. 
C. Side view of upper molar. 



ecd^ entoconid 

he, h3'pocone 

hcd^ hypoconid 



;«(-, metacone 

mcd, metaconid 

ins, mesostyle 



mts, metastyle 

pc, paracone 

pcd, paraconid 



pre, protocone 

p)xd, protoconid 

ps, parast3'le 










Fig. 2.— Young Pipistrelle. (Natural size.) 

From a specimen in the British Museum, to shovv the small wings, but 
large feet and thumbs. 




PLATE VIII. 



i 




PlPlSTRELLE 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. "- Letter xi., dated 9th Sept. 1767. 

3 Zoologist, 1847, 1 766- 1 767. 



126 VESPERTILIONID^— 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 in its 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 fiies 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 orround, usincr its tail as a 
basket until it had obtained a firm hold. Mr Cuming writes 
somewhat similarly in his pleasant book, The Arcadian 
Calendar,^ and in reply to my inquiry for further details, 
he adds the information that he has "often watched the 

1 Midland Naturalist, 1883, 149-153, arranged and contributed by H. A. Mac- 
pherson. 

'-^ Animal Life and Intelligence (London : Edward Arnold), 1 890-1 891, 65. 
3 George Newnes, Ltd., London, 1903, 35 (illustrated by J. A. Shepherd). 



VESPERTILIO 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 manoeuvre 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 manoeuvre 
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 Linn£eus, Systema Nattirce^ x., 31-32 ; based on Vesperftlw 
muriftus of Linnsus (not V. miirinus of Schreber, 1775, see above, page 51). 

1820, Eptesicus, C. S. Rafinesque, Amials of Nature^ 2 ; based on Eptesicus 
inelanops of Rafinesque = Vesperiilio fuscus of Beauvois. 

1829. Cneph^US, 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 
fiir Naturgeschkhte, i., 312 ; based on Vespertilio serotinus of Schreber, and eleven 
other species. 

1839. Vesperus (sub-genus), auct. et op. cit., 313; based on the thirty-two toothed 
species of the genus Vesperugo, e.g., serotinus of Schreber, discolor of Natterer, 
nilssoni of Keyserling and Blasius, savii of Bonaparte, leucippe of Bonaparte, and 



128 VESPERTILIONID^— VESPERTILIO 

aristippe of Bonaparte ; preoccupied by Vesperus of Latreille, 1829, a genus of 

Coleoptera. 
1 84 1. NOCTULA (sub-genus of Pipistrellus\ C. L., Prince Bonaparte, Fatma Tfalica, 

i., xxi. (under Vespertilio alcythoe) ; based on Vespertilio serotinus of Schreber. 
1856. Cateorus (sub-genus), F. A. Kolenati, Allgemeine deutsche Naturhist. Zeitiing 

(Dresden), Neue Folge, ii., 131, 162-163 > based on serotinus^ i.e., La Serotine of 

Daubenton. 
1856. Meteorus (sub-genus), F. A. Kolenati, op. cif., ii., 131, 163-167 ; included 

nilssoni of Blasius, discolor of Kuhl, leucippe, aristippe, and savii, all of Bonaparte ; 

preoccupied by METEORUS of Halliday, 1835, a genus oi Hymenoptera. 
1858. AmblyOTUS, F. a. Kolenati, Sitzungsberichte der Kaiserlichen Akad. der 

Wissenschaften (Vienna), xxix. (9), 252 ; based on A. atratiis of Kolenati = Vesperugo 

nilssoni of Keyserling and Blasius. 

1863. "Aristippe, F. A. Kolenati, Beitriige zur Kenntniss der Phtkiriomyiarien, 
Petersburg, 1863" (thus in Koch, Das Wesentliche der Chiropteren, etc., 471, 473, 
1863, under Meteorus) ; included V. discolor of Natterer= V. muri)ius of Linnaeus 
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 

Scotophihis pachyojnus of Tomes. 
1870. Nyctiptenus, L. J. Fitzinger, Sitzungsberichte cit. supra, Ixii. (i), 424; based 

on Vespertilio sviithii of Wagner. 
1872. MARSlPOLiEMUS, W. Peters, Monatsberichte der Koniglich Akad. der Wissen- 

schafteri (Berlin), 260, a sub-genus of Vesperugo based on Vesperus {M.) albigularis 

of Peters = V. murinus of Linnaeus. 
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 
Vespertilio has been fully explained on pages 50-51. 

M. Mehely uses Rafinesque's name Eptesiciis for the serotiniis- 
fuscus group, restricting the name Vespe^'tilio to the remainder of 
the genus as here understood. Mr Thomas has pointed out 
the objections to this proposal i^Ann. and Mag, Nat. Hist., July 
1901, 31-32). A more recent arrangement, that of Mr Miller, 
makes the division between V. murinus as sole representative of 
Vespertilio, and the remaining species as Eptesiats. This will 
probably meet with general acceptance, since the short broad 
tragus and large nares of typical Vespertilio 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 V. serotinus, V. nilssoiti of Keyserling 
and Blasius ( = /^. borealis audoruni) of northern Europe and 
Siberia generally to the Arctic circle ; V. vmrimis of Linnaeus 
( = K discolor midoruin) of the temperate portions of the Palae- 
arctic region, mainly in moun- 
tainous districts (for distinctions, 
see page 140, under V. miirintts) ; 
and Cabrera's V. ochroinixhis 
with forearm only 32 mm., in 
central Spain and the Balearics. 
There are representatives in 
North and South America (see 
under V. serotinus^ 

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- 
tion. The wing (Plate IX., 
Fig. 2, p. 126) is broader than in 
Nyctalus, but springs from the 
base of the toes, as in Pipi- 
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 — 




Fig. I r,— Diagram of Arrangement 

OF Teeth of Vespertilio serotinus. 
(i) Upper and (2) Lower Jaw. 



inc 



pm 



3-3' I-r ' 2-2' 3-3 

The skull (Fig. 4, No. 2, p. 56), as exemplified by serotinus, 
is heavily built, flattened, larger and altogether stronger than that 
of Pipistrellus, but, as compared with that of Nyctalus, 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 bullae smaller. In dorsal profile the skull presents 
nearly a straight line, rising gradually from nose to occiput. 

N 



I30 VESPERTILIONIDyE— VESPERTILIO 

Only one species, V. serotinus, is a regular member of the 
British Fauna, but V. murimis 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 

Histoire Naturelle, viii., 129, pi. xviii., fig. 2 ; also, Man. de VAcad. Roy. des Sd., 

2,77 and 380, pi. ii. (15), fig. 2, 1759, published 1765 ; described from France. 
1774. VESPERTILIO SEROTINUS, J. C. D. von Schreber, Die Sdiigthiere^ i, pi. liii., 

167, evidently naming Daubenton's La Serotine ; Jenyns ; Bell (ed. i) ; Clermont ; 

Dobson ; Flower and Lydekker ; Miller ; Thomas, Zoologist, 1898, 100 ; Johnston ; 

Cabrera ; Millais. 
i8o6. VESPERTILIO NOCTULA, Isidore Geoffroy, Ann. dii Miis. d^Hist. iVa/., viii., 193, 

pis. 47 and 48 ; transference of name in error. 
181 1. VESPERTILIO MURINUS, P. S. Pallas, ZoograpMa Rosso-asiatica, i., 121, No. 

46 ; preoccupied by V. murinus of Linnaeus, 1758. 
1811. VESPERTILIO SEROTINA, auct. et Op. cit., 123. 
1827. VESPERTILIO WIEDII, C. L. Brehm, Ornis (Jena), iii., No. 3, 24 ; described 

from Jena, Germany. 
1827. VESPERTILIO OKENI, atect. ct Op. cit., iii., No. 4, 25 ; described from Jena, 

Germany. 
1829. VESPERTILIO RUFESCENS, ai(ct. cit., Oken's Isis (Jena), 643 ; described from 

Jena, Germany. 

1838. SCOTOPHILTJS SEROTINUS, J. E. Gray, Mag. Zool. and Bof., 4g7 ; MacGillivray; 
Bell (ed. 2). 

1839. Vesperugo (vesperus) serotinus, a. Graf von Keyserling and J. H. 
Blasius, Wiegmann's Archiv fiir Naturgeschichte, i., 313 ; Dobson ; Blanford. 

1844. VESPERTILIO INCISIVUS, J. Crespon, Faune Mcridionale, i., 26 ; described from 

Nimes, France. 
1844. VESPERTILIO PALUSTRIS, auct. et Op. 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, Sdugethiere DeutscJilands, 76 ; Lydekker. 
1 862- 1 863. Cateorus serotinus, var. typus, var rufescens, and var. pallidus ; 

Carl Koch., Jahrbiicher des Vereins fiir Naturkunde im Herzogthu7n (Nassau), xviii., 
463-467, etc. ; described from middle Europe, south Europe, and the "die steppen 
Siid-Russlands " respectively : the latter apparently = V. turconianus of Eversmann. 

1870. Vesperus serotinus, L. J. Fitzinger, SitzimgsbericJite der Kaiserlichen Akad. 
der Wissenschaften (Vienna), Ixii. (i), 88. 

1900. Eptesicus serotinus, Mehely Lajos, Monographia Chiropteroritm Hungariae, 
209, 340, pi. xiv. ; Miller ; Trouessart (1910). 

La Serotine of the French ; die spdtfliegende Fledernimis of the 
Germans. 

Serotine from the French serotin, from the Latin serotinus, i.e., late^ 
or backward. 

" Rattle-mouse" of Wight (Guyon, Zoologist, 1856, 5216). 



THE SEROTINE 131 

Distrilbution : — Bats of this type have a remarkably wide distribu- 
tion, being found over the greater part of the known world. In 
the Palaearctic 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. fusais of Beauvois ranges 
throughout the austral, transitional, and the lower edges of the 
boreal zones, and has sub-species in (i) 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. i), 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 185 1 of Bury and Borrer in 
the Isle of Wight {Journ. cit., 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, Field, 
14th Oct. 1893, 597, and Zoologist, 1893, 45^; Jeffrey, 1894, 261; 
Butterfield, 1897, 141); in Hampshire {lAX^oxd, Joiirn. cit., 1887, 65; 
Kelsall, 1891, 395); in Kent (Harting,/c'?^;7/. cit., 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 Forth (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 VESPERTILIONID^— 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. at., 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 {Zoologist, 1892,403). A reputed example 
in the Nev/castle Museum has been examined by Southwell, and proves 
to be a Noctule {Journ. cit., 1887, 234), and Lilford's belief that it 
occurs in Northampton {Joiirn. cit., 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 {Op. cit. supra, 
p, 83, I, 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 Nyctalus {?\2Xq. IV., Fig, 3, p. 60). 

The wing (Plate IX., Fig. 2, p. 126) is broader than in N. noctula, the 
lower leg, tail, forearm, and fifth metacarpal being all longer than in 




Natterer's Bat (J natural size.) 



THE SEROTINE 



133 



that species. The third metacarpal is slightly shorter than in noctula, 
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 vertebrse being free. The thumb 
is longer, and more slender. The total expanse is actually about the 
same as in N. noctula, but is less relatively to the forearm length. 

The fur is soft and longer than in Nyctalus, 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 N. 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 mciilt. 
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 
melanic and a brown, and seems to be 
contrary to Bell's statement that "the 
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 

^ A similar sexual coloration is, however, present in Miniopterus dasythrix 
(Temminck) and M. fraterculiis of Thomas and Schwann (see Proc. Zool. Soc. 
(London), 20th Feb. 1906, 161 -162). 




Fig. 12. — Front View of Incisors 
AND Canines of Vespertilio serotinus 
(enlarged and diagrammatic). 



N 2 



134 



VESPERTILIONIDtE— 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 transylvamis, and Cabrera one with the forearm measuring 
51 mm. from the Balearics as insularis ; the same author's (^t^j-^^/ 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 isabellinus (Tem- 
minck, 1827, forearm 41, from Tripoli), turcomanus (Eversmann, 
1840, expanse about 200, from the Aralo - Caspian steppes), and 
bottce (Peters, 1869, forearm 40, from Botta, Arabia), are clearly not 
conspecific. Mirza (Philippi, 1865, forearm 53), and shiraziensis 
(Dobson, 1 87 1, 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 








_^ 












o 




•a 
a 


■a 

_2 




'5 


u 




o 


a 


■a 


Vo 

S 






a 








OS 


H 


J 




1 


a 


o 




1 


1^ 




w 




i-i 
cm 










H 


1^ 








Male (in the flesh), Essex 


























(Miller Christy, Zoologist, 


























1S94, 424) .... 
























355 


Male do., Yalding, Kent, 2Sth 


























July 1906 .... 


79 


20 


8-5 


51-5 


20 


11-5 


50-5 


8 


85 


47 


47 


348 


Male, do., Surrey, 6th Aug. 


























(W. R. Ogilvie-Grant) . 


68 


19 




52 


22 




53 




89 


47 


43-5 




Male, Brede, Sussex . 














49 












Female (in the flesh), Yalding, 


























Kent (W. R. Ogilvie-Grant), 


























7tb Aug. 1897 


72 


15-5 




52 


20-5 


9 


53 


7 


89 


49-5 


46 


354 


Female, do., Sth Aug. 1897 


73 


16 




55 


21 


10 


54 


7 


87 


47 


45 


355 


Female, do., 13th May 1898 


80 


19 




52 


21 




51 


7 


84 


45-5 


41 


374 


Female, do., 3rd July 1898 


73 


23 


c" 


54 






53-5 


6-5 


85 


48 


44-5 


374 


Female, do., 2nd Oct. 1898 


73 


17 




48 


20-5 


10 


53 


7 


90 


47 


44 


365 


Female, do., 2nd Oct. 1898 


75 


15 




50 


21 


12 


52-5 


7 


84 


45 


41-5 


356 


Female, Surrey, 6th Aug. 1902 


























(W. R. Ogilvie-Grant) . 
Average of females 


75 


20 


9 


57 


















73 


17-9 




52-5 


20-8 




52-8 


6-9 


86-5 


47 


43-5 


368 


Do. of both sexes . 


74-2 


18-25 




52-3 


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 145 and 
15 inches = 86S-8 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, f oz.= 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 orained 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 

' Zoologist^ 1874, 4126. 2 Proc. Linn. Soc. (London), 6th Dec. 1853, 260 (1855). 
^ Field., 14th March 1874, 263 (quoted by Edward Newman). 
* Zoologist., 1 891, 305-306 and 425. 



136 VESPERTILIONID^— 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 Myotis. 

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 around. 

o 

To the above notes may be added the information kindly 
supplied by the Rev. E. N. Bloomfield, of Guestling Rectory, 

' E.g.^ Blasius in Germany, Borrer and Bond in England; see Zoologist, 1891, 
203. 



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. 

' Porthesia chrysorrkcea, Linnaeus. 

- 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 VESPERTILIONID^— 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 a colony 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 loth ; 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 /^/^^ i3c,4^ 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.). 

* 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, Linnajus.i 

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 jttst 
deady It was taken (as he wrote to Mr Thomas Southwell), 
" I 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. 
Possibly 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 eng^aofed 
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 Syste7na Naturce, x., 32 (7), 1758 : until recently cited as Vespertilio discolor of 
Natterer. 

- No. 37a. 

^ Southwell, Trans. Norfolk and Nonvich Nat. Soc, 1873-74, 80 ; (I am indebted 
to Southwell for further particulars by letter); E. Newman, Field, 7th March 1874, 
218, and Field, 5th Sept. 1874, 246. 

■* V. nilssoni. 



I40 VESPERTILIONID^— MYOTIS 

It ranges from southern Sweden to Dalmatia, and from 
France to Asia, and is, perhaps, representative of the American 
genus Lasionyderis. 

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, Sy stein der Europaischen Thierwelt, i., 105-106, 188 ; 

based on Vespertilio tnuHniis of Schreber (not V. miirmus of Linnceus) ; Gray, 

Ann. and Mag. Nat. Hist., December, 1842, 258. 
1829. Nystactes, Jakob Kaup, op. cit., i., 106, 108-109; based on Vespertilio 

bechsteini of Leisler \ name preoccupied by Nystactes of Gloger, 1827, a genus of 

birds (see Palmer, Index, 467, 1904). 

1829. Vespertilio, Jakob Kaup, op. cit., i., no; based on Vespertilio daubentoni, 
V. nattereri, and V. mystacinus ; A. Graf von Keyserling and J. H. Blasius, 
Wiegmann's Archiv f'iir Naturgeschichte, i., 307, 1839; Harrison Allen, 1864 and 
1893 ; Dobson, 1878 ; not VESPERTILIO of Linnaeus, 1758. 

1830. Leuconoe, Friedrich Boie ; Oken's his (Jena), 256-7 ; based on die Wasser- 
fledermause, type V. daiibentonii. 

1841. Selysius, C. L., Prince Bonaparte, Fauna Italica, i., introduction (3); based 
on Vespertilio mystacinus of Leisler. 

1841. Capaccinius, C. L., Prince Bonaparte, Fauna Italica, i., introduction (4) ; based 
on Capaccinius megapodius of Bonaparte = Vespertilio capaccinii of Bonaparte. 

1842. Trilatitus, J. E. Gray, Ann. and Mag. Nat. Hist., 258, December: in- 
cluded Vespertilio hasseltii of Temminck, V. viacelliis of Temminck— V. adversus 
of Horsfield and T. blepotis of Gray =^ Miniopterus sp. 

1849. Tralatitus, F. L. P, Gervais in Charles D'Orbigny's Dictionnaire Universel 

d'hist. Nat., xiii., 213 ; a misprint for Trilatitus. 
1856. Brachyotus (sub-genus), F. A. Kolenati, Allgemeine deutsche NattirJiist. 

Zeitung (Dresden), Neue Folge, ii,, 131, 174-177 ; based on mystacinus of Kuhl, 

daubentonii 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, op. cit., ii., 131, 177 ; based on nattererii of 

Kuhl, and ciliatus (p. 131) or emarginatus of Geoffroy (pp. 177-179). 
1856. Myotus, F. a. Kolenati, op. cit., ii., 131, 179-181 ; based on bechsteinii oi Kuhl 

and murinus of Linnaeus. 



MYOTIS 141 

1866. Tralatitius, J. E. Gray, Ann. atid Mag. Nat. Hist., 90, February; a mis- 
print for TiHlatitiis. 

1867. Pternopterus, Wilhelm Peters, Monatsberichie der Koniglich Akad. der 
Wissenschaften (Berlin), 706 ; a sub-genus of Vespertilio = MVOTIS, based on 
P. lobipes of Peters = V. muricola of Hodgson. 

1870. EXOCHURUS, L. J. Fitzinger, Sitzungsberichie der Kaiserlichen Akad. der 
Wissenschafteji (Vienna), Ixii., i, 75-81 ; based on Vespertilio inacrodactylus of 
Temminck, V. //^/-^T?]?/^// of Temminck, and V. inacrotarsus of Waterhouse. 

1870. Aeorestes, L. J. Fitzinger, op. cit., 427-436 ; included Vespertilio villosissiiniis 
of Geoffroy, V. albescens of Geoffroy, V. nigtHcans of Maximilian, and V. levis of 
Geoffroy. 

1870. CoMASTES, L. J. Fitzinger, op. cit., 565-579 ; included Vespertilio capaccini of 
Bonaparte, V. nicgapodius of Temminck, V. dasycneme of Boie, and V. limnophilus 
of Temminck. 

Classification and synonymy : — As shown by Mr Miller, the 
genus Vespertilio of Linnaeus 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 Myotis 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- — daubentoni, myslacinus, 
the rare becksteini, and nattereri. A fifth, the common con- 
tinental M. myosotis (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, M. emarginatus, is stated 
to have occurred once (see page 189). Other European 
species are M. capaccini of Bonaparte, a bluish-grey bat, with 
narrow, hairy, interfemoral membrane and elongated backward 
pointing, falcate tragus ; M. dasycneme of Boie (see page 157) ; 
and the Mediterranean M. oxygnatJms (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 Nyctalus, Pipistirlhis, 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 strenorth accord- 
ing to the size of the species : it has 
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 Nyctalus and Vespertilio ; the zygo- 
mata are flattened, the auditory bullae 
moderately developed. 

The number of teeth (Figs. 8, p. loi, 
13, and 16, p. 174) is increased to 
thirty-eight by the addition of a third premolar in each jaw and 
side, so that the formula is — 

1 ..,3-3 





Fig. 13.— Diagram of Arrange- 
ment OF Teeth in Genus 
Myotis. 

(i) Upper and (2) Lower Jaw. 



pni 



38- 



.2-2 I - I 

3-3 i-i' ^ 3-3' 3-3 

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 Leitconoe : — 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 M. 
daubentoni is the sole British representative, and the members 
of which are characterised by their large feet. This character 
in dattbentoni, sufficiently evident as it is, reaches an extreme of 
development in ricketi of Thomas. For these bats the name 
Letcconoe, 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 M. nattereri is a more 
distinct character. 

DAUBENTON'S, OR THE WATER BAT. 

MYOTIS DAUBENTONI (Kuhl). 

1819. Vespertilio daubentoni, Heinrich Kuhl, Neue Antt. der WetterauiscJien 
Gesellschaft fiir die gesammte Naiurkimde, i., ii., 195, pi. xxv., fig. 2 ; described from 
Leisler's MSS. from Hanau, Germany ; Bell (ed. i) ; 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. LeuconoE daubentoni, Friedrich Boie, Oken's Isis (Jena), 256-257. 

1839. Vespertilio ^dilis, Leonard Jenyns, Ann. Nat. Hist.., iii., -Ji., pi. iii., April ; 
described from an albinic variety, from Auckland St Andrew, Durham, England. 

1840.? Vespertilio volgensis (? species), Eduardo Eversmann, Bull. Soc. Impcriale 
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, Faune MMdionale, i, 16 ; vide 
Trouessart, Bull, de la Soc. d' Etude des Sci. Nat. de Nimes, 7, i., 35-39, 1879 ; 
described from Nimes, France. 



144 VESPERTILIONID^— MYOTIS 

1855. Vespertilio daubentonii, var. emarginatus, G. B. Buckton, Proc. Linnean 
Soc. (London), 6th Dec. 1853, 260. 

1856. Brachyotus daubentonii, F. a. Kolenati, Allgemeine deiitsche Naturhist. 
Zettung {'Drfsd&n), Neue Folge, ii., 131, 175-176. 

1887. Myotis CILIATA,^ J. Daday, Ertekezesek a Tenneszettudomdfiyok Korebola 
(Budapest), xvi., 7, 35, 1886 ; not Vespertilio ciliatiis of Blasius, vide Mehely, 330. 

1898. Myotis daubentoni, Oldfield Thomas, Zoologist, 100; Collett ; Johnston; 
Mehely ; Cabrera ; Millais. 

1 9 10, Myotis (leuconoe) daubentoni, E.-L. Trouessart, Faune des Mammiferes 
d' Europe, 27. 

Die Wasserjiedernians 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 
gardens 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 

^ Vespertilio capucinellus and Vespertilio minutillus are cited under this species 
as of " Koch, Bayr. Fauna" but I cannot find the original reference ; see Fitzinger, 
Sitztcngsberichte der Kaiserlichen Acad, der Wissenschaften (Vienna), Ixiii., i, 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, Field, 31st July 
1897, 222) ; it was found abundantly in Perth by Grieve {Ann. Scott. Nat. 
Hist., October 1894, 193-195), and in Solway is commoner than P. pipi- 
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 {Irish Naturalist, 1897, 39). It was next encountered 
by J. R. Kinahan in Kildare in 1853, ^i^d was at first identified with M. 
nattereri {Nat. Hist. Review (Dublin), i., 23-25 and 87, 1854), but assigned 
by Bell to the present species {op. cit., i., 148-149; also vi., 383, 1859). 
Subsequently Jameson {Irish Naturalist, loc. a'/., 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 {Irish 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 Myotis niystacinus, by E. T. Newton {Quart. Jour n. Geol. 
Soc, August 1899, 420; see also Ahhot, Journ. cit.. May 1894, 171-211), 
as have leg-bones from superficial deposits in the caves of Co. Clare, 
Ireland (Scharfif, Trans. 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 oi Pipistrellus pipistrellus. 

Breeding season and number of young : — A single young one is 
born, most frequently in June and July. 



146 VESPERTILIONID^— MYOTIS 

Description : — In general form and appearance this bat is typical 
of the group Leiiconbe. In size it is somewhat larger than Pipistrellus 
pipistrellus^ but distinctly smaller than Nyctahis leisleri. 

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 -mng (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 vertebrae. 

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 XI., 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- 
tilio (Bdilis was based on an albino specimen. 

In the skull (Fig. 8, No. 2, p. loi) the profile of the cranium is almost 



DAUBENTON'S, OR THE WATER BAT 



147 




Fig. 14. — Front View of In- 
cisors AND Canines of 

My Otis dauhenloni (enlarged 
and diagrammatic). 



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 M, carissivia of North 
America been taken in England, it would 
have passed muster as daubejitoni. 

Dimensions in millimetres (see page 
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 111^ 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 {^Zoologist, 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. 

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 



VESPERTILIONID^—MYOTIS 



DIMENSIONS IN MILLIMETRES:- 



Males. 






-a 








^ 












<*- 




>. 


M) 


.d 






o 




^ 


. 


,_; 




o 

0) 




•tt 


a 








.a 




.5 


40 






m 




.8 


1 

bD 


a:) 


I 


to 

% 
o 


■73 


a 

a 
o 


"3 
a 

a 

3 


]3) 

60 
PI 
O 


1 


1 

cS 
a 


ID a 




a 




to 










.a 
H 




^ 


•^ 


£ 


Male (from spirit), North- 


























ampton .... 










17 


9 


37-5 


5 


57 


31 


29-5 




Male, Middlesex 










17-5 


8 


38 


6 


58 


32 


30 




MaIe(f'rom .spirit), Temple 


























Combe, England, Oct. 


























1907 .... 


50-4 


12-6 


6 


30'6 


15 


9 


35-6 


7 


59 


32 


29-2 


244 


"Male, Merioneth, Wales, 


























31st March, 1891 . 


41 


12-5 


6-75 


35-5 






33 










202 


*Male, Strathspey, Scot- 


























land, 6th Sept. 1891 . 


43 


9-5 


7 


36 


17 


lit 


36 










226 


*Male, Loch Dochart, W. 


























Perth, Scotland, 25th 


























July 1894 


40 


12-5 


7-4 


35 


17 


7-5 


38 










227 


*Male, do., 25th July 1894 


4t} 


14-25 


7-4 


33-5 


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 


lOt 


38 










227 


*Male, do., 15th July 1894 


49-5 


17-5 


6-4 


31-5 




lOt 


34-25 










201 


Male, Bohoe Cave, Fer- 


























managh, Ireland, 11th 


























July 1895 (Dublin 


























Museum), from dried 


























skin .... 








26 


15 


10-25 


35 


5-6 


57 


31 


30 




Male, Co. Louth, 6th July 


























1896 (Dublin Museum), 


























from dried slciu . 








31 


15 


10 


34-25 


5-5 


60-5 


32 


30 




Two males (from spirit),^ 
Bohoe Cave, Fer- | 


























44 






30 


18 


10-25 


38-5 


7 


64 


34 


32 




managh, Ireland (Dub- C 
lin Museum) . . J 

Average of males . 


46 








17-5 


9 


37 


7 


60 


32 


30 




























49-1 


13-4 


6-6 


32-7 


16-3 


9-5 


36-4 


6-1 


58-8 


32 


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. 


51 


13 




35 




10 


35 












Female, Hampshire (mea- 


























sured from spirit at 


























British Museum) . 










17 


9 


38 


6 


63 


34 


31-5 




*Female, Gordon Castle, 


























N.-W. Banff, 14th July 


























1892 .... 


45-5 


13 


6-5 


37 


10-5 


8-5 


36 


10-5 








227 


*Female, Loch Dochart, 


























W. Perth, 15th July 




. 






















1894 .... 
Average of females . 


44 


18 


6-5 


37 


16-75 


10-5t 


39 










227 


46-7 


12-7 




34-8 




9-4 


36-7 












* Female, Yorkshire, one 


























day old (A. Whitaker) . 


29-25 






15-75 














76 


82 


*Female, Loch Dochart, 
























approx. 


W. Perth, young, 15th 


























July 1894 . 


29 






18-5 


9-5 


8-5 


18-5 








140 


101 



Measured by Eagle Clarke in inches, and converted to mm. 



t With claw. 



PLATE IX. 




Wings (A milural size and diagrammatic) OF 

I . Nyctalus noclula. 2. Vespertilio serotinus. 

3. Rhmolophus ferrum-tujuinuin. 



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- 

* Letter xi. to Pennant, dated 9th Sept. 1767, original edit., 1789, 32. 
'^ V, emarginatus of Geoffroy. •* Vespertilio cedilis. 

■* Zoologist, 1889, 434. 

O 2 



1 50 VESPERTILIONID^— 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. 

^ 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 151 

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 nth 
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 

* Ann. Scott. Nat. Hist., 1896, 201. ^ Irish Naturalist, 1896, 94. 

3 Proc. Dublin Nat. Hist. Soc, Dec. 9, 1853, reported in Nat. Hist. Review 
(Dublin), i., 1854,23-25 ; Zoologist, 1853, 4012-4013, 



152 VESPERTILIONIDiE— MYOTIS 

polished as to Indicate a large colony, and the little animals 
voiced their protests by a loud outcry, descrfbed 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 

^ In the quotation, Daubenton's has throughout been substituted for Natterer's 
Bat, for which species Kinahan mistook it (see p. 145, under "Distribution"). 
^ Ann. Scott. Nat. Hist., 1894, 193-195. 



PLATE X. 




SeroTINE Bat. (Natural size.) 



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. 
^ Irish Naturalist, 1897, 39. 



154 VESPERTILIONIDyE— 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 Irish Naturalist^ 1899, 31 and 124. ^ Op. cit., 202. 

^ Irish Naturalist, 1905, 106-107. 

* Oldham has seen it flying at 2.45 A.M., or seventy-five minutes before sunrise, 
on 13th July. 

^ Zoologist, 1874, 4128. " Irish Naturalist, 1899, 33. 



DAUBENTON'S, OR THE WATER BAT 155 

27th April/ Mr Oldham tells me that he has seen it on 
the wing in Cheshire on the 19th, 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 igth,^ 
while Messrs Coward and Oldham, on the 28th, received two 
fledglings, as well as an adult female, recently a mother. 

^ Journ. Birmingham Nat. Hist, and Philosoph. Soc.^ Jan. and Feb, 1896, II., i., 7. 

2 Zoologist, 1887, 293. 

3 Faune de la Normandie, i., 149 : Paris, 1888, * Zoologist, 1892, 403. 

^ In an instance observed by Professor R. Collett, the mother hung head down- 
wards {in lit.). 

^ Naturalist, March 1907, 74 : unfortunately this bat died almost immediately. 



156 VESPERTILION ID^— 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 

^ Xylophasia polyodon, Urapteryx sambucaia, Triphcena pronuba, Mamestra 
brasskte, Cidaria populata, and Porthesia auriflua, the latter accepted with reluctance : 
Zoologist, 1899, 472-473- 



PLATE XI. 






Mmvr^'' 




Heads (natural size) OF 

1. Mvotis niystaciiius. 3. M vol is nattereri. 

2. Myotis dattbenlonL 4. Pipishellus pipistreHiis. 



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 
manoeuvre, 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).^ 

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 emargmatus, 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 

' Dobson, Catalogue of Chiropter a, 285. 

^ F. Boie, Oken's Isis (Jena), 1825, 1200. 

^ Zoologist, 1854, 4361 ; 1856, 4938. 

* Proc. Linnean Soc. (London), 1853, 260 (pub. 1855), 

^ Catalogue of Chiroptera, 296. 



158 VESPERTILIONID^— 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, Nciie Ami. der Wetterauischen 
Gesellschaft fiir die gesaminfe Naturkunde^ i., ii., 202 ; described from Leisler's 
MSS., from Hanau, Germany; Bell (ed. i) ; Jenyns ; MacGillivray ; Blasius ; 
Clermont; Fatio ; Bell (ed. 2); Dobson ; Harting, Zoologist, 1888, 161-166, pi. 2, 
and 441, pi. 3 ; Blanford ; Flower and Lydekker ; Winge. 

1821. Vespertilio COLARIS, H. R. Schinz, Cuvier's Das Thierreich, i., 177; 
described from the collared phase (see description below), from Meisner's MSS., 
from Mont Blanc, Switzerland; type lost, vide auct. cit., Etiropciische Fauna, 17, 
1840. 

1827. Vespertilio schinzii, C. L. Brehm, Omis (Jena), iii., 27; described from 
Renthendorf, Germany. 

1833. Vespertilio humeralis, L. A. F. Baillon, Man. de la Soc. Roy. d^ Emula- 
tion d'' Abbeville, 50 ; described from the phase with black shoulder-patches (see 
description below), from Abbeville, France. 

1838. Vespertilio emarginatus, William MacGilHvray, British Quadrupeds, 96 ; 
not V. emarginatus of Geoffroy. 

1841. Selysius MYSTACINUS, C. L., Prince Bonaparte, Fauna Italica, i., introduc- 
tion (3). 

1844. Vespertilio latipennis, J. Crespon, Faune Mcridionale, 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. Imperiale des Nats, 
de Moscou, xviii., 505, pi. 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. 

Zeitutig (Dresden), Neue Folge, ii., 131, 174-175 ; not Vespertilio ciliatus of 

Blasius, 1853 = V. emarginatus of Geofifroy. 
1862-63. Brachyotus mystacinus, var. nigricans, var. rufofuscus, and van 

AUREUS, Carl Koch, Jahrbiicher des Vereins fiir Naturkunde itn Herzogthum 

(Nassau), xviii., 440-448 ; see also Zoologische Garten, xi., 3689, 1870 ; described 

from Nassau, Germany. 
1869. Vespertilio mystacinus, var. nigricans, Victor Fatio, Vertebrcs de la Suisse, 

i., 92 ; described from Switzerland : see Mottaz, loc. cit. 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) van 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, Sitzioigsberichte 
der Kaiserlichen Acad, der Wissenschaften (Vienna), Ixiii., i, 217; renaming 
Brehm's Vespertilio schinzii. 

1871. Vespertilio mystacinus schrankii,i auct. et op. cit., 219. 

1871. Myotis brandtii, auct. et op. cit., 284. 

1898. Myotis mystacinus, Oldfield Thomas, Zoologist, 100 ; Collett ; Mehely ; 

Johnston ; Cabrera ; Millais, 99, pi. 12. 
1910. Myotis (myotis) mystacinus, E.-L. Trouessart, Faune de Mammiferes 

d'Europe, 33. 

Le Vespertilion AToustac of the French; die Bartfledermaus of the 
Germans ; but these are book-names only, the bat being probably 
unknown to popular observation either in this country or 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. M. siligorefisis (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 Pipistrellus pipistrellus 
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 Pipistrellus pipistrellus (^Journ. cit., 1888, 164-165). 
Similarly, Oldham's Shropshire record of 1890 {Journ. 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 filr Naturgeschichte, ix., 25, 1843) mentions 
V. schrankii as of Koch, but I cannot find the original description. 



i6o VESPERTILIONID^— MYOTIS 

To come to details ; in spite of scarcity of information, Harting, 
who collected the available data up to i8S8 {Zoologist, 1888, 161-166), 
was able to cite occurrences in Somerset, Dorset, Hampshire with 
Wight (see also More, Joiirn. cit., 1894, 148; Wadham), Sussex, Kent, 
Essex, Cambridge, Northampton, Warwick, Worcester, Stafford, and 
one or two other counties (see also Cocks, Journ. cit., 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 {Joiuni. cit., 1889, 68-69), ^"*^ 
Lancashire {Journ, 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. pipistrellus 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, Journ. cit., 1906, 70), is very likely the most abundant 
bat of Merioneth (Caton Haigh, Journ. cit., 1887, 294), and has a 
general reputation for being not uncommon in the north of the princi- 
pality (Caton Haigh, Journ. cit., 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 m»ay 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, op. cit., 165), but Macpherson mentions four for Cumber- 
land, including one near the Scottish border. 



THE WHISKERED BAT i6i 

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 {pp. cit., 165), a clue which 
enabled W. Evans to work out its entire history {Mammalian 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 {Ann. 
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 
{Nat. Hist. Review (Dublin), i., 24, 1854; see also, vi., 383, 1859). This 
was at first recorded as an example of AT. daiibeyitoni, but the mistake 
was corrected at a meeting of the same Society on 12th May 1854 
{Nat. Hist. Review (Dublin), i., 148, 1854). In 1897 {Irish 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, /(3Z^r;2. cit., 1898, 272; AXcock, Journ. cit., 1899, 57). 
including Lambay Island (Baring, Journ. cit., 1907, 19) : Down (Lett, 
Proc. Belfast Nat. Field Club, 20th March 1900; R. Patterson, Irish 
Naturalist, 1900, 162): Wexford iJAo^dX, Journ. cit., 1902, 103; Barrett- 
Hamilton, Journ. cit., 1908, 207) : and Carlow (Pack Beresford, Jourji. 
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 M. da?ibentoni. 

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 Leuconoe, and it is thus readily 
distinguished from the Pipistrelle, than which it is larger. It is smaller 
than M. daubentoni. 



i62 VESPERTILIONIDvE— MYOTIS 

The forehead is elevated, the occiput prominent, the muzzle pointed, 
the nostril moderately tumid (Plate XI., Fig. i, 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. i,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 AT. daiibentoni. The relations of the lower leg and 
foot are remarkably different, the length of the latter being only about half 
that of the formier as against nine-sixteenths in M. dmibentoni. 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 colaris 
and humeralis. 

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. loi) the profile is saddle-shaped, or like 
a shallow S ; the frontal region is depressed, and the narrow muzzle 
about horizontal. The bullae are small. 

The teeth are similar to those of M. 
daubentoiii, but the outer lower incisors are 
not so long in section (Fig. 15). 

Individual variation runs frequently 
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. auremn" from 
south Germany and the plains, and the 
typical " var. rufofuscum!^ Probably all 
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, 575 ; from posterior border of m^ 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 



Fig. 15.— Front View of In- 
cisors AND Canines of Myotis 
mystacimis (enlarged and dia- 
grammatic). 



164 



VESPERTILIONID^— 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 

ci 


,a 

-a 

c 

1 

£ 
to 


ta 
to 


■5 





§ 
.a 

Si 

■a 
-S 
W 


a 

03 

i 




"O 

a 

s 

.a 
H 


'So 

M 
§ 


t 

03 

1 
1 


> 

a 
1 



a 

a 
a . 

£ 



1 


Male (in spirit), Burring- 
ton, Cheddar, Jan. 1907 

Male, Macclesfield (dry 
skin in British Mu- 
seum) .... 

Male, Co. Louth, ISth 
July 1005 (Dublin 
Museum) . 

Male (in spirit), Co. Fer- 
managh, nth July 
1895 (Dublin Museum) 

Male, do. (do.) (no dale) 

Female, Burrington, 
Cheddar, Jan. 1907 . 

Female, Merioneth, 8th 
July 1892 (W. E. 
Clarke) 

Average (approximate) 


50 

44 

42 

48 


11 
14 


7-5 


30 

31-5 

35 


18 
14 

If) 
lG-5 

10 
14 


7'5 

7 
7 

7-5 
8 


37 

35 

32 

33 
34 

34 
33 


6-0 

5-25 
5-25 

5 
5-5 


55 
53 

53-5 
53-25 

55 


30 
30 

28-5 
28 

29 


30 
28 

27 
26-25 

27 


228 


5-45 
grms. 


46 








15-5 


7-5 


34 


5-5 


54 


29 


27-5 






Male, young (in spirit), 
Burrington, Cheddar, 
Jan. 1907 . 


48 








le 


7-5 


34 


5-5 


54 


28 


26-75 







The Whiskered Bat is probably the commonest British 
representative of its genus. It was described by the German 
naturaHst 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 

^ Zoological Journal, ii., 109. 



PLATE XII. 




Whiskered Bat. (Natural size.) 



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, as a 
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. " Naturalist^ 1899, 74. 

3 As remarked also by Caton Haigh, Zoologist^ 1887, 294. 



i66 VESPERTILIONID^— 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 

^ Worcestershire, i., 174. - Naturalist, 1897, 242. ^ Zoologist^ 1874, 4128. 
* Naturalist, 1899, 74. ^ 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 lisfht 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 

' Zoologist^ 1874, 4128. - Tomes in BelL 

^ N. H. Alcock, Irish Naturalist, 1898, 272. 

■^ W. D, Roebuck, Naturalist, 1886, 113. ■'"' Zoologist, 1896, 255, 



\ 

V 

i68 ^^ VESPERTILIONID^— 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.^ 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 G. 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 r.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. 

^ Gonoptera lihairix and Scotosia dubitata. 

2 Of which the most abundant are Blepharoptera serrata, Borborus niger, and a 
species of Culex. 

3 Zoologist, 1906, 186 ; also, Wilson, in lit. * Zoologist, 1874, 4128. 
^ Naturalist^ 1897, 242. " Charbonnier, see Somerset, i., 164. 



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 i8th 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 fiies.^ 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. 

"It evinced," writes Mr Oldham, "little disposition for 

' Zoologist^ 1901? 315 ; 1904) 311- ^ Irish Naturalist, 1902, 103. 

^ Diptera, and that some pieces of legs and wings clearly belonged to a small 
Tipulid — very possibly a species of Trichocera. 

* Naturalist, 1897, 242. ^' Irish Naturalist, 1899, 56. 

'' Zoologist, 1899, 49-53. "^ Scotosia dubitata and Gonoptera libatrix. 

Q 



I70 VESPERTILIONID^— 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 J 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 a lamp. 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. In a 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. 

- Zoologist, 1899, 51-53. Norton {loc. cit., 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. 



172 VESPERTILIONID^— 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, Neue Ann. der Wetterauischen 

Gesellschaft fiir die gesammte Naturkunde, I., i., 30, pi. 22 ; described from Hanau, 

Germany, from Leisler's MSS. ; Jenyns ; Bell (ed. i) ; MacGillivray ; Blasius ; 

Clermont ; Fatio ; Bell (ed. 2) ; Dobson ; Flower and Lydekker ; Lydekker ; 

E. T. Newton, Quart. Joum. Geol. Soc. (London), August 1899, 420 ; Winge. 
1829. Nystactes bechsteinii, Jakob Kaup, System der Europdischen Thierwelt, 

i., 106, 108-9. 
1842. Myotis bechsteinii, J. E. Gray, Ann. and Mag. Nat. Hist., Dec, 258 ; 

Fitzinger ; Thomas, Zoologist, 1898, 100; Mehely ; Johnston; Millais. 
1856. Myotus bechsteinii, F. a. Kolenati, Algemeine deutsche Naturhist. Zeitung 

(Dresden), Neue Folge, ii., 131 and 179. 



PLATE XIII. 






Wings (| natural size and diagiammatic) OF 
I. My Otis jnyslacinus. 2, M. bechsteini. 3. M. 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, Ann. and Mag. Nat. Hist.., 

Sept., 220 ; described from La Granja, Sierra de Guadarrama, Central Spain. 
1910. Myotis (myotis) bechsteini, E.-L. Trouessart, Faune des Mammiferes 

d^ 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 M. inurinus = myosotis {Quart. Jourji. 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 Nyctalus leisleri, the lower leg is but 
little shorter than that of Vespertilio 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 B. barbastellus. 

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. 

Q 2 



174 



VESPERTILIONID^— MYOTIS 



The wing is therefore of expanse under the average, broad at its 
base, with an ample interfemoral membrane. It is poorly developed 

at the extremity. The thumb 
is long. 

The fur is soft and woolly. 

The face (Plate XIV., Fig. i) 

is thinly haired except on the 

I forehead, and there are a few 

moustache - like hairs on the 







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 M. fiatterefi, the bases of the 
hairs being everywhere dusky, with the tips some shade of wood-brown 




Fig. 16, 



-Side View (diagrammatic and enlarged) 
OF Skulls and Teeth of 



I. Nyctalus leisleri. 2. Myotis bechstemi. 
3. M. nattereri. 



BECHSTEIN'S BAT 



175 



above, whitish beneath. The moderately distinct Hne of demarcation 
passes along the Hne of junction between wing and body and thence 
to the base of the ear. 

The skull, as compared with that of M. nattereri, is larger, but 
narrower ; it carries more prominently developed cranial ridges and 
much bigger bullae (Fig. 16, No. 2, p. 174). 

The teeth are similar to those of M, 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 favotiicus on the ground of their 
markedly smaller ears and bullae. I cannot distinguish the few British 
specimens available from the type of favonicus, but await further 
material before identifying the British Bechstein's Bat with that form. 
Both are distinctly smaller than the Hungarian. 





DIMENSIONS 


IN MILLIMETRES 


: — 














^ 


































3 




+3 








o 




>^ 


"S 


.£3 






O 




!3 








<o 




13 


S 








fi 




O 






> 






a 

1 


.2 

1 
t 


^1 


s 


IS 

3 




a 


=1 


s 
o 




1 

O 


<o a 




W 




u 






a 

13 




A 


ij 


% 


a 


s 
o 


Two, New Forest, July"j 
1886 (B. W. H. Blagg), - 
from old skins . .) 




19 


10 


35 


20 


10-75 


43 


7 


64 


36 


34-5 


248 




lS-5 


9 


35 


19 


10 


42 


7 


64 


33 


33 


252 


Male, Henley-on-Thames, 


























10th March 1901 (J. G. 


























Millais, British Museum), 


























from dried skin 




IS 






19 


10 


39 


7 


59 


30-5 


31 




Male, Isle of Wight, 14th 


























Aug. 1909 (P. Wadham) 


[52] 


19 


8 




20 




42 


7-f 


67 


35 


34+ 




Female, do. 31st July 


























1909 .... 
Average . 


[52] 


19 


8 


34 


20 


8-5 


39 


7-f 


65 


35 


34 






18-7 


8-7 


35 


19-5 


? 


41 


7 


64 


33-5 


33 


[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 n^ 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 VESPERTILIONID^— 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, M. myosotis 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.^ 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 1901, 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.* 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 

^ Zoologist, 1888, 260. ^ Sussex, i., 301. 

3 In Berkshire, not Oxfordshire (see A. H. Cocks, Zoologist, 1910, 74). 
* Recorded by J. E. Harting, Field, 20th Nov. 1909, 889 ; and by J. E. Kelsall, 
Zoologist, 1 9 10, 30. 



BECHSTEIN'S BAT 177 

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. KelsalP 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 unHke 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, i., 98. 

2 Midland Naturalist, 1883, 153. ^ Z^(3/<?gw/, 1884, 483 ; 1885, 146. 
'' Ornis (Jena), 1827, iii.. No. 3, 20. 



178 VESPERTILIONIDyE— MYOTIS 

NATTERER'S BAT. 

MYOTIS NA TTERERI (Kuhl). 

1818. Vespertilio nattereri, Heinrich Kuhl, Neue Ann. der Wetterauischen 
Gesellschaftfiir die gesammte Natttrkunde, i., i, 33, pi. xxiii. ; described from Hanau, 
Germany ; Jenyns ; Bell (ed. i) ; 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, Ann. and Mag., Nat. Hist., Dec, 258 ; 
Fitzinger ; Thomas, Zoologist, 1898, 100 ; Johnston ; Mehely ; Millais. 

1856. ISOTUS NATTERERli, F. A. Kolenati, Algemeine deutsche, Natiirhist. Zeitung 
(Dresden), Neue Folge, ii., 131 and 177. 

1862-63. IsOTUS NATTERERI, var. TYPUS, and var. spelaeus, Carl Koch, Jahrbiicher 
des Vereins fiir Naturkunde im Herzogthum (Nassau), xviii., 426 and 430 ; described 
from Nassau, Germany. 

1 9 10. Myotis (myotis) nattereri, E.-L. Trouessart, Faune des Mammiferes 
d'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 M. n. boinbitius of Thomas 
{Proc. Zool. Soc. (London), 28th Nov., 1905, 337). It ascends to 2300 feet 
in the Caucasus (Satunin). 

M. thysanodes of Miller {North 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 M. 
nattereri 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 Zoologist, 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, Zoologist, 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 
{not 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 {Maniinalian Fauna of Edinburgh, 22-23, ^^d Anii. Scott. 
Nat. Hist., 1 90 1, 1 29- 1 31), 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, ^^^^ presented through M'Coy to 
the Dublin Natural History Society {Proc, 12th February 1845 ; 
J. R. Kinahan, Proc, 9th December 1853, published in Dublin Nat. 
Hist. Review, i., 23, 1854; see also ii., 8, 1855, also vi., 383, 1859; 
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, 

' I 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. 



i8o VESPERTILIONIDiE— MYOTIS 

Louth, Longford (Jameson, Irish Naturalist, 1897, 39), Cork (Folliott 
Darling, Zoologist, 1883, 294), Galway (Kane, Irish Naturalist, 1894, 
116, and 1897, 88; also a specimen from Woodpark, in the British 
Museum), Antrim (Patterson, Journ. cit., 1900, 274), Carlovv (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 M. daubento7ii with this species may have been 
founded upon an actual mixture of the two — see Journ. cit., 1 897, 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 XL, 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 M. daubentojii. 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 

I. Myotis bechUeini. 2 and 3. Barbastella barbastellus (front and side views). 

4. Pie CO tits auritus. 



NATTERER'S BAT 



i«i 



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. i6. No. 3, p. 174) the 
profile resembles that of M. 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. 

Geograpliical variation has not been 
studied, except in the sub-species M, n. 
bombinus of Japan, which is stated to be 
characterised by more abrupt and consider- 
able inflation of the frontal region of the 
skull and by darker colour. 

Dimensions in millimetres (see following 
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 -^fi 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 m^ 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. 




Fig. 18. — Front View of 
Incisors and Canines of 
Myotis nattereri (enlarged and 
diagrammatic). 



l82 



VESPERTILIONID^— MYOTIS 





DIMENSIONS 


IN MILLIMETRES 


:— 












Males. 






.d 




















ti 














;3 












o 




>, 


60 


.a 






O 









1^ 


• 


03 




o 


a 


■a 




■zh 


^ 




O 

.a 


'to 


t^ 


> 


la 




.Q 




oTc' 
















a 


C3 • 




a 


S 




'3 


S 




g 

2 




-3 


1 


03 






03 

•a 


"1 
to 


^1 

TO 


H 


o 


'0 

a 


O 


-a" 


a 
o 


a 


1 
XI 




Male (British Museum) 


42 


16-5 


10 


38 


17 


8 


40 


6 


66-5 


33 


32-5 




Male, Bedfordshire (mea- 


























sured from dried skin) . 


42 


15 




41 




8 














Male, Galway (Dublin 


























Museum) .... 










17 


8 


39 


6 


68-25 


34 


33 












Females. 
















Female, received from A. 


























Whitaker, Yorkshire, 10th 


























October 1906 (measured 


























in the flesh) 


43 






32 


















Female in alcohol (Tomes' 


























collection, in British 


























Museum) .... 




14 


10 




16-5 


8-25 


40 


5 


64 


33 


32 






Female (do.) 






14 


9 




16 


8 


40 


6 


65 


33 


32 






Female (do.) 






14-5 


9-5 




16 


8 


39 


6 


66-5 


34-5 


34 






Female (do.) 






14 


8 




16 


8 


40 


4-75 


69 


35-6 


33 






Female (do.) 






14 


9-75 




16-5 


8-5 


38 


6-5 


63-75 


33 


32 






Female (do.) 






17-5 


9 




16-5 


8 


39 


6-75 


65-5 


33 


32 






Female (do.) 






14 


8 




17 


7-5 


39 


6 


65-5 


34-6 


33 






Female, Nortliants 


46 


14 


9 


s'e 


17 


7-5 


39 


6-5 


65 


36 


33 






Female, do. 


47 


14 


9 


33 


16 


8 


39 


6 


65 


34 


33-5 






Female, England (Bartlett : 


























British Museum) 


42 


16 


9-5 




16-75 


8 


39 


5 


66-75 


34-6 


34 




Female, Berks, measured 


























from dried skin 


49 


16-5 


9 


40 
















267 


Female, Ilminster 


46 


17 




43 






42-5 










285 


Female, N. Wales (the last 


























three in British Museum) 


43 


16 


8 


42 
















272 


Female, Co. Fermanagh 


























(Dublin Museum) . 

Average of females \ 
(approximate) . / 










17-25 


9 


89 


6-0 


67-5 


35 


34-75 




45-1 


14-9 


9 


37-6 


16-5 


8 


39-4 


5-8 


65-6 


33-1 


83 


[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 Mamtal), 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. 

* 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, ibid., 1898, 493. 

2 William Borrer, ibid., 1874, 4127. 

•' Joseph Armitage, Naturalist, 1900, 114, and other references. 
< A. H. Cocks, Zoologist, 1906, 186-187. 



i84 VESPERTILIONID^— 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 

^ Zoologist^ 1889, 241-248. 



NATTERER'S BAT 185 

allowing so near an approach as to be identified while on the 
wing. Its flight when feeding was by no means rapid, but on 
leavinof 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 oft 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.^ 

^ Zoologist, 1883, 294. 2 Ji)id.^ 1898, 493. 

^ Paper read before the Biological and Geological Section of the Cardiff 
Naturalists' Society, March 1905, 5 ; also in MSS. 

R 



i86 VESPERTILIONIDiE— 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 

^ Zoologist^ iQOSj 51-56. ^ Ibid.^ 1889, 382. ^ 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 Vespertilionidce and the Rhinolophidcs 
(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 Rhino lop kidce. In the Horseshoes the short tail is 
carried recurved over the back ; in Pterygistes^ PipistrelluSy 
Plecotus, and some species of Myotis, it is usually carried 
curved beneath the body ; in M. nattereri 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 

^ I.e., Nye talus. 



i88 VESPERTILIONID.E— MYOTIS 

to be constant in the Rhino lop hides, and seldom noticeable in 
the VespertilionidcB, 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, 



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.' 

" I 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 (Geoffrey). 

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 cPHist. Nat., viii., 198. 

•^ As ciliatus, Wiegmann's Archiv fitr Naturgeschichte^ XIX., i., 288-293. 

R 2 



I90 VESPERTILIONIDiE— 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 
A^tgleterreT 

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 M. nattereri, and R. F. Tomes,^ 
although he believed that Geoffrey's type-specimen was a 
M. mystaciuMs, 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 M. 
mystacinuSy with a narrow interfemoral membrane, its posterior 
border ciliated but less thickly so than in M. nattereri. 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 MY sons (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 Beige, 1842, 20. - Zoologist, 1856, 4938-4939. 

^ Ibid., 1887, 161-162, footnote. 



THE MOUSE-EARED BAT 191 

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 
broupfht it alive to Dr Hans Gadow. 

This species has suffered much at the hands of slipshod 
nomenclaturists. It is clearly the Vespertilio 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," Vespertilio murinus of Linnaeus, 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 murinus 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. ?>iyotis hy Bechstein, in Gemeinut' 
zige Naturgeschichte Deutschlands, Qr'c., I., ed. 2, 1154, 1801. 



192 VESPERTILIONID.E— 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 ; and a 
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 Museutn, i ; based 
on Macrotus eitropceus of Leach, the " European Longear," from Devonshire, Eng- 
land, presented by G. Montagu ; named, but without description, hence Plecotus 
has priority. 

18 18. Plecotus, Etienne Geofifroy, Descriptio7i des Mammiferes qui se irouvent en 
Egypte^ ii., 112, 118-119, pi. ii., No. 3 (for date see Sherborn, Proc. Zool. Soc. 
(London), 1897, 287-288 ; based on Poreillard of Daubenton, la barbastelle, and 
an undescribed species from Timor. 

1829. Plecautus, Francois Cuvier, in Dictionnaire de Sciences Naturelles, lix., 415, a 
misprint for Plecotus ; no species mentioned. 

This genus, although small in known species, has a wide 
distribution, mainly in the Palaearctic 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. auritiis. P. puck, which I described 
from Murree, northern India, differs in cranial characters, while 
my P. teneriffcB, from Teneriffe, is larger, having the fore- 
arm measuring about 44 mm. (see Ann. and Mag. Nat. Hist.y 
December 1907, 520-521); G. M. Allen's P. sacrimontis, from 
Mt. Fusi-Yama, Japan, has a long thumb (117 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 CorynorhinuSy 
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 



PLECOTUS 



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. i) 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 
long as the head and body, and has 
the tip slightly exserted. 

The skull (Fig. 8, No. 4, p. loi) 
is weak ; the profile descends gradu- 
ally from the inflated brain - case to 
the depressed, broad, not saddle - 
shaped facial region ; the zygomata 
are flattened ; the cranial crests are 
very slightly developed ; the auditory 
bullae are very large, fully equalling, or, 
perhaps, exceeding, in size those of 
much larger bats, such as Nyctalus 
noctula, Vespertilio serotinus or Myotis 
myosotis. 

There are two less teeth (Fig. 8, 
No. 4, p. loi ; Fig. 19 ; and Fig. 20, 
p. 197) than in Myotis, one pair of upper premolars being absent ; 
two more than in Barbastella, the formula being : — 




Fig. 19. — Diagram of Arrange- 
ment OF Teeth in Genus 
Plecotus. 

(l) Upper and (2) Lower Jaw. 



2-2 



I - I 



pm 



3-3' 



3-3 
3-3 



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 VESPERTILIONID.E— PLECOTUS 

THE LONG-EARED BAT. 

PLECOTUS AURITUS (Linnseus). 

1740. The Double-Eared Bat, Eleazar Albin, A Natural History of Birds, iii., 95, 

pi. ci. 
1758. Vespertilio AURITUS, Carolus Linnaeus, Systema Natura, x., 32 (6) ; xii., 47 (5), 

1766 ; described from Upsala, Sweden ; Bingley ; Donovan ; Jenyns ; Clermont. 
1789. (?) Vespertilio auribus, Gilbert White, Selborne, Letter xi. to Thos. Pennant, 

9th September 1767 ; named, but' without description, from Selborne, England. 
1 8 16. (?) Macrotus EUROPiEUS, W, E. Leach, Systematic Catalogue of the Specimens 

of the Indigenous Mammalia and Birds that are Preserved in the British Museum, 

etc. (London), i ; named, but without description, from Devonshire, England, from 

Montagu's specimens. 
18 1 8. Plecotus AURITUS, Etienne Geoffroy, Description des Mammifires qui se 

trouvent en Egypie, ii., 118, pi. ii., No. 3 ; Fleming ; Bell (ed. i) ; MacGillivray ; 

Blasius ; Fatio ; Dobson ; Blanford ; Flower and Lydekker ; Lydekker ; Thomas ; 

Mehely ; Johnston ; Cabrera ; Millais ; Trouessart (19 10). 

1825. Vespertilio otus, Friedrich Boie, Oken's /$•« (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 Mammalogie, 95, No. 232. 

1827. Plecotus cornutus, auct. et op. cit., 96, No. 234. 

1828. Plecotus brevimanus, Leonard Jenyns, Trans. Linnean Soc. (London), xvi., 
55, pi. i., fig. 2 ; described from an immature specimen from Grunty Fen, Isle of Ely, 
England ; not P. brevimanus of Bonaparte, 1841 ; Bell, eds. i and 2 (figures). 

1829. Plecotus vulgaris, A. G. Desmarest, Faune Frangais, Mammiferes, 
livr. 19, 18 ; pi. 2, livr. 16, fig. 3, described from France. 

1829. Vespertilio brevimanus, J. B. Fischer, Synopsis Mammalium, 118 and 553. 
1832. P. peronii, Isidore Geoffroy, Guerin's Magasin de Zoologie, ii., Classe I., 

No. 2, pi. iii., fig. I ; described from a male and female obtained, locality unknown, 

during the voyage of Peron and Lesueur. 
i860. Plecotus kirschbaumii, Carl Koch, Achter Bericht der Oberhessischen 

Gesellschaft fiir Natur- und Heilkunde, 40 ; described from Oberhessen, Germany. 
1862-63. Plecotus auritus, typus, montanus et brevipes, Carl Koch, fahrbiicher 

des Vereins fiir Naturkunde im Herzogthum (Nassau), 406-408; described from 

Nassau, Germany. 
1873. Plecotus leucoph^eus, N. A. Syevertzoff, Trans. Soc. Nat. Moscow, viii., 2 ; 

translated by Craemer in Ann. Mag. Nat. Hist., July 1876, 42 : named, without 

description, from north-western Turkestan. 

LOreillard'^ of the French, die langohrige Flederniaus 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 ^ 

^ Horned Bat, of Cheshire (Coward and Oldham) ; see also under PiPlSTRELLE, 
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; Barcelo), Specimens from more than one locality have 
formed the basis of specific names, e.g., bonapartii of Gray {Mag. 
Zool. and Bot., ii., 495, 1838) from Sicily; cBgyptiacus of Geoffroy from 
Egypt; 2 cJiristii of Gray {Joe, cit.) and ustus of Heuglin, from Wadi 
Haifa, north Africa ; and homochrous of Hodgson {Joum. 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, Zoologist, 1844, yjT, Wadham), and Scilly (Clark), and almost 
with equal certainty in every county of Wales (see Forrest, Trans. 
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. ^'c'c (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. Hist., 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 Pipistrellus 
pipistrellus. 

In Ireland it is probably found commonly in every county, having 
been known to Alcock and Moffat from all except eleven {Irish 
Naturalist, 1901, 250-1), and there can be but little doubt that the 

^ Specimens seen. 

^ Or cEgyptius, as frequently cited, but original description not found. 



196 VESPERTILIONID^— 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 (Harrington, Migration^ 284), and by 
Sheridan on Achill, Co. Mayo (Dublin Museum). 

Distribution in time : — A portion of a humerus from layer 1 1 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. i) 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, Field, 2 1 St March 1874, 271 ; Borrer, 
Zoologist, 1874,4128 (two); V>'d.x\n%, Journ. 
cit.y 1898, 261. I have also records of 
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). Millais has seen a true albino 
and a cream-coloured variety from Nor- 
folk, and Eagle Clarke {Naturalist, iZg4.,62>) 
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 Fig. 2o.-Front View of Incisors 
represent a hitherto undescribed sub- 
species. Those from Africa and others 
from sandy regions are said to be of pallid coloration (? P. christii 
of Gray), while Hodgson's P. homocJirous 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. 

Skxill: — Greatest length, 16-5; basal length in middle line, 12; 
palatal length in middle line, 7 ; from posterior border of w^ 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. 



AND Canines of Plecotus auntus 
(enlarged and diagrammatic). 



198 



VESPERTILIONID^— PLECOTUS 



DIMENSIONS IN MILLIMETRES:— 



Males. 




•a 

.s 

a 
1 




.a 

CO m 

s 


'3 


n 

5 


11 


03 

O 




4^ 

•a 
s 

60 

s 




> 
t 

a 

1 


® 13 
1° 

o 


1. 
1 


Male, Grimsby, Lincoln- 




























shire, 10th Sept. 1890 




























(W. Eagle Clarke) . 


47 


35-5 


18-25 


43-5 






38 










259* 




Male, Manifold Valley, 




























North Staffordshire, 




























16th March 1907 (C. 




























Oldham) . 


47 


34-5 


16 


35 


19 


8-5 


36 


7 


63 


31 


30 


247 




Male, Tring, Herts, 




























dried skin at British 




























Museum . 


51 


36 




45 


17 


9 


37-75 


5-5 


62 


31-5 


30 






Male, London, do. 


51 


38 




48 


17 


9 


37-5 


6 


63 


31-5 


31 






Male, Montrose, do. . 


45 


36 




45 


16-5 


8-5 


37-75 


7 


63 


31 


30-5 






Male (dried skin), Co. 




























Louth, 3rd August 




























1893(Dublin Museum) 










18 


9 


38 


7-25 


64 


32 


30-25 






Male (do.), do. 4th 




























August 1893 (Dublin 




























Museum) . 










17 


9 


37 


7 


62-75 


31-75 


80 






Male (do.) do. 5th 




























August 1893 (Dublin 




























Museum) . 








33-5 


17 


9 


39 


7-5 


62-5 


32 


81 






Male, Achill Island (do.) 










16-25 


8-5 


35 


6 


57-75 


29-25 


27-75 






Male (In the flesh), Co. 




























Fermanagh (do.) 

Average of males \ 
(approximate) / 










18 


9 


39 


7 


62-5 


32 


29-5 






48-2 


36 




41-6 


17-3 


8-8 


37-5 


6-6 


62-2 


81-3 


30 






Male, young, Devon 


50 


37 




44 


17 


9 
















Male (dried skin), Co. 




























Louth (hardly ma- 




























ture) .... 










18 


9 


38-5 


7 


61 


31 


29 






Male, young, Co. An- 




























trim (Dublin Museum , 




























in alcohol) . 










11 


8 


32-5 


7-5 


27-5 


14-5 


13 






Females. 



Female (in flesh), Mani- 
fold Valley, Stafford- 
shire, 16th March 1907 
(0. Oldham) 

Female, Devon, dried 
skin, at Brit. Museum 

Female, Essex, do. . 

Female, Herts, do. 

Female (1), Montrose, 
do 

Female (2), do. do. . 

Female (3), do. do. . 

Female, North Ber- 
wick, 16th Sept. 1894 
(W. Eagle Clarke) . 

Female (dried skin), Co. 
Louth, 5th July 1893 . 

Female (do.) do. 5th 
August 1893 (Dublin 
Maseum) . 

Female (do.) Co. Ar- 
magh, 4th September 
1893 (Dublin Museum) 

Female (from spirit), 
Co. Louth (Dublin 
Museum) . 

Female do. 

Female do. 

Female, Co. Meath (do.) 

Female, Co. Dublin (do.) 

Female, Co. Louth (do.) 

Average of females \ 
(approximate) / 



48 


35 


42 


36 


49 


35 


46 


34 


45 




44 





35-5 



17 



15 



45-7 



40-5 

42 
45 
42 

46 
42 
40 



37 



17-5 

18 

18-5 

18 

18-5 

17-5 



19-75 

19 

18-5 

19 

20 

19 



9-5 
8-5 



9-5 



18-3 



40-5 
40 



(!)35-5 



39 



7 

6-5 

6-6 

6-5 

6 

7 



7-5 



61-75 

65 

65 

68 
67 
63 



63 



40-25 



39 
40 
40 



7-5 
7-5 
8-5 



7-1 



Average total expanse of eight individuals, 



63 



67-6 

64 

64-5 

66-5 

66-6 

65-6 



66 



33 



32 



32 



33 

31 

82 

81-5 

83 

82 



82 



29-75 

32 

31-5 

82-5 
31-5 
29-5 



30-5 



31-5 



30 



31-6 

30-25 

80 

81 

32-25 

81 



260 



5 grms, 



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 

^ Irish Naturalist, 1901, 241-251. ^ See Introduction, pages 40, &c. 

^ The folding is well described by A. H. Cocks, Buckingham. 



200 VESPERTILIONID^— 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, 

^ Zoologist^ 1853, 3937* " Ibid.^ 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 

* Zoologist^ 1897, 231. - Ibid.^ i890> 7i- 

Fieldy 23rd July 1887, 149. * Lord Lilford, Zoologist, 1887, 66. 



° Zoologist, 1889,382. ^ Spilosorna lubricipeda. 

'' TriphcBna. ^ Plusia gamma. '•* Zoologist, 1883, 294. 



S 



202 VESPERTILIONID^— 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," Irish Naturalist^ 1906, 278. 
^ Ibid.^ 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 
in sight. . . . 

" 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. 
j> 5tu „ „ 3" " )' 

,, 6th „ „ 36 „ „ 

j> 7tn 5j J, 33 )j » 

„ 8th „ „ 34 „ „ 

)j loth „ „ 33 „ „ 

5j iith. ,j „ 37 „ „ 

„ 1 2th „ „ 34 „ „ 

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 mornlnor twilio-ht.^ 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 

^ I.e., Leisler's Bat, 2 ^/^/^^ 23rd July 1887, 149, 

^ Gordon DalgHesh 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 i8th 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 in situ. The sleeping-place was a hole in a Beech-tree, 
5|- 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 nth 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. 



PLATE XV. 




Long-eared Bats, (g 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 
2ist 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. 

'* It returned for a few days about March 17th, during the 
observer's absence from the locality, and deserted again on the 
25th. On April loth 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 loth, nth, and 
1 2th — it did not fly at all. On the loth it was visited with a 
lantern at 7.35, 9.5, and 11.5 p.m. ; on the nth, at 8.45 p.m., 
and on the 12th, at 8 and 10.30 p.m. ; and on all these occasions 
was seen sound asleep. 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 
loth 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 

^ See also Mem. and Proc. Manchester Literary and Philosoph. Soc, 49, ii., 9, 1-4, 
31st March 1905. 



2o6 VESPERTILIONID^— 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 20th 
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, 

' Compare Goldsmith's " Lazy bats in drowsy clusters cling." - In 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 nth 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, 

^ R. C. R. Jordan, Zoologist^ 1843, 75 ; Alcock and Moffat, Irish Naturalist^ 1901, 
248 (nth July). 



2o8 VESPERTILIONID.E— 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 fiy 
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 

' Original not seen : quoted by Rollinat and Trouessart, vide supra, p. 36. 
^ J. Armitage, Naturalist, Feb. 1905, 39. 



HISTORY OF BRITISH BIRDS— continued. 

histories of very many others, which were formerly little known, 
have 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 towards a more satis- 
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 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 
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 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- 
known bird artist. 



PATERNOSTER ROW, LONDON, E.C. 



In the Press, to be p ublished 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 Illustrations and Maps 



WITH 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 
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. 

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 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), 
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 oi 
Bird-Migration, but should occupy a place essentially its own in 
ornithological literature. 

OXJRNEY & JACKSON 

PATERNOSTER ROW, LONDON, E.G. 



Oliver and Boyd, Printers, Edinburgb. 



PLECOTUS 209 

feeding Noctiice ... 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 lenofth. 

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 TcEniocampa, 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, pupae, 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 

> Zoologist, 1899, 472. - Calliphora eryt/irocephala. ^ Zoologist, 1889, 245. 

^ Gonoptera libatrix and Scotosia dubitatd. ^ Naturalist, 1905, 39. 

^ F. W. L. Ross, Zoologist, 1845, 11 58; Couch, Journ. cit., 1853, 3942; Lilford, 
Journ. cit., 1887,66; Oldham, yo«^r«. cit., 1890, 349; R. Newstead in Coward and 
Oldham, 1895, 166; W. Evans ; Forrest in MS., etc., etc. 

T 



2IO VESPERTILIONID.E— 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. 

1 82 1. BARBASTELLA, J. E. Gray, London Medical Repository^ xv., 300; based on 

Vespertilio barbastellus of Schreber. 
1829. Barbastellus, Jakob Kaup, Systetn der Europdischen Thierwelt, i., 96 ; based 

on Vespertilio barbastellus of Schreber. 
1839. Synotus, a. Graf von Keyserling and J. H. Blasius, Wiegmann's Archiv fiir 

Naturgeschichte, i., 305 ; based on Vespertilio barbastellus of Schreber. 

Synonymy: — The earliest generic name known to have been 
applied to the Barbastelle is certainly Barbastella of Gray, a 
name revived by Mr Miller in 1897 i^^^^^- and Mag. Nat. Hist., 
Oct. 1897, 384-385). It clearly antedates both i5^r^^i^/^/6^5 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, B. barbastellus. 

The distribution is imperfectly known, but extends at least 
from Skandinavia to Abyssinia, Egypt (Heuglin, Reise in Nord- 
ost A/rika, ii., 30, 1877), and Arabia, where the local form was 
named leucomelas by Riippell {Atlas zur de^^- Reise ini n'drd- 
lichen A/rika, ^2)^ P^- xxviii. B, 1826); and from Great Britain 
to Assam, where the representative is Hodgson's darjeling- 
ensis, a common mountain bat of the Himalayas, ascending 

^ R. M. Barrington, The Migration of Birds, 284 ; R. H. Porter, London, 1900. 

^ Since the above article was written, Oldfield Thomas has described P. wardi, 
a large pale-coloured form, from Leh, Ladak, Kashmir {Ann. and Mag. Nat. Hist., 
19x1, 209) ; and P. ariel, allied to P. wardi but of much darker colour, from Ta-tsien- 
lu, Szechwan, Western China {Abst. Zool. Soc. (London), 7th February 191 1, 3). 



BARBASTELLA 



211 




to at least 8000 feet (Blanford). It is said to have a larger ear 
and longer forearm (44.5 mm.) than barbastellus. 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. 

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- 
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. loi), as 
exemplified by B. barbastellus, is weak, 
with prominent rounded brain-case ; 
broad, somewhat concave facial region ; 
weak, flattened zygomata ; weak or no cranial crests ; and 
moderately developed auditory bullae. 

There are thirty-four teeth (Figs. 21 and 22) arranged as — 

• 2 — 2 I — I . 2 — 2 3 — 3 

I , c , pni , 111 - — ^ — 

l-l i-i 2-2 3-3 




Fig. 21. — Diagram of Ar- 
rangement OF Teeth in 

genus Barbastella. 
(i) Upper and (2) Lower Jaw. 



= 34- 



The upper incisors are oblique, the outer pair small, the inner 
large and bifid ; the upper canine is provided with small anterior 



212 VESPERTILIONID^— 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 (Schieber). 

1760. La BARBASTELLE, L. J. M. Daubenton in E. L, le Clerc, Comte de Buffon's 

Histoirc Naiurclle, viii., 119 and 130-131, 135-137, pi. xix., fig. 2 ; described from 

France. 
1774. Vespertilio BARBASTELLUS, J. C. D. von Schreber, Die Siiiigthiere^ i., p!. Iv., 

168, evidently naming Daubenton's La Barbastelle ; James Sowerby, British 

Miscellany, pi. v., g, 1804-6 ; George Montagu, Trans. Linnean Soc, London, ix., 

171, 1808; Bingley ; Pennant; Jenyns ; Clermont. 
1785. Vespertilio barbastella, P. Boddaert, Elenchus Animaliiwi, i., 69. 
1821. Barbastella barbastellus, J. E. Gray, London Medical Repository, xv., 300 ; 

Miller ; Thomas ; CoUett ; Johnston ; Millais ; Trouessart (1910). 
1829. BARBASTELLUS BARBASTELLUS, Jakob Kaup, System der Eiiropiiischen Thier- 

•welt, i., 96. 

1836. Plecotus BARBASTELLUS, Georges Cuvier, Le Rcgne Animal, i., 74 ; Fleming. 

1837. BARBASTELLUS DAUBENTONll, Thomas Bell, British Quadrupeds (ed. i), 63 ; 
(ed. 2), 81, 1874, renaming Daubenton's La Barbastelle ; MacGillivray. 

1838. BARBASTELLUS COMMUNIS, J. E. Gray, Mag. Zool. and Bot., ii., 495 ; evidently 
renaming V. barbastellus of Schreber. 

1839. Synotus BARBASTELLUS, A. Graf von Keyserling and J. H. Blasius, Wieg- 
mann's Archiv fiir Naturgeschichte, i., 305 ; Blasius ; Fatio ; Dobson ; Flower and 
Lydekker ; Lydekker ; Cabrera. 

1900. BARBASTELLA BARBASTELLA, Mehely Lajos, Monographia Chiropterorum 
Hungariae, 131 and 326, pi. v. 

Barbastelle from the French barbastelle and Italian barbastella, from 
the Latin barba, i.e., 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 B. barba- 
stelhis is preferably a mountain bat ; as is B. darjelhigensis, 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. 



THE BARBASTELLE 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 
Jouryial, ii., no, 1826) on the ground that the individual in the British 
Museum marked by him barbastellus, is undoubtedly referable to Myotis 
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, Jonrn. cit., 1869, 
1768). Thus between Cornwall and Kent the sprinkling of occurrences 
perhaps indicates a continuous distribution. There are several records 
for Surrey (Mitford,/t'/^r//. cit., i860, 6953 ; Millais, 42 ; Thorburn, Field, 
19th July, 1902, 142; Dalgliesh, Zoologist, 1907, 299; Bucknill and 
Murray); and Sussex (Borrer, Zoologist, 1874, 4128-29; Brazenor, 
Joiirn.cit., 1887, 152 ; Millais, 42); for Hampshire, one or two (Hart and 
Kelsall) ; for Wiltshire, one at Salisbury (Blackmore, Zoologist, 1869, 
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 
{Jouni. cit., 1869, 4128-29), and by Dale as "not common" {Journ. cit., 
1887, 234). North of the Thames Valley it is known from Essex, 
where Doubleday found it not uncommon in Epping Forest {Journ. cit., 
1843, 6"]), 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 {Journ. cit., 1874,4128); in Hert- 
ford, Oldham found one on Berkhampstead Common {Journ. 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 {in lit., also see Jourti. cit., 1892, 329; Witchell, yi?«r;A cit., 
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. cit., 1857, 5420; see also for Suffolk, ^O'^q, Journ. cit., 1891, 347) ; 
and in Norfolk Southwell describes it as not rare and generally distributed. 

T 2 



214 VESPERTILIONID^— BARBASTELLA 

Jenyns knew of its occurrence in Cambridge ^ and Northampton ; Lord 
Lilford in the latter county {Journ. cit,. 1894, 187), and in Hunting- 
don {Journ. ciL, 1894, 395); Steele Elliott in Bedford; and Montagu 
Browne in Leicester (y(9?/r«. aV., 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,/(??^r/^. cit., 1857, 5590, see also Tomes). North 
of the Wash, Caton Haigh's description {Journ. cit., 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, Zoologist, 1904, 262; 
Trans. Caradoc and Severn Valley Field Club, iv., i, 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 {^Naturalist, 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. 

^ 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. 



2l6 



VESPERTILIONID/E— BARBASTELLA 



The nearly full-grown young are said to differ from the adults in 
having the underside darker. 

The sktill (Fig. 8, No, 5, p. loi) 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 
taken on 4th September {Zoologist^ 1894, 
187 and 395). 

As regards size, the forearm in eight 
British specimens measured by me aver- 
ages 1-5 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 : — Foot, 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, io-6; 
palatal length in middle line, 5 ; from posterior border of m^ to anterior 
border of canine, 5 ; greatest breadth at zygoma, J-y ; 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 Barbastellc 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. 




Fig. 22. — Front View of 1n- 
cisoKS AND Canines of Barha- 

stella barhastellus (enlarged and 
diagrammatic). 



PLATE XVII. 





f 




Wings (h natural size and diagrammatic) OF 

I. Pkcolus aurilus. 2. Barbaslella harhastellus. 

3. Rlii)wIophus hipposideros. 



PLATE XVIII. 







Heads (natural size) OF 

id 3. Rhinolophus hipposideros (front, three-quarter, and side views). 
4. R hinoloplms ferrum-equinum. 



THE BARBASTELLE 



217 



DIMENSIONS IN MILLIMETRES: 







■a 
§ 

1 

a 




I 


bD 



3 


.a 

'% ^ 

"Si 

-^ 


i 


5=' 

"o 

s 


a 

H 


1 
bo 



1-; 


s 


> 


a 
a . 

a 
£ 


Male, Arrow Lodge, War- 
wickshire, 7th October 
1S4S(R. F.Tomes's col- 
lection), measured from 
dried skin at British 
Museum 

Male, Ouudle, Northants, 
5th April 1900 (Lord 
Lilford), do. . 

Male, Milton, Hants, 
7th April 1900 (Rev. J. 
E. Kelsall), do. . 

Male, Llanelwedd, Rad- 
norshire, Cth June 
1904 (H. E. Forrest), 
do 

Male (received from E. 
A. Wilson), 5th Oct. 
1906, measured in 
flesh .... 

Male, Cheshire (G. E. 
Dobson), measured 
from spirit in British 
Museum 

Average of males ^ 
(approximate) . / 


50* 
5^* 

46 

45 


16* 

1-2 -7 
14-5 


6* 

7-5 
6-5 


42 

48* 

44* 

43 
40 


18 
IS 
lS-5 

19 

19-5 

19 


7 
6-5 

6-5 

7-5 

6 


37 
38 
3« 

38 

38 

39 


4 
4 
3-25 

4-5 

5 

5 


65 
approx. 

08 
06 

68-5 

63-5 

67-5 


34 
35 

34 

35-5 
35-5 
35 


32 
33 

31-5 

33 5 
34 

31 


200 

254* 


4S 






43-4 


18-6 


6-7 


37-6 


4-3 


00-4 


34-8 


32-5 




Female, Elton, Hunts, 
4th Sept. 1894 (Lord 
Lilford), measured from 
dried skin . 

Female, Epping Forest 
(H. Doubleday), mea- 
sured from alcohol 

Female, Herts, received 
in flesh from C. Old- 
ham, 16th Sept. 1908 . 


GO* 
44 
51 


10 
13-5 


S 
9 


43 
54 


18 
19 
20 


6-5 

6 

7 


36 
37 
41 


4 
4 
4 


03-5 

05 

07 


35 
32 
37 


31 
30 
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 



2i8 VESPERTILIONID^— 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 

' See A. Newton, Zoologist, 1857, 5421 ; Lord Lilford, ibid., 1894, 187. 
- See H. P. Blackmore, ibid., 1869, 1558 ; C. W. Brazenor, ibid., 1887, 152 ; A. H. 
Cocks, Jenyns, etc. 



THE 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 I 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 

> In Bell. 



220 VESPERTILIONID^— BARBASTELLA 

of repose during the day, but durinor 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 in a 
ruined monastery in Arragon, at the foot of the Pyrenees, . . . 
and in smaller numbers in a similar locality at Botes, 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. 

^ Zoologist, 1857, 5590. - Ibid., 1887, 66-67. 

"^ 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, Naturalist, 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. As a 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 VESPERTILIONID^— 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. 

^ See H. P. Blackmore, Zoologist, 1869, 1558. 

- R. Mitford, ibid., i860, 6953 ; Lilford, ibid., 1894, 395 ; Owen, per Forrest. 

^ For use of Nycteris (replacing Lasiurus), see Miller, Proc. Biol. Soc, Wash- 
ington, 17th April 1909, 90. 

■* Under the name Vesperiilio 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 BAT 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 

1 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. 

^ Miller, North American Fauna, No. 13, 112, 1897. 
* .Science, New Series, v., 542-543, April 2, 1897. 



224 VESPERTILIONID^— 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.] 

• Nycteris semota (Allen). - N. borealis (Miiller). 

^ Lasionycteris nocHvagans (Le Conte). 



RHINOLOPHID^. 

HORSESHOE-NOSED BATS. 

Characters :— These bats have a conspicuous nose-leaf; no 
tragus ; a short tail and moderately developed interfemoral 
membrane ; premaxillae 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, Tableau elt'mentaire de PHistoire Natiir- 
elle des Animaux, 105. 

1799. RHINOLOPHUS, Le C. Lacepede, Tableaux . . . des Mamtnlfires, etc., 15, No. 
73 ; based on Rhinolophus ferru7n-equinuin. 

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), i ; based, without description, on Phyllorhifia minuta of Leach = Vespertilio 

mi7iutus 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 Pittoresque d'Histoire Naturelle, 

iv., 2, 617, quoting from Cuvier and Geoffroy, Magazine Encyclopedique, 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., Rhifiolophus philippinensis ; preoccupied 

by Phyllotis of Waterhouse, 1837, a genus oi Muridce. 
1866, Speoriferra, J. E. Gray, Proc. Zool. Soc. (London), 13th Feb., 82 ; based on 

Speoriferra vulgaris, i.e., Rhinolophus vulgaris of Horsfield. 
1866. CCELOPHYLLUS, W. Peters, Proc. Zool. Soc. (London), 22nd Nov., 427 ; based 

on Rhinolophus ccelophyllus of Peters. 
1887. PSEUDORHINOLOPHUS, Max Schlosser, Bcitrdgc stir Paliiontologie Osterreich- 
226 U 



226 



RHINOLOPHID^— RHINOLOPHUS 



Ungarns, etc., VI., i. and ii., 55, 61, pi. ii., fig. 42 ; based on Rhinolophus antiquus 
of Filhol and other extinct species. 
1 90 1. EURYALE, Paul Matschie, Sitzungs-Berichte der Gesellschaft naturforschender 
Freunde zu Berlin^ 225 ; based on E. mehelyi of Matschie. 

The genus is the only one of its family. There are two 
British species, R. ferruvi-eqtiimtm and R. hipposidei'os. 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 R. vidhelyi of Matschie and the 
Spanish R. carpetamts of Cabrera. Peters's R. blasii of south- 
eastern Europe has peculiar nose-leaf and lower incisors. 

Distribution: — The seventy-one species and numerous sub- 
species recognised by Andersen (Ann. and Mag. Nat. Hist., 
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 (i?. hipposideros) or 
southern Korea (i?. ferrum- 
eqtiinum nippon and R, cor- 
nutus). Andersen refers exist- 
ing species to six "types," all 
of which he believes to have 
had their origin in the Oriental 
Region. 

The genus is characterised 
by moderate or small size, 
highly developed nose - leaf 
with triangular posterior por- 
tion, and deeply notched ear 
with well - developed anti- 
tragus. 

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 




%, 






Fig. 23.— Diagram of Nose-leaf of 
Rhinolophus. 

a, sella. d, division between the 

3, lancet. nostrils. 

c, c, eyes. e, e, horseshoe. 



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 mammae 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- 
tion, and are not in communication 
with any gland : they are attached by 
fibrous tissue to the pubis (see Rol- 
linat and Trouessart, Mdm. Soc. Zool. 
de France, x., 1897, 124). 

The teeth (Figs. 25 and 26) are 
thirty-two, arranged as — 



2-2' 



I - r 



}>-i 




The lower incisors are tricuspid. 
The upper incisors, although minute, 
are always present in the always 
ossified premaxillai. The anterior fig. 24.— front View of In- 
upper premolar is minute, and often cisoks and Canines of Riuno. 

i^ i- ^ /o///z« (enlarged and diagrammatic). 

crowded out of the tooth-lme 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 RHINOLOPHID^— RHINOLOPHUS 



THE GREATER HORSESHOE BAT. 

RHINOLOPHUS FERRUM-EQUINUM (Schreber). 
RHIXOLOPHUS 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, pi. xx., figs, i and 2 ; also, Mem. de 

VAcad. Roy. des Sd., :^77, 382, pi. 15, fig. 4, 1759, published 1765 ; described from 

France. 
1774. Vespertilio FERRUM-EQUINUM (in part), J. C. D. von. Schreber, Die 

Sdiigthiere, i., pi. Ixii., upper of two figs., 174, 190; evidently naming Daubenton's 

Le Fer-a-cheval; Montagu, Trafisactiofts Linnean Soc. (London), ix., 162, 1808 ; 

Bingley ; Donovan ; Clermont. 
1776. Vespertilio equinus (in part), P. L. S. Miiller, Naiursystems Supplemettis 

und Register Band., 20, renaming Schreber's Vespertilio ferruvi-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, Elenchiis Animalium, i., 71, 

renaming Schreber's V. ferrum-eqidnwn. 
1788. Vespertilio ferrum-equinum, a major, J. F. Gmelin, Systema Naturae., i., 50. 
1792. Vespertilio ferrum equinum major, Robert Kerr, Animal Kingdom., ()C) 

(not V. molossus major of Kerr, op. cit., 97) ; described from France. 

1797. NOCTILIO ferrum-equinum, J. M. Bechstein, Der Zoologe, i, i.-viii., 62. 

1798. Vespertilio hippocrepis (in part), F. von P. Schrank, Fauna Boica, i., 64. 
1803. RHINOLOPHUS MAJOR, Etienne Geofifroy, Catalogue Mammifcres Mus. d'Hist. 

Nat. (Paris), 56 ; described from Burgundy. 
1807. Vespertilio ferrum equinum majus, Lazare ST^aWa-nzSini, Rapports de Pair 

avec les etres organises., etc., edited by Jean Senebier, ii., 77 (Geneva) ; described 

from Nonantola, near Modena, or from Scandiano, Italy. 
1813. RHINOLOPHUS UNi-HASTATUS, Isidore Geoffroy, Ann. du Mus. d^Bist. Nat., 

XX., 257 ; described from Europe. 
18 1 7. RHINOLOPHUS FERRUM-EQUINUM, W. E. Leach, Zoological Miscellany, iii., 

2 ; Harting, Zoologist, 1887, 2, pi. i. (G. E. Lodge), et pluritnorum auctorwn. 
1829. RHINOLOPHUS UNIFER, Jakop Kaup, System der Europdische ft. Thierwelt, 

\., 104 ; without description, hence a ?totnen Jiudum. 
1862-63. RHINOLOPHUS FERRUM-EQUINUM, var. GERMANICUS et van ITALICUS, Carl 

Koch, Jahrbiicher des Vereins fiir Naturkunde im 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), i6th May 1905, 113 

footnote). 

1872. RHINOLOPHUS LIBANOTICUS, CONCHIFER, et RUFESCENS, '^ Ehrbg. et Lichst. 

Mpts," W. Peters, Monatsberichte der Koniglich Preussischen Acad, der Wissen- 

schaften (Berlin), 1 871, 310 ; without description, hence a nomen nudum. 
1885. RHINOLOPHUS UNIHASTATUS var. HOMORODALMASIENSIS, Daday Jeno, Orvos 

— Termeszettudomdnyi Ertesito, etc., x. (3), 274 ; described from Homorod — 

Almas, Hungary, in error (see Mehely, Monographia Chiropterorum Hungariae, 

120-122 and 323, 1900). 
1887. RHINOLOPHUS FERRUM EQUINUM, var. HOMORODENSIS, Daday Jeno, Erte- 

kezt'sek A Tertneszettudomdnyok Korcbol, Budapest, xvi., No. 7, 13, pi., 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-a-cJieval of the French ; die grosse Hufeisemiase 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. hipposidej'os.) 

Distribution: — R. fernun-equinnni 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. pipistrellus, the commonest 
bat of Normandy (de Kerville, Naturaliste, 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, Mem. Soc. Zool. de 
France, x., 1897, 1 14). 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, British 
Zoology, ed. iv., 129, pi. xiv., 1776), and later was encountered by Mon- 
tagu in considerable numbers in company with R. hipposideros, 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 {Zoologist, 1884, 483) 
states that one was shot on the Berkshire side of the river at Oxford 
(lat. 51°, 46') about 1875 ; ^5 miles further south Noble took two in 
his cavern in the same county, near Henley-on-Thames, on 14th 
March 1909 (see Cocks, Journ. cit., 1909, 154); the first-mentioned 

^ Not, as stated erroneously, at Dartmouth, Devon (see Rowe). 

U 2 



230 RHINOLOPHID^— 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 {vide 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, Zoologist, 1906, 69) and Taunton 
(Millais) ; in Wiltshire, Great Cheverell {Joiirn. cit., 1910, 307) ; in 
Surrey, Godstone (rare, Bucknill and Murray) ; in Kent, Rochester 
Cathedral and Margate (Bell), Maidstone and Dover (Millais), and 
Canterbury (abundant, Borr er, /ourn. cit., 1874, 4129); in Hampshire, 
Portsmouth (Borrer, loe. cit.), and Christchurch (Trevor-Battye and 
Lascelles) ; in Wight, Sandown, Farringford, Bonchurch, and the 
Underclifif (the commonest large bat of the latter, see More, Bury, 
Millais, Wadham) ; in Dorset, Tomson Manor (Salter, Zoologist, 1865, 
9835); in Devon, Teignmouth (Jordan, /i^^/r;/. cit., 1843, 75), Torquay 
(De Hiigel, Journ. cit., 1869, 1768), Hooe, Plymouth, and Plympton 
(Rowe) ; in Cornwall, several localities (see Clark) ; a Yorkshire record 
in tho: Jou7'n. 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 {Field, ist 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 Nat. Hist. Review, vi., 383, 1859). 

The almost entirely south-western distribution oi R.ferriini-equinuni 
(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. 

Distritoution 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, hycnena, and rhinoceros found with them, 
while others are quite recent (see Owen, British Fossil Mammals, 
etc., 15-16). 

Breeding season, and number of yoimg: — 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 {Coviptes rendus Soc. 
Biol. (Paris), 26th January 1895, 53-54. and 6th July 1895, 534-536; also 
Mem. Soc. Zool. de France, x., 1897, ^ 4-134) have been strangely over- 
looked, find that in France the latter number is never exceeded. 

As in the Vespertilionidcz, copulation takes place in the autumn (but 
see under R. 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 {DxivdA, Jonru. de rAnat. et de la Physiol., 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 RHINOLOPHID.-E— 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 
Vavancenient des Sciences, x. ; Coinpte 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 Sac. Phil, de Paris (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-Sequard 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 

' L. Camus and E. Gley, Comptes reiidus Soc. Biol. (Paris), i8th July 1896, 
787. 



THE GREATER HORSESHOE BAT 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 : — 

r. Metacarpal Metacarpal Metacarpal 

Forearm. jjj '^ jy^ y 

1000 644 724 743 

The false nipples are very long, transversely wrinkled, and flattened ; 
their colour is yellowish or brownish. 



234 RHINOLOPHID.^— 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. i) 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 as a quarter. The auditory bullae 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, 
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 nippon of Japan and south China, 
in which the size is moderate, the horseshoe 
broad, and the teeth rather small; Hodgson's 
iragatus, of the Himalayas, with large size 
and broad horseshoe ; and Andersen's 
I'eguhts 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 
Iberian Peninsula, to Transcaspia and the 
Euphrates Valley, and has the forearm 
usually 57 mm. and upwards. Cabrera's 
ohsairus, 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 





Fig. 25. — Diagram of Arrange- 
ment OF Teeth in Rhinolophus 
ferriim-equimvn . 
(l) Upper and (2) Lower Jaw. 



236 



RHINOLOPHID/E— RHINOLOPHUS 



undoubtedly a shorter forearm than the type, although with a skull of 
about equal size. It may be known as R. ferrum-equinuin insulanus 
{Ann. Mag. Nat. Hist., March, 1910, 292). Its forearm is even shorter 
than that of obscurns, with which form it cannot be phylogenetically 
identical, even if apparently indistinguishable. 



DIMENSIONS IN MILLIMETRES 







.a 

60 


+3 "W 






o 




^' 








o 




>a 


□ 


■^ C8 






.£3 




rt 


+j 




> 






■a 
o 


O 


§£.• 




to 




S 

1 

* 


"3 


!§? 


""* 




3 . 




c 
ca 

w 


1 


O H 


'5 


o 


S 


S 


•5 


"3 

u 

a 
a 

0) 


1 
1 


o 


Male, Cheddar, Somer- 


























set (T. A. Coward), 


























6th Jan. 1907 . 


66 


23 


15x8-75 


33 


24-5 


10 


54-5 


6 


85 


36-75 


39 




Male, do. . 


61 


22 


14-25 X 8-5 


32-5 


25 


11 


53-5 


6 


82 


34 


38-5 




Male, do., 7th Jan. 


























1907 .... 


67 


22 


15x8 


33 


24 


12-75 


55 


6 


S7-5 


36 


38 


330 


Male, do., a little dried 


61 


21-5 


14x8 


34 


24 


11 -.5 


51 


6 


75 


34 


38-5 


312 


Male, do. 


65 


22 


15x8 


33 


25 


11 


54-5 


6 


82 


38 


41 


342 


Male, do. . 


65 


23 


15x8 


36 


25 


11 


54-5 


6 


87 


36 


40 


342 


Male, do. 


61 


21 


15x8 


30 


23-5 


11-5 


53-5 


7 


85 


36 


39 


342 


Male, do. . 


60 


22 


15x8 


32 


25 


11-5 


51 


6 


82 


33 


36 


342 


Male, do. . 


65 


23 


15X8 


33 


25 


11 


53 


6 


86 


35 


40-25 


342 


Male, do, . 


61 


21 


14x8 


33 


25 


11 


54 


6 


83 


36 


39 


330 


Male, Somerset (W. 


























Eagle Clarke) . 

Average of males \ 
(approximate) / 


63-5 


22 




33 






55-5 










330 


63-2 


22 


14-7x8-1 


32 


24-5 


11-2 


53-6 


6-1 


83-4 


35-4 


38-9 


334-6 


Female, Cheddar, Som- 


























erset (T. A. Coward), 


























5th Jan. 1907 . 


64 


22 


14-5x8 


33 


24-5 


10-5 


55 


6 


84-5 


35-5 


39-5 




Female, do., 7th Jan. 


























1907 .... 


69 


22 


13x8 


33 


26 


13 


54 


7 


88 


37 


42 


340 


Female, do. . 


63 


22 


15x8 


34 


25 


11-5 


54 


7-5 


87-5 


37-5 


41 


355 


Female, do. . 


68 


21-5 


14x8 


32 


25 


13 


54-5 


7 


84 


37 


41 


350 


Female, do. . 

Average of females ■) 


65 


23 


14x8 


34 


26 


11 


55 


7 


85 


34 


39 


355 


65-8 


22-1 


14-1x8 


33-2 


25-3 


11-8 


54-5 


6-9 


85-8 


36-2 


40-5 


350 


(approximate) J 



























Andersen gives forearm of thirteen specimens of both sexes as averaging 55-4, with extremes 

of 58 and 63-8. 



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, in inches : 
—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 i-ii 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 11 fi to 



THE GREATER HORSESHOE BAT 237 

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.). 

Distingmshing characters: — R. ferriivi-cqiiinuvL 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 RHINOLOPHID/E— 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 tiying 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, form a 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 
a man'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 qui vive. 

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 manoeuvre 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 Mein. and Proc. Manchester Lit. and Philosoph. Soc, xlix., 2, i-i i, pi. 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 RHINOLOPHID^— 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, op. cit. st4pra, the 
present account being adapted from these two writers. 

- 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 meetins: 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 either side, while the finders 
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 aflect 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 Hysena, 
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.^ 

• A fact observed by J. Couch before 1853 {Zoologist^ 1853, 3940- 
2 See Proc. Zool. Soc. (London), ist August 1907, 312-324; also Mem. and P roc. 
Manchester Lit. and Philosop]i. Soc, 52, xi., 1-12, pi. and figs. 1-4, 21st April 1908. 

X 



242 RHINOLOPHID^— 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 Mem. Soc. Zool. de France^ 1897, x., 1 14 and 121. 



THE GREATER HORSESHOE BAT 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 1 6th January. Detailed records for the succeeding months 
are not yet available, but Couch's account of a Greater 

' Zoologist^ 1907? 288-294. 



244 RHIN0L0PHID.41— RHINOLOPHUS 

Horseshoe taken on the wing in Cornwall^ on nth 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 Covi^ard 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 
tiuttering 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 debris 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. - Ibid., 1887, 262. 

■^ The following insects are also mentioned by Coward as forming part of the 
diet : — the large beetles Melolotttha vulgaris of Fabricius, Geotrupes spiniger of 
Marsh, and G. stercorarius of Linnseus ; a staphylinid beetle of the genus Quedius ; 
the flightless beetle Nebria brevicollis of F'abricius, and a Pterostichus ; a geode- 
phagous beetle, perhaps of the genus Aviara, and, perhaps, a Dytiscusj 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. typhoeiis 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 LinniEus, Gonoptera libatrix of Linnaeus, Triphccna orbona of 
Fabricius, T. promiba of Linnteus, and Xylophasia polyodon of LinricCus. Several 
Diptera, including Miiscidce apparently of the genus Liiciliaj a wasp-like Hyvicrt- 
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 Triphana fimbria and 
Agrotis saucia were identified by J. E. Harting for Macpherson {loc. cit.). 



246 RHINOLOPHIDyE— 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 in a 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 RHINOLOPHID.E— RHINOLOPHUS 

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, loc. at., 291. 

2 Loc. cit. ; also, Cofnptes 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 20th 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 RHINOLOPHIDvE— 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,\n\., 131-133, pi. xx., figs, i and 2 ; also, Man. de VAcad. Roy. 

des Set., m, 382, pi. XV., fig. 4, 1759, published 1765 ; described from France. 
1774. Vespertilio ferrum-equinum (in part), J. C. D. von Schreber, Die 

Siiugthiere, i., pi. Ixii., two lower figures, 174, 190; evidently naming Daubenton's 

Le Fer-a-cheval; Montagu, Transactions Linnean Sac. (London), ix., 162, 1808; 

Bingley ; Clermont. 
1776. Vespertilio equinus (in part), P. L. S. Miiller, Naiursystems Supplements 

und Register Band, 20. 
1785. Vespertilio ungula (in part), P. Boddaert, Elenchus Animalium, i., 71 ; 

renaming Schreber's V. ferruin-equinum. 
1788. Vespertilio ferrum-equinum, ^. minor, J. F. Gmelin, Sy sterna Naturae., i., 50. 
1792. Vespertilio ferrum equinum minor, Robert Kerr, Animal Kingdom., 99 

(not V. molossus minor of Kerr, op. cit, 97) ; described from France. 

1797. NOCTILIO HIPPOSIDEROS, J. M. Bechstein, Der Zoologe, I., i.-viii., 65. 

1798. Vespertilio hippocrepis (in part), F. von P. Schrank, Fauna Boica, i., 64. 
1800. Vespertilio hipposideros, J. M. Bechstein, in Thomas Pennant's Allgemeine 

Uebersicht der vierfiissigen Thiere, ii., 629 footnote, see also 615 and 736; 
Donovan. 

1803. RHINOLOPHUS MINOR, Etienne Geoffroy, Catalogue Mammiferes Mus. d'Hist. 
Nat. (Paris), 57 ; described from Paris. 

1804. Vespertilio (? Rhinolophus) hippocrepis, Johannes Hermann, Observa- 
tiones Zoologicae, 18. 

1808. Vespertilio minutus, George Montagu, Trans. Linnean Soc. (London), ix., 
163, etc.; described from Devonshire, England; Pennant, ed. of 1812, 181, i., 
pi. xiv., figs. 3 and 4. 

1813. Rhinolophus bi-hastatus, Isidore Geoffroy, Ann. du Mus. d'Hist. Nat., 
XX., 259 ; renaming Daubenton's Le petit Fer-a-cheval. 

1816. Phvllorhina minuta, W. E. Leach, Systematic Catalogue of the Specimens 
of The Indigenous Mamtnalia and Birds that are Preserved in The British Museum, 
etc. (London), i ; named, without description, from Torquay, Devon. 

1817. Rhinolophus hipposideros, W. E. Leach, Zoological Miscella7iy, iii., 2, sp. 2, 
pi. 121, et pltcrimorum auctorum. 

1829. Rhinolophus bifer, Jakob Kaup, System der Europdische Thierwelt, i. 104 ; 

without description, hence a nomen nudum. 
1862-63. Rhinolophus hipposideros, var. typus, alpinus, et pallidus (in part), 

Carl Koch, fahrbilcher des Vereins fiir Naturkunde ijn Herzogthum (Nassau), 530- 

531 ; described vaguely and "scarcely determinable" (Andersen). 
1885. Rhinolophus bihastatus, var. kisnyiresiensis, Daday Jeno, Orvos- 

Termeszettudomdnyi Ertesito, etc., x, (3), 274 ; described from Kis-Nyires, Trans- 

sylvania. 



THE LESSER HORSESHOE BAT 251 

1887, Rhinolophus hipposideros van troglophilus, Daday Jeno, Erfekezesek 
A Termeszettudomdnyok ^t?>/^tf/ (Budapest), xvi., Nos. 7, 8, pi., figs, i 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 hipposideros : 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-equimun, but 
hipposideros would give place to minor. 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. hipposideros 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 minimus, inhabit- 
ing the Mediterranean region, Egypt excepted, with the Balearics, Malta, 
Corsica, and Crete, to Sennaar and Keran ; hipposiderus (sic) 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 minutus, of the British Islands. The distribution in France, 
as given by Rollinat and Trouessart {Mem. 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. hipposideros repeats and amplifies the 



252 RHINOLOPHID^— 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 wi-th 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 {loc. 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 {Trans. Linnean Soc, ix., 1808, 162- 171) ; and after- 
wards (with R. ferruvi-equinuDi) 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, Journ. cit., 1887, 63; Hollis, 
Joiirn. cit., 1907, iio-iii). Further records up to 1887 have been 
summarised by Kelsall {loc. cit.), showing that it occurs, often abund- 
antly, in Cornwall (see also Couch, Jotu^n. cit., 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 speciall)'- 
favoured haunt (see also 'Lqw'is, Journ. cit., 1906, 69, and Millais); and in 



PLATE XVI. 




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, Joiirn. 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 {Ann. 
and Mag. Nat. Hist., May 1840, 185); by Wallis near Ross {Trans. 
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 {Jotirn. cit., 1903,430) ; and Newstead, Caton Haigh, Oldham 
and Coward in Denbigh and Flint {Joiirn. cit., 1887, 152; 1896,255; 
1897, 537-538; 1906, 70-70- 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. cit., 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 Field, 12th 
April 1884, 499; Nat2imlist, 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 

Y 2 



254 RHINOLOPHID^— 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 {New Stat. Ace. 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 loth March 1859. 
The former obtained only one specimen {Proe. Dublin Univ. Zool. 
and Bot. Ass., 15th April 1859, in Dublin Nat. Hist. Review, \\., 522- 
525); but the latter adduced strong evidence to prove that it is 
the common bat of the district around Ennis iProc. Dublin Nat. Hist. 
Soc., 1st April 1859, in Dublin Nat. Hist. Reviezv, vi., 379-38 0-^ 
These localities have since been confirmed for Clare by J. R. Kinahan 
{Zoologist, 1861, 7617-7624), and Scharff {Irish 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, loc. cit., 
92-93), and later from the same district by Forrest {in lit. 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 oi 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 {Naturalist, March 
1907, 74, and in lit.). Proger, however, saw young ones born in 
South Wales about 20th June {in lit.). He has also once observed 
spring courtship (see Proe. 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, Irish Naturalist, 1897, 36, gives the dates of occurrences differently, 
but, as I think, incorrectly. 



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." 

The wing, ear, and foot are coloured as 
in R. ferruni-equinuin. 

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 skxill (Fig. 26) the auditory 
bullae 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. 

The anterior upper premolar, though 
small, is in the tooth-row, the upper canine 
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 




Fig. 26.— Diagram of Teeth of 

Rhinolophus hipposideros. 

(i) Upper and (2) Lower Jaw, 



•56 



RHINOLOPHIDyE— RHINOLOPHUS 



detail. Kelsall {Zoologist, 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 417 and 347 to 38, and of the skull 16 and 
14-5 to 15-5 mm. in the typical sub-species and in Jiiiniinus respectively. 
Minutus agrees in skull with the type, in forearm with minimus. The 
differences are slight, but constant for a series. 



DIMENSIONS IN MILLIMETRES:— 







J 


^ 






^ 

















o 


Ml 


1| 




ti 


o 


E 




]5; 


"eS 


"3 






73 
1= 


S 


-"■H-g 


rt 


^ 




rt 


a 

c3 


m 




04 


^ to 




■a 


C8 




H 


1 






to 
S 


C3 
C3 


1 


c; 




w 


C 


§"1 






a 




2 


iJ 


^ 


8 


cS 








^5 






K. 




H 











Three males (H. J. Char-^ 
bonnier) . . . / 










17 
17 
15 


\ ( 
/■•\ 


34-4 
37-3 
35-3 


}■ 


f 
•l 


23 
24 
24-5 


20 ■^ 
28 - 
20 j 




Thirty specimens "j Max. 
measured by K. [- 
Andersen . J Min. 




15'u 


10-n-s 


27 


lS-5 


S-7 


3'J 






24-9 


28-2 






























\i-l 


0- 6-7 


23-5 


'10-3 


7'5 


30-3 






22-8 


24-7 




Female, Devonshire (1), 


























measured in dried skin 


























at British Museum. 


39 


15-5 




20 


8 


S-8 


30 


4 


51 


23-5 


25 


241 


Female, do. (2), do. . 


38 


15-5 




2S 


10 


10-S 


38 


4 


53 


23 


20 


250 



Note :— Embryos within the mother reach an expanse of 100 to 110, and by July the young attain to 190. 

Proportionate lengths : — Foot, without claws, about -50 of lower leg ; 
fifth metacarpal, about i-i 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 ?n^ to 
anterior border of canine, 57 ; greatest breadth at zygoma, 8 ; posterior 
breadth, 6-8. 

The weight of three is given by Couch (Zoologist, 1853, 3941), as 
yy, 74, and y2> grains = 5, 4-8, and 47 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 R. 
ferrum-equiniim. 

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 



THE LESSER 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 is, 
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 winQTs. 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 feeling for an 
opening by which to escape. But, at the same time, its 
shyness when approached gave Tomes the strong but, as it 



258 RHINOLOPHID/E— 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 Nat. Hist. Review^ vi., 379-381, 
1859. 

^ Zoologist^ 1 86 1, 7617-7624. 



THE LESSER HORSESHOE BAT 259 

males, and of the four females two were found on ist 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 too^ether 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 re^-ard 
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, iir. 
^ Irish Naturalist, 1907, 140-144. 



26o RHINOLOPHID^— RHINOLOPHUS 

have supplied me with independent information that they 
have found the bats inhabiting caves opening on to bare 
hillsides. 

At Cefn, on i8th 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 ist 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. ^ Meta menardi. 

3 In Kent's Hole, Devonshire, on the 8th (Oldham). ^ Zoologist, 1903, 430. 



PLATE XIX. 




Greater Horseshoe Bats. 



THE LESSER HORSESHOE BAT 261 

faecal 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. 

^ /?. curyale of Blasius. ^ Myotis emarginatus. 

^ In lit. ^ Naturalist., March 1907, 74, etc. 



262 RHINOLOPHID^— 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, as a 
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. 



PLATE XX. 




Lesser Horseshoe Bats. (.Natural size.) 



MicroTormea by 

Preservation 

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