;-jj5^]^
o
B
CATALOGUE
OF THE
MAMMALS
OF
WESTERN EUEOPE
(EUROPE EXCLUSIVE OF RUSSIA)
IN THE
COLLECTION
OF THE
BEITISH MUSEU])^^^?^
BY
GEREIT S. MILLER
/-/■
LONDON
PRINTED BY ORDER OF THE TRUSTEES OF THE
BRITISH MUSEUM
SOLD BY
Longmans, Green & Co., 39, Pateenostkr Row, E.G.
B. QuARiTCH, 11, Grafton Street, New Bond Street, W.
DuLAU & Co., Ltd., 37, Soho Square, W.
AND at the
British Museum (Natural History), Cromwell Road, S.W.
1912
All rights reserved
LONDON :
PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,
DUKE STREET, STAMFORD STREET, S.B., AND GREAT WINDMILL STREET, W.
hh.y^i '^1^' '^'^-^ dhu't.
PEEFACE
Although the idea of a publicatiou ou the Mammals of
Europe was suggested many years ago by the late Lord
Lilford, who kindly contributed an annual sum towards the
collecting necessary for its realization, the possibility of
issuing the present Catalogue has mainly grown up from the
work which its author, Mr. Gerrit S. Miller, of the United
States National Museum at Washington, has for some years
been doing independently on the subject.
It is true that European Mammals had not been neglected
here, and that the collection had begun to grow, both by
the help of the Lilford Eund and by the efforts of Major
G. E. H. Barrett-Hamilton, who published many papers on
the subject, and of Mr. Oldfield Thomas, E.E.S., who devoted
a number of his vacations to collecting in various parts of
the Continent.
There was, however, no prospect of being in a position
to prepare a Catalogue until about 1905, when Mr. Miller
arranged to devote his entire time for a considerable period
to the study of European Mammals. The opportunity was
taken of having the results of this work published here
instead of in America, by inducing him to write a British
Museum Catalogue ; thus utilizing his knowledge, and com-
bining for the purposes of his studies the material of both
the American and the British National Museums.
Collections were then made in various selected areas,
partly by Mr. Miller himself and partly by trained collectors,
such as Messrs. A. Eobert, C. Mottaz, Eev. S. Gonzalez and
N. Gonzalez, the cost of whose services were contributed by
friends of the Museum, notably Mr. Oldfield Thomas, the
Hon. N. C. Itothschild and ^Ir. J, I. S. Whitaker. The
Catalogue could hardly have been contemplated if it had
not been for Mr. Thomas' unremitting efforts in developing
IV PREFACE
tlie collection. He has not merely regarded tliese efforts as
an official duty, but he has in addition been a generous
donor who has frequently supplied funds for the purpose of
obtaining specimens. Mr. Miller has thus had at his
disposal a collection fairly representative of all parts of
Western Europe, and immensely superior to anything that
had been thought of before he began his work.
Marine Mammals (Cetacea and Pinnipedia) are not
included in the present Catalogue. For a definition of
" Western Europe " reference must be made to page vii of
the Author's Introduction.
As Mr. Miller is on tlie staff of the United States
National Museum the special and cordial thanks of the
Trustees of the British Museum are due to the authorities of
the former Institution for the facilities granted to him for
carrying through the preparation of the Catalogue, a work
which involved a furlough of two years and a half from his
usual duties at Washington.
The thanks of the Trustees are also due to Mrs. Oldfield
Thomas and to Mr. E. C. Wroughton for their kindness
in undertaking the considerable labour of preparing and
verifying the lists of the specimens in the Museum collection
after Mr. Miller had made his scientific determinations.
SIDNEY E. HAEMER,
Keeper of Zoology.
British Museum (Natural History),
London, S.W.
October, 1912.
INTEODUCTION
The collection of European Land-mammals in the pjiitish
Museum consists of about five thousand specimens. One
hundred and twenty-four of these are types. It has for the
most part been brought together during the past thirty years
through the efforts of the late Lord Lilford, of Mr. Oldfield
Thomas, and of Major G. E. H. Barrett-Hamilton. The older
material, though not extensive, includes much that is of historic
interest, such as the numerous specimens received from the late
Baron E. de Selys-Longchamps, the types of various species
described by Gray and Bonaparte, and Darwin's Porto Santo
rabbits which have been the subject of so much groundless
speculation. It is, however, from the recently-obtained material
that the collection derives its true value. These specimens are
almost without exception carefully-prepared skins accompanied
by skulls and measurements, together with full records of sex,
date, and exact locality. The more important sources from
which they were obtained are as follows : collections brought
together by Professor W. Wolterstorff from central and eastern
(Jermany, and presented by the late Lord Lilford ; collections
made in Roumania by the late E. Dodson under the direction
of Major G. E. H. Barrett-Hamilton, and presented by the late
Lord Lilford ; material from southern Spaia presented by the
late Lord Lilford ; extensive collections made in south-western
France, in southern Italy and in Sicily by A. Pvobert and
presented by Mr. Oldfield Thomas; collections from south-
central France and the vicinity of Strassburg, Germany, made
by C. Mottaz and presented by Mr. O. Thomas ; small collections
from Denmark, Holland, Pas-de-Calais, Brittany, Portugal, the
Balearic Islands, Switzerland and northern Italy, made and
presented by Mr. O. Thomas ; collection from central and
northern Spain made by N. Gonzalez and presented by i\Ir. O.
Thomas ; a large collection from miscellaneous sources brought
together and presented by Major G. E. H. Barrett-Hamilton;
a collection from Greece made by C. Mottaz and presented by
VI INTRODUCTION
Mr. J. I. S. Whitaker and the Hon. N. C. Rothschild; a
collection from Spain and southern France made by G. S.
Miller and purchased by the Museum ; several collections
from Transylvania made by C. G. Danford ; collections from
Hungary made and presented by the Hon. N. C. Eothschild
and Mrs. Rothschild ; smaller collections and single specimens
have been received from many other persons,* whose names will
be found in the detailed lists of material in this Catalogue.
Although unquestionably forming the largest of all collections
of European mammals the material in the British Museum is
not sufficient to be made the basis of a monographic study of
the fauna. Free use has, therefore, been made, throughout the
preparation of this Catalogue, of specimens in other collections.
Chief among these are the United States National Museum in
Washington and the private collection of Charles Mottaz in
Geneva. The material at Washington, about 4000 specimens,
is mostly from the following sources : (a) Sweden, Germany,
Switzerland and Belgium, collected by J. Alden Loring ; (b)
Sicily, Italy and the region of Barcelonnette, Basses-Alpes,
France, by Dane Coolidge ; (e) south-western France, by
Robert T. Young ; (d) north-eastern Germany, the Riesengebirge
and Hai'z Mountains, by F. L. J. Boettcher. There are also
miscellaneous smaller collections from Switzerland (G. S. Miller,
L. Stejneger, E. H. Zollikofer), Belgium (de Selys-Longchamps),
Holland (G. S. Miller), Denmark (L. Stejneger), Norway (T.
Stejneger) and Sweden (Sundevall. Tullberg, Lonnberg). Finally,
the Merriam collection, now the property of the U.S. National
Museum but not yet catalogued, contains numerous European
specimens, for the most part received from de Selys-Longchamps.
The Mottaz collection, about 3000 specimens, is especially rich
in series of the smaller mammals of Switzerland and the adjoin-
ing portions of France ; it also contains useful material from
Italy and western France (Charente). Other supplemental
material to which I have been given free access, or which has
been sent for examination in London or Washington, is contained
in the museums of Madrid (types of Cabrera), Nimes (types of
Crcspon), Paris (types of Geoffroy and other historic speci-
mens), Genoa (Italian Bats, Microtines and Ungulates), Turin
(Italian mammals, especially Ungulates), Naples (type of Mi/otis
oxygnathus Monticelli), Geneva (types of Fatio, authentic Swiss
* This is particularly true of the many friends of the Museum who
have aided in procuring the large series of British mammals.
INTRODUCTION VU
specimens of Lynx), Lausanne Agricultural School (skull of
Ursus "formicnriiis " from the Alps), Munich (type of S/ialnx
(jrsecus Nehring), Berlin Agricultural High School (type of
Arvicola rattireps stimminiji Nehring), Breslau (skulls of foxes),
Leiden (co-types of Arvicola arenarius de Selys-Longchamps),
Copenhagen (Mus fseroensis and small carnivores), Christiania
(Sorex, Evotomys, etc), Stockholm (Swedish carnivores and
rodents), Cambridge {Mustela erminea ricinse, Lemmus lemmus
" n-nssidens") and Edinburgh (rodents from northern Scotland).
Private collections which have been in the same generous
manner placed at my disposal are those of Mr. Angel Cabrera,
of Madrid (Spanish mammals, including several types). Dr.
Enrico Festa, of Turin (Italian mammals), Mr. Angelo Ghidiui,
of Geneva (Swiss and north Italian mammals), and Dr. Fernand
Lataste, of Cadillac-sur-Garonne, France (carnivores and micro-
tines).
The total number of sj^ecimens on which this work is based
approximates 11,500. All those of which definite record has
been made are enumerated in the paragraphs headed : Specimens
examined* Absence of a note to the contrary indicates that
all the specimens from a given locality are in the British
Museum. Discrepancies frequently occur between the number
of " specimens examined " and the number tabulated in the final
paragraph as forming part of the Museum collection. These
result from the fact that under "specimens examined" are
included duplicates as well as registered specimens, while only
the latter appear in the final lists. f
For the purposes of this Catalogue, " Western Europe " is
regarded as including the continent of Europe outside the
frontiers of Russia; also the immediately adjacent islands, and
Spitzbergen, Iceland, and the Azores. The members of the
living mammal fauna of this region, exclusive of the cetaceans,
pinnipeds, and species such as Buhalus huhalis in Italy and
Simia sylvamis J on the Rock of Gibraltar, which certainly owe
* In these lists 11,372 specimens are recorded. They are distributed
as follows: Insectivora, 1,777 ; Chiroptera, 2,210; Carnivora, 877 ; Rodentia
Duplicidentata, 379, Rodentia Simplicidentata, 5,854 ; Ungulata, 28i.
t Certain duplicates have been transferred to the United States
National Museum since the lists of " specimens examined " were prepared.
No attempt has been made to alter the records in the lists on this account ;
but the U.S.N.]\I. numbers of such specimens are frequently to be found
in the Tables of cranial measurements. (All numbers above 10,000 indicate
specimens in Washington.)
X For use of this name in place of " Macacus inuus," see Thomas, Proc.
Zool. Soc. London, 1911, pp. 125-126, :March, 1911.
VIU INTRODUCTION
their presence to artificial introduction, are treated mono-
graphically on the basis of the material already enumerated.
This material has been found sufficient, in most of the groups, to
give what appears to be a fairl}' satisfactory idea of the essential
features of the fauna. In the ungulates and the larger car-
nivores, however, it is so totally inadequate that no attempt
could be made to revise the genera by which they are repre-
sented. This is especially to be regretted on account of the fact
that some of these larger mammals are nearly extinct, while
others are being modified by the introduction of foreign stock
to replenish exhausted game preserves. Immediate action is
necessary if the final opportunity to gain a clear understanding
of this part of the European fauna is not to he lost.
The litei'ature of European mammals is so voluminous,
particularly as regards local lists and special notes on distri-
bution, and it is for the most part based on conceptions of
species and local races so different from those underlying the
present work, that an amount of labour incommensurate with
the importance of the results would be required to prepare
extended bibliographical Tables for each form recognized. The
citations are, therefore, restricted to those which seem of impor-
tance in giving a clear idea of the systematic history of each
animal ; that is, to the specific and sub-specific names under
which it may have been described, to the first use of the actual
binomial or trinomial here adopted, to the names used in the
monographic works of Blasius, 18-57, and Trouessart, 1910, and
to any uthei- publication which might seem pertinent to a
particular case.
In deciding questions of nomenclature, an attempt has been
made to apply the International Code and the rulings of the
Commission strictly and consistently, even to the reluctant
acceptance of the terms applied to genera by authors who
f(jllowed a system different from that now in use.
With the exception of figure 121, lent by the Smithsonian
Institution, all the illustrations are original. The di'awings of
teeth were made in London by Mr. A. J. Engel Terzi ; part
of those of the skulls were made by Mr. Terzi ; the rest were
done in Washington by Mr. H. B. Bradford.
A few words in conclusion regarding the actual making of
the manuscript. I prepai'ed all the descriptions, synonymies,
lists of specimens examined, and Tables of cranial measurements.
The external measurements, which are not to be regarded as
INTRODUCTION IX
more thau approximately accurate, are mostly given as recorded
on the labels, though much verification and correction for ears
and hind feet has been done from the dried specimens. In order
to economize time, the records of registered material were made
directly from the specimens which I had identified. Mrs.
Oldfield Thomas and Mr. R. C. Wroughton carried out this
portion of the work.
G. S. M.
Washington,
July 1, 1912.
Order.
Number of
genera
recognized.
Number of
forms
recognized.
Number of
forms not
represented
in B.M.
Number of
recognized
forms not
seen.
Insectivora
7
45
5
Chieoptera
10
33
Carnivora
15
47
1
Rodentia Duplicidentata
2
19
1
„ Simplicidentata
26
139
6
2
Ungulata
9
31
9
4
Total
m
314
22
G
SYSTEMATIC INDEX
ORDER INSECTIVORA.
Family TALPID^ . .
Sub-Family 1. Talpin-tj: .
1. Talpa Linnaeus
1. europgea Linnfcus .
2. casca Savi
3. occidentalis Cabrera
4. romana Thomas
Sub-Family 2. Desmanin.e
1. Galemys Kaup
1. pyrenaicus Geoffroy
a. pyrenaicus Geoffroy
h. rufulus Graells .
Family SORICID^ . .
1. Sores Linnseus
1. araneus LiniuBUs .
a. araneus Liniueus
b. castaneus Jeoyns
c. santonus Mottaz
d. euronotus IMiller
e. bergensis Miller .
/. tetragonurus Hermann
g. pyrenaicus Miller
h. fretalis Miller
i. granarius Miller .
2. rainutus Linna»us .
a. minutus Liniia'us
h. lucanius Miller .
3. alpiuus Schinz .
a. alpinus Schinz .
h. hercynicus Miller
2. Neomys Kaup .
1. fodiens Schrebcr
a. fodiens Schreber.
h. bicolor Shaw .
2. milleri Mottaz .
3. anomalus Cabrera .
3. Pach>'ura de Selys-Long
champs ....
1. etrusca Savi.
4. Crocidura NVagler.
1. leucodou Hermann
2. mimula Miller .
a. mimula Jlillor .
h. iculisma Mottaz ,
c. cantabra Cabicra
2
3
3
15
15
18
20
20
21
26
2G
28
29
31
35
37
40
41
41
42
44
45
52
53
55
GO
CO
62
63
65
66
69
73
78
81
81
83
86
88
94
95
98
99
PAGE
8. russula Hermann ... 99
a. russula Hermann . . 101
h. pulchra Cabrera . . . 103
c. cintrre Miller. ... 108
4. sicula :Miller .... 108
5. cauere Miller .... 109
6. caudata Miller .... 110
7. cyrnensis Miller . . . Ill
8. balearica Miller ... 112
Family ERINACEID.-E ... 114
1. Erinaoeus Linnaeus . . . 114
1. europseus Linna'us. . . 115
a. europreus Linna'Us . . 120
h. hisiaanicus Bariett-
Hamilton . . . . . 122
c. italicus Barrett-Hamil-
ton 123
d. consolei Barrett-Hamil-
ton 126
2. roum aniens Barrett-
Hamilton 127
3. nesiotes Bate .... 129
4. algirus Duvernov and
Lereboullet . ". . . 130
a. algirus Duvernoy and
Lereboullet .... 131
b. vagans Tliomas . . . 133
ORDER CHIROPTERA.
Sob-Ordee MICROCHIROPTERA.
Family RHINOLOPHID.^ . . 136
1. Rhiuolophus Lac6pede . . 137
1. ferruin-oquinum Schrober 189
a. ferrum-equinum Schre-
ber 142
b. i n s u 1 a n u s Barrett-
Hamilton 147
2. hipposideros Bcchstein . 147
a. hipposideros liechstein . 149
b. minimus Heuglin . . 151
c. minutus Montagu . . 154
8. euryalc Blasius .... 155
4. meholyi Matschie . . . 159
5. blasii Peters 162
SYSTEMATIC INDKX
Order Chikopteka — Sub - Order
MlCBOCHlKOPTEBA— aw^i.
Family VESPERTILIONlD.t: . 165
Sub-Family Vespkktiuo:sis.t: . 165
1. ilyotis Kaup 166
1. mystaciuus Kohl . . 169
2. nattereri Kohl .... 17-i
3. emargiuatus Geoffroy . . 177
4. bechsteimi Kuhl . . 179
5. daubentonii Kuhl . . 1S4
6. capaccinii Bouaparce . . 1S7
7. dasycneme Boie . . . 189
8. myotis Borkhauseu . 192
9. oxvgnathus Monticelli . 199
2. Pipistrellus Kaup ... 202
1. pipistrellus Schrebor . . 201
2. uatkusii Keyserliug aud
Blasius 213
3. kuhlii Kuhi 215
4. savii Bonaparte .... 219
3. Eptesicus Rafinesque . . 224
1. serotinus Schreber . . 226
2. sodalis Barrett-Hamiltou . 231
3. uilssonii Kevserling and
Blasius. .".... 234
4. Vespertilio Linnaeus . . . 238
1. murinus Lin me u-^ . . . 238
5. Nyctalus Bowdieh ... 242
1. maximus Fatio. . . . 244
2. noctula Schreber . . . 245
3. leisleri Kuhl .... 252
4. azoreuni Thomas . . . 254
6. Plecotus GeofEroy. . . 256
1. auritus Linnwus . . . 256
7. Barbastella Gray .... 263
1. barbastellus Schrebt-r . . 263
Sub-Family Minioptesisj. . . 266
1. Miniopterus Bonaparte . . 268
1. schreibersii Kuhl . . . 269
Family MOLOSSID.E ... 276
1. Nyctrnomus Geoflroy . . 276
1. teniotis Rafinesque . . 277
ORDER CARNIVORA.
Family URSID.E 2S4
1. Ursus Linnseus .... 285
1. arctos T.inn.f n< . . . 285
2. Thalarctos Gray .... 297
1. maritimus Fhipp> . . . 298
Family CAXID.F 303
1. Canis Linnfeus .... 304
1. lupus Linnseus .... 305
a. lupus LinnKUs . . . 313
6. signatus Cabrera . . 314
c. deitanus Cabrera . . 315
2. aureus Liunieus . . . 315
VAGK
2. Alopex Kaup 318
1. lagopus Liiuuvus . . . 319
2. spitzbergenensis Barrett-
Hamilton aud Bouhote . 324
3. Vulpes Oken 325
1. vulpes Linn*us . . . 326
a. vulpes Liuuams . . . 330
6. crucigera Bechsieiu. . 331
c. silacea Miller. . . 333
2. ichnusse Miller .... 336
Family MUSTELID.E . . . 340
Sub- Family Mklix.?; .... 341
1. Meles Brissou 341
1. meles Linnaeus .... 343
a. meles Linnseus . . . 348
b. mariauensis GraeUs. . 352
2. arealus Miller .... 352
Sub-Family LuxRrs.t: .... 354
1. Lutra Brissou 354
1. lutra Linmvus .... 355
Sub-Family Mustehn.*; . . . 364
1. Martes I'inel 365
1. martes Linnieus . . . 366
a. martes Linnseus . . . 372
b. latinorum Barrett-Ha-
milton 373
2. foina Erxleben .... 374
(I. foina Erxleben . . . 375
h. mediterranea Barrett-
Hamilton 360
3. bunites Bate .... 380
2. Mustela Linnseus. . . . 381
Sub-geiius Mustela Linnseus . 384
1. erminea Linnseus . . . 385
a. erminea Linuseus . . 387
b. sBstiva Kerr .... 389
c. stabilis Barrett-Hamil-
ton 390
d. riciuifi MiUer. ... 397
2. hibernica Thomas and
Barrett-Hamilton . . 398
3. nivalis Linnseus . . . 401
a. nivalis Linnseus . . 402
6. boccamela Bechstoiii . 405
c. iberica Barrett-Hamilton 407
4. africana Desmarest . . 412
5. galinthias Bate . . . 414
Sub-Genus Lutreola Wagner . 415
6. lutreola Linnseus . . . 415
Sub-Genus Putorius Cuvier . 418
7. putorius Linnseus . . , 419
a. putorius Linnseus . . 423
b. aureolus Barrett-Hamil-
ton 425
3. Vormela W. Blasius ... 428
1. peregusna Gueldenstaedt . 429
Sub-Family Gulonls-E . . . 432
1. Gulo Storr 433
1. ?ulo Linnseus .... 434
KTSmUTIC IKDEX
PAGK
(jrder Cabsivoea — conlinu^l.
Family VIVERKID>iE . .
440
1. Mungofi Geoff roy and Cuvier 440
1. widdringtonii Gray
441
2. Genetta Oken ....
44e
1. genetta LinnasuB .
447
a. genetta Ltinnmnn
451
b. balearica Thomasj .
452
c. rhodanica Matschie .
452
Family FBLID>E ....
1-5.5
1. Felk Linrijeus.
456
1. Hilvestrii-; Schrel>er .
457
a. silvestris Schreter .
462
fc. grarapia Miller .
464
c. tartesBia Miller .
465
2. sarda Lataste .
468
3. agrius Bate ....
470
2. Lynx Kerr
470
1, lynx Linnajus .
471
2, pardellus Miller . .
475
ORDER RODENTIA.
Sub-Okdkk duplicidentata.
Family LEPORID^ .... 484
1. OryctoLagus Lilljeborg . . 484
1. cuni/.-uluH Linrijeiis . , 48-5
o. cxiniculu.8 Linrijeus . . 490
6. huxleyi Haeckel . . . 491
2. Lepus Linnaeus .... 495
1. europfftUB PaJlas . . . 498
a. europaius Pallas. . . 502
b. occidentalis de Winton. 50^
c. pyrenaicus Hilzheimei* . 506
d. meridiei Hilzheimer . 506
e. corsicanus de Winton . .507
/. hybriduB Desmarest. 508
g. tranBsylvanicus Mat-
schie 509
2. creticus Barrett-Hamilton 512
3. mediterraneus Wagner . 513
4. granatensis Flosenhauer . 515
a. granat^jnsis Pi^jsenhauer 516
6. gaUiecius Miller . . 517
c. iturisBiuB Miller . 518
d. pamasBius Miller . . 519
5. timidus Linnaeus . . . 522
a. timidus Linnaeus . 526
b. varronis Miller . . . 528
c. scoticus Hilzheimer. . 529
6. hibemicus Bell. . . . 531
Scb-Obdee SIMPLICIDENTATA.
Family ZAPODIDiE .... 5-35
Scb-Family S1CIETIH.S; . . . 536
1. Sicista Gray 536
1. loriger Nathusius . . . 537
2. trizona Pet^nyi .... 539
J-ACE
Family HYSTBICIDiE . .
.542
1. Hyetrix Linnaeus .
542
1. crigtata Linnseus
543
Family MUSCARDINID^^ .
549
1. Eliomys 'SVagner .
5.50
1. quercinus LinnseuB
5-51
2. gymnesicus Thomas .
558
3. paUiduB Barrett-Hamilton 559
4. sarduB Barrett-Hamilton
500
5. lusitanicus Pi^uvens .
560
2. DyromyB Thomas.
.566
1, nitedula Pallas . . .
567
a. nitedula Pallas .
568
6. intemiediuB Nehring
569
c. wingei Nehring .
570
2. robustus Miller.
572
3. Glis BrisBOn ....
572
1. glis Linnaeus
.573
a. glis Linnaeus .
.577
t. italicus Barrett-Hamil
ton
.578
c. melonii Thomas .
579
d. pyrenaicus Cabrera .
582
4. Muscardinus Kaup .
583
1. aveDanarius Linnaeus .
583
2. pulcher Barrett-Hamilton 590
Family MUEID.S: . . .
591
Scb-Family Ceicetot^ . .
592
1. CriMjtultiB Milne-Edwards
.593
1. atticus Nehring
593
2. Cricetus Leske
.596
1. cricetus Linnaeus .
597
a. cricetus Linn3?UB
602
6. canesoens Nehring .
603
c. nehringi Matschie .
605
3. ilesocricetuB Nehring .
605
1. newtoni Nehring .
606
Sub-Family Micbotis^ . .
610
1. MyopuB Miller
611
1. Bchisticolor Lilljeborg.
611
2. LemmusLink.
614
1. lemmuB Linnaeus .
615
3. Evotomys Coues .
623
1. glareolus Schreber .
626
a. glareolus Schreber .
a32
b. britannicus Miller .
6-34
c. suecicuB JliUer .
6-36
d. istericus iliUer .
6-37
e. norvegicns ililler
6.38
/. vasconiae Miller .
639
g. helveticus Miller
640
k. nageri Schinz
641
i. hallucalis Thomas .
643
2. skomerensis Barrett -Ha
milton ....
. 644
3. caesariuB ililler .
. 645
4. rutilus Pallas .
. 646
5. rafocanus Sundevall .
. 648
SYSTEMATIC INDEX
and
Order Rodentia — Sub - Order
SiMPLiciDENTATA — Family
MuEiD.E — Sub-Family ili-
CBOTIN.E —continued.
4. Microtus Schrank
Sub-Genus Microtus Schrank
1. agrestis Linuieus .
a. agrestis Linnseus
b. exsul Miller .
c. levernedii Crespon
d. bailloni de Selys-Long-
champs
e. hircus Bellamy .
/. neglectus Jenyns .
g. rozianus Bocage .
2. arvalis Pallas
a. arvalis Pallas
b. meridianus PaUas
c. duplicatus Rorig
Borner
d. levis Miller .
3. incertus de S61ys - Loug-
champs
4. asturianus Miller .
5. orcadensis Millais .
6. sandayensis Millais
a. sandayensis Millais
b. westrse Miller
7. sarnius Miller .
8. cabrerge Thomas
9. dentatus Miller.
10. hartingi Barrett-Hamilton
11. angularis Miller
12. ratticeps Keyserling and
Blasius .
Sub-Genus Chionomys Iililler
13. nivalis Martins
a. nivalis Martins .
b. aquitanius Miller
14. lebrunii Crespon .
a. lebrunii Crespon.
b. leucurus Gerbe .
15. ulpius Miller .
5. Arvicola Lac^pede
1. amphibius Linnaeus
a. amphibius Linnaeus
b. reta Miller
2. sapidus Miller .
a. sapidus JMiller
b. tenebricus Miller
3. terrestris Linnaus.
4. italicus Savi.
5. illyricus Barrett-Hamilton
6. musignani de Selys-Long-
cbamps
7. scherman Shaw
a. scherman Shaw .
b. exitus Miller .
c. monticola de
Lougchamps .
Selys-
658
659
662
668
669
671
672
673
675
680
681
683
686
686
687
690
693
694
696
697
698
700
701
703
704
706
708
712
713
716
717
718
719
722
723
723
725
730
732
732
733
735
738
740
741
744
744
745
746
749
6. Pitymys MclMurtrie .
1. subterraneus de Selys-
Longchamps ....
a. subterraneus de Selys-
Longchamps ....
b. capucinus Miller
2. dacius ^Miller ....
3. drueutius Jliller
4. fatioi Mottaz ....
5. multiplex Fatio
6. savii de Selys-Longchamps
7. nebrodensis Mina-Palumbo
8. pyrenaicus de Selys-Long-
champs
a. pyrenaicus de Selys-
Longchamps ....
b. brunneus Miller .
9. planiceps Miller .
10. gcrbii Gerbe ....
11. lusitanicus Gerbe.
12. marite Major ....
13. pelandonius MiUer .
14. depressus Miller .
15. ibericus Gerbe
a. ibericus Gerbe .
b. centralis Miller .
c. pascuus Miller .
d. regnlus Miller
16. duodecimcostatus de
Selys-Longchamps
17. provincialis Miller
18. thomasi Barrett-Hamil-
ton
19. atticus Miller ....
Sub-Family IMurin.e ....
Apodemus Kaup ....
1. epimelas Nehring .
2. sylvaticus Linnaeus
a. sylvaticus Liunaus .
b. callipides Cabrera
c. dichrurus Rafinesque .
d. creticus Miller .
3. hebridensis de Winton
4. hirtensis Barrett-Hamil-
ton
5. fridariensis Kinuear .
6. liavicollis jNIelchior
a. flavicoUis Melchior .
b. wiutoni Barrett-Hamil-
ton
7. agrarius Pallas . . . .
. Micromys Dehne . . . .
1. minutus Pallas . . . .
a. soricinus Hermann .
b. xjratensis Ockskay
. Epimys Trouessart .
1. rattus Linnteus . . . .
a. rattus Linnaeus .
b. alexandrinus Geoft'roy .
2. uorvegicus Ersleben .
PAGE
752
755
758
760
760
762
763
764
768
770
770
771
772
772
773
776
777
778
779
780
782
782
783
784
784
785
786
787
791
791
794
797
803
809
810
818
824
825
825
828
829
831
836
840
841
844
846
848
849
853
854
858
SYSTEMATIC INDEX
Order Rodentia — Sub - Order
SiMPLiciDENTATA — Family
MuBiD.E — Sub-Family Mu-
KiNyE — continued.
4. Mus LimisBus ....
1. musculus Linua3us
fl. musculus Linna3US .
b. aiioricus Schinz .
2. muralis Barrett-Hamilton
3. fteroensis Clarke
4. spicilegus Pet6nyi .
a. spicilegus Petlnyi .
b. hispanious Miller
c. lusitanicus Miller
5. Acomys Geoffrey .
1. miiious Bate
Family SPALACID^ .
1. Spalax Gueldenstaedt
1. dolbrogeae Miller .
2. hungaricus Nehring
3. greecus Nehring
Family SCIURID^ . .
1. Sciurus Linnaeus .
1. vulgaris Linnajus .
a. vulgaris Linnaeus
b. varius Gmelin
c. leucourus Kerr .
d. russus Miller.
e. fuscoater Altum .
/. italicus Bonaparte
g. lilajus Miller .
h. alpinus Desmarest
i. numantius Miller
j. infuscatus Cabrera
A;, segurse Miller
I. bffiticus Cabrera .
2. CitcUus Oken . . .
1. citellus Linuajus .
2. suslica Gueldenstaedt
3. Marmota Blumenbach
1. marmota Linnoeus .
2. bobak Miiller . .
Family PETAURISTID^
1. Sciuropterus F. Cuvier
1. russicus Tiedemann
Family CASTORID^
1. Castor Linnaeus .
1. fiber Linnaeus .
863
865
869
871
874
875
877
878
879
882
883
883
887
887
889
894
895
897
898
898
905
906
907
909
910
912
913
914
914
916
917
923
924
924
929
931
932
937
940
941
941
947
947
947
ORDER UNGULATA
page
Family SUID^ 956
1. Sus Linnteus 956
1. scrofa Linnaeus. . . . 957
[attila Thomas] .... 960
2. meridionalis Major . . 960
Family CERVID^ .... 962
1. Cervus Liunteus .... 963
1. elaphus Linnaeus . . . 964
a. germanicus Desmarest . 965
b. elaphus Linnaeus . . 967
c. atlanticus Lonnberg . 967
d. scoticus Lonnberg . . 968
e. hispanicus Hilzheimer . 969
/. corsicanus Erxleben. . 969
2. Dama Hamilton Smith . . 970
1. dama Linuffius .... 970
3. Capreolus Gray .... 972
1. capreolus Linnaeus. . . 973
a. capreolus Linnaeus . . 974
b. transsylvanicus Matschie 975
c. canus Miller .... 975
d. thotti Lonnberg. . . 975
4. Alces Gray 976
1. alces Linnaius .... 978
5. Rangifer Hamilton Smith . 979
1. tarandus Linnaeus . . . 980
2. fennicus Lonnberg. . . 981
3. platyrhynchus Vrolik . . 985
Family BOVID.E 986
1. Ovis Linnaeus .... 986
1. musimon Pallas . . . 987
2. Capra Linnaeus .... 988
1. ibex Linnffius .... 989
2. pyrenaica Schinz . . . 989
a. pyrenaica Schinz . . 1)90
b. liisitanica Franca . . 991
c. victoriae Cabrera. . . ',)91
d. hispanica Schimper. . 991
3. segagrus Erxleben . . 992
3. Rupricapra BlaiuviUe . . 992
1. rupicapra Linnasus . . 993
2. ornata Neumann . . . 994
3. pyrenaica Bonaparte . . 995
4. parva Cabrera .... 995
APPENDIX.
Crocidura ichnusas Festa . . . 998
Evotomys glareolus norvegicus —
cranial measurements . . . 999
Spalax — Forms recognized by
M6hely. . 1000
CATALOGUE
OP THE
LAND-MAMMALS OF WESTERN
EUROPE.
Order INSECTIVORA.
1827. Insectivora Gray, Griffith's Cuvier, Aoim. Kingd., v, -p. 100.
Gcoijraphical distribution. — Africa (iucluding Madagascar),
Europe, Asia (including the Malay Archipelago), North America,
^^ Greater Antilles, and extreme north-western portion of South
America.
Characters. — Terrestrial, non volant, placental mammals with
low development of brain, the cerebral hemispheres without
convolutions ; teeth of a primitive or modified tuberculo-sectorial
type, the postei'ior upper premolar and anterior lower molar
never specially modified as carnassials.
Bemarhs. — The mammals of this order present such diversity
of form and structure that it is difficult to frame any definition
by which all members of the group may be invariably recognized
by skeletal or external characters. Ten families of Insectivora
are currently recognized, three of them occurring in Europe.
KEY TO THE EUROPEAN FAMILIES OP INSECTIVORA.
Dentition of a semi-crushing type : first and second
upper molars with four subequal cusps, their styles
and commissures rudimentary ; sutures in skull
persistent ; a large external pterygoid plate ; zygo-
matic arch complete, heavy. (In European species
back covered with spines.) (Hedgehogs) Erinaceidie, p. 114.
Dentition of a strictly sectorial type : first and second
upper molars with three or four cusps strongly
contrasted in size, their styles and commissures
highly developed and forming an important func-
tional part of tooth ; sutures in skull mostly dis-
appearing early in life ; no external pterygoid
plate ; zygomatic arch slender or incomplete. (Back
covered with soft, dense fur.)
Anterior lower incisor greatly elongated in axis of jaw;
mandibular articulation double ; zygoma absent ;
floor of brain-case with largo lateral vacuities ;
no auditory bulla; general form mouse-like, the
neck evident ; external ear present (Shrews) Soricidie, p. 28.
B
-a INSECTIVOEA
Anterior lower incisor not elongated in axis of jaw ;
mandibular articulation single (normal) ; zygoma
present ; floor of brain-case bony throughout ; a
small auditory bulla ; general form not mouse-
like, the neck concealed between the greatly
enlarged shoulders ; no external ear (Moles and
Desmans) Talpidie, p. 2.
Teeth in front of molars sharply differentiated by
form into incisors, canines and premolars, the
upper incisors small, sub-equal ; front feet
highly modified for burrowing, the palms
everted ; tail scarcely as long as head (Moles) Talpinx, p. :2.
Teeth in front of molars not differentiated by
form into incisors, canines and premolars, the
inner upper incisor greatly enlarged, vertical,
trenchant ; front feet not modified for burrow-
ing (habits aquatic), the palms in normal
position ; tail (in European members of the
group) longer than head and body (Desmans) Desmauinie, p. 20.
Family TALPID^.
1826. Talpidx Gray, Thomson's Annals of Philosophy, xxvi, p. 339.
Geographical (listribuiion. — North temperate portions of Old
and New "Worlds ; in Europe south to the Mediterranean coast
and west to England.
Characters. — Skull long and narrow, strongly tapering an-
teriorly, most of its sutures disappearing early in life ; zygomatic
arch complete, slender ; floor of brain-case completely ossified ;
tympanic bone attached to skull, forming a ilattened bulla ;
mandible with single articulation, the glenoid surface normal ;
no external pterygoid plate ; crowns of upper molars low, much
narrower internally than externally, the paracone and metacone
near middle of crown, the commissures and styles well developed
and forming with corresponding portions of lower teeth an
effective cutting apparatus ; body heavy, cylindrical, the short
neck concealed between the greatly developed shoulders ; eye
minute, often covered by the integument : snout much elongated,
tei'ete or depressed ; no external ear.
Beinarhs. — The members of the family Talpidie are at once
recognizable among European mammals by the great development
of the shoulder girdle and apparent absence of neck, the auditor}-
orifice seeming to lie at the shoulder. Though excessively modified
in general form the Talpidse are much less specialized than the
Soricidse in the more fundamental characters of skull and teeth.
The family is divisible into several very distinct groups or sub-
families, two of which are represented in Europe. In one of
these the animals are specially adapted to subterranean habits :
in the other they are modified for aquatic life.
Sub-Family TALPIN.-E.
Geot/raphical distribution. — Temperate portions of Europe and
Asia, from England to Jajian : in Europe south to the Meditei-
ranean coast.
Characters. — Teeth in front of molars sharply differentiated
by form into incisors, canines, and premolars, the incisors, both
above and below, small, sub-equal, chisel-shaped, the upjaer
canine large, strongly trenchant ; external form highly moditied
for subterranean life, the greatly enlarged orbicular front feet
with palms permanently turned outward, the hind feet much
smaller, not peculiar in form ; tail short ; muzzle terete.
Remarks. — This group, comjwsed of the true moles of the
Old World, and specially characterized by the relatively primi-
tive condition of the anterior teeth, is represented by four or five
genera, one of which occurs in Europe.
Genus TALPA Linn;eus.
1758. Talpa Linnaeus, Sysfc. Nat., i, 10th ed., p. 52.
Type species. — Talpa europsea Linnteus.
Geographical distribution. — Europe and western and central
Asia. Eastern limits of range not known.
Characters. — Dental formula: i 3Z3, CjEti, pm xEr^, »h 3=3 = 44.
Up23er premolars small, distinctly spaced, showing no tendency
to become imbricated. Lower canine slightly but evidently
differentiated from incisors in size and form ; auditory bulla very
slightl}^ inflated, its outlines usually indistinct, the meatus small,
sub-circular ; external form strictly talpine ; ear-conch absent ;
eye minute, often covered by the skin.
liemarks. — The genus Talpa contains half a dozen or more
species. Four of these occur in Europe, one of them generally
distributed, the three others confined to the Mediterranean
region.
KEY TO THE EUROPEAN SPECIES OF TALPA.
Greatest diameter of «i' about 4 mm. ; three lower
molars together about 8 mm. (vicinitj' of
Rome) T. romana, p. 18.
Greatest diameter of 5k' about 3 mm. ; three lower
molars together about 7 mm. or less.
. Condylobasal length of skull 33 to 37 mm. (dis-
tribution general) T. curopxa, p. 3.
Condylobasal length of skull 29 to 32 mm.
(southern).
Posterior border of anteorbital foramen over
front of last molar (Italian) T. cxca, p. 15.
Posterior border of anteorbital foramen over
middle of .second molar (Iberian) T. occidentalis, p. 15.
TALPA EUROP^A Linnaeus.
1758. [Talpa] europxa Linnseus, Syst. Nat., i, 10th ed., p. 52 (Sweden).
17G6. Taljici /n',sn(.s- P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band,
p. 36 (Ostfriesland).
1772. [Talpa] caudata Boddaert, Kortbegrip van het zaraenstel der Natuur,
1, p. 50 (Renaming of europxa).
B 2
4: INSECTIVORA
1777. [Talpa europxa] a albo-maculata Erxleben, Syst. Regni Anim., i,
p. 117 (Ostfriesland).
1785. [Talpa] vulgaris Boddaert, Elenohus Anim., i, p. 126 (Europe).
1789. [Taliya europsea] variegata Gmelin, Syst. Nat., i, 13th ed., p. 110
(Sweden).
1789. [Talpa europma'\ y alba Gmelin, Syst. Nat., i, 13tli ed., p. 110
(Sweden).
1789. Talpa europxa t cinerea Gmelin, Syst. Nat., i, 13th ed., p. 110 (Eifel,
Germany) .
1792. Talpa etirop[iea] nigra Kerr, Anim. Kingd., p. 200 (Renaming of
euro2)iea).
1797. Talpa curopxa riifa B[orkhause]n, Der Zoologe (Compendiose Biblio-
thek gemeinntitzigsten Kenntnisse fiir aUe Stande, pt. xxi),
Heft v-viii, p. 13 (Southern France).
1836. Talpa europxa flavescens Reichenbach, Pracht.-gemoinn. der
Siiugeth. des In- und Auslandes, fig. 473 (Saxony).
1852. Talpia europsea albida Reichenbach, Vollstiindigste Naturgesch. des
In- und Auslandes, iv, p. 336 (Germany).
1852. Talpa europxa lutea Reichenbach, Vollstiindigste Naturgesch. des
In- und Auslandes, iv, p. 336 (Germany).
1857. Talpa europxa Blasius, Saugethiere Deutschlands, p. 109.
1869. Talpa curopasa, flavescens Fitzinger, Sitzungsber. kais. Akad.
Wissensoh. Wien, Math.-Naturwiss. Classe, lix, pt. i, p. 400.
1869. Talpa europsea, maculata Fitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lix, pt. i, p. 401
(Renaming of albo-maculata).
1869. Talpa europxa 3 grisea Fitzinger, Sitzunsgber. kais. Akad. Wissensch.
Wien, JNIath.-Naturwiss. Classe, lix, pt. i, p. 403 (synonym of
cinerea wrongly attributed to Zimmermann, Geogr. Gesch., ii,
p. 390, 1780, where vernacular name only is used).
1897. ITaliM] scalops Schulze, Abh. u. Vortr. Gesammtb. Naturw. iv,
no. 10, p. 19. (Substitute for europxa.)
1910. Talpa europxa Trouessart, Pauue Mamm. d'Europe, p. 61.
Type locality. — Upsala, Sweden.
Geographical distribution. — Europe from Great Britain and
the Channel Islands eastward, and from the Mediterranean coast
to Scotland and central Sweden.
Diagnosis. — Size medium (hind foot about 18 mm., condylo-
basal length of skull rarely less than 33 mm.) ; skull with orbit
rather long, so that distance from posterior border of orbit to
posterior border of anteorbitai foramen is about equal to that
from latter point to front of hist j^remolar or back of canine ;
teeth of moderate size, the length of upper tooth-row (exclusive
of incisors) less than 14 mm., the greatest diameter of wi^ about
3 mm. ; mesostyle of m- and m^ entire or with apex slightly
notched.
E.itrrnal rliararters. — General form highly moditied for sub-
terranean burrowing habits, the neck so short that the conical
head with much produced snout ajapears to be joined directly,
between the very large, everted front feet, to broad, powerful
shoulders, behind which the unusually long, nearly cylindrical
body tapers gradually to somewhat weak and narrow pelvic
region. Fur dense and velvety, the hairs all of the same length,
about 12 mm. long throughout body, shorter on head. Muzzle
pad well developed, slightly wider than high, its entire margin
free, the upper edge folded back and with a deep nari-ow median
emargination, the whole surface finely and evenly rugose.
Behind pad the skin of the muzzle is naked and wrinkled along
median line above, the exact size and form of the naked area
varying in different individuals, but its length usually about
7 mm., its breadth anteriorly equal to greatest diameter of pad,
that posteriorly somewhat less. Under side of upper lip with
deep median groove extending forward to muzzle-pad. Eyes
excessively minute, less than 1 mm. in diameter, in some
individuals completely covered by the integument, in others with
a pin-hole aperture.* No external ear, the meatus about 2 • 5 mm.
in diameter. Legs so short that scarcely more than the feet
project beyond general integument of body. This is particularly
true of the front legs, which are entirely concealed to wrist.
Front feet very large, the palms orbicular, permanently turned
outward, their surface naked, finely and evenly tuberculo-reticu-
late, without trace of pads. Toes, five, each armed with a long,
slightly-curved nail broadly grooved along under surface, that of
fifth digit not so large as the others. Third digit longest, second
and fourth sub-equal and slightly shorter, first and fifth slightly
shorter than second and fourth. Hind foot short, somewhat
triangular in outline, broad through base of toes but narrowing
rapidly toward heel, the five toes with well developed, curved,
but not flattened claws, much less enlarged than those of front
foot ; second, third and fourth digits sub-equal and longest, fifth
and first successively shorter, soles naked, reticulate, with
five rudimentary tubercles at bases of digits, and a fifth neai-
middle of inner margin, its anterior extremity projecting so as to
suggest a supplemental (clawless) toe. Dorsum of pes thinly
clothed, the hairs at its edge forming a slight fringe. Tail
thickened and fleshy, about 1^ times as long as hind foot,
subterete but somewhat compressed, much constricted basally ;
scales arranged in rather irregular rings, of which there are
about twelve to the centimeter at middle ; hairs of tail sparse,
not concealing rings, sometimes forming a thin pencil. Mamma? :
/> 2 - 2 ; i2-2 = 8.
Colour. — Fur everywhere dark slaty grey, sometimes almost
l>lackish, at others more neai'ly a dark smoke-grey, the hairs
every wliere with a noticeable metallic or purplish iridescence ;
underparts sometimes a little less dark than back, and occa-
sionally with a yellowish brown sufl'usion : tail concolor with body.
Skull. — The skull is long and narrow, tapering gradually
forward from middle of brain-case, the widest region, to just
behind canines, the extreme tip of rostrum widening a little, the
zygomatic arches not standing out beyond general contour.
* JTuoh confusion has been caused by the supposition that the presence
or absence of this minute aperture is a specific or racial character.
IXSECTIVOKA
Outline when viewed from the side long wedge-shaped, rounded
oti' posteriori". Surface of skull smooth, except for a slight
sagittal crest, present in old individuals along course of parietal
suture, and a crest-like elevation extending ftirward and outward
from autero-external angle of interparietal along edge of slightly
intlated mastoid region, and terminating anteriorly in a slightly
jxiinted in'ojection. Brain-case with posterior mai'gin nearly semi-
circular to projecting points already mentioned, then abruptly
conical to back of interorbital region, its outline when viewed
from behind fusiform, a little less than half as deep as wide :
condyles slightlv projecting, but not noticeably breaking general
posterior outline. Interparietal large, strap-shaped, slightly
convex in front, similarly concave behind, the lateral extremities
squarely truncate, its antero-pos-
terior diameter about one-third
transverse diameter. Its anterior
and lateral sutures remain visible,
but the lambdoid suture is nearly
obliterated early in life, though
its position is usually marked by
a slight ridge representing the
lamlxloid crest. Base of brain-
ca.se smooth, without conspicuous
ridges, furrows or open spaces,
the bones for the most part some-
what inHated : two ill-tielined pits
in surface of basioccipital in front
of foramen magnmu ; a shallow,
broadlv triangular median furrow
between the low, flattened bulke,
the sub-circular, slightly triangu-
lar outline of which is sometimes
distinctly indicated, but more
often very obscui-e : auditory
meatus small, nearly circular.
Iinier pterygoid plate small but well developed, the hamulars
short, turned slightly outward : mesopterygoid space much
narrower posteriorly than anteriorly, its greatest breadth less
than half length, its length much more than width of palate
bctN\ ccn posterior molars, its anterior border broadly rounded :
region outside pterygoid plate inflated and with low but distinct
longitudinal ridge, convergent anteriorly with that forming edge
of mesopterygoid space. Interorbital region sulvcylindrical, dis-
tinctly expanded at middle. Rostrum narrower and somewhat
abruptly lower than interorbital region, its narrowest point just
behind canines : nares with evident posterior emargiuation.
Anteoi-bital foramen moderately large, its jxisterior border OAcr
metastyle of */('-, the plate forming outer wall of canal usually
nnich nari-ower than foramen Lachrymal foramen above anteor-
YW. 1.
Titlpii eur,tpK.t. y-.n. size.
r
bital foramen and slightly in front of its middle ; its orifice over
metastj'le of j/ji. Temporal fossa rather large, the distance from
its posterior' border to posterior border of anteorbital foramen
equal to that from latter point to front of first premolar or back
of canine. Palate moderately broad, without special peculiarities
of form, terminating posteriorly, a little behind tifi, in a slightly
raised crescentic ridge ; a vacuity about as large as the minute
incisi\e foramen on each side of palate opposite space between
//(' and »;«'-. Mandible rather slender, the ramus curved downward
near middle and upward posteriorly, the angular process nearly
on level with alveolar line. Coronoid process large, its broadly
rounded off extremity rising considerably above articular level,
its height above alveolar line about eijual to least breadth of
l)Osterior segment of mandible. Articular process slender, the
single articular surface small, normal in position. Angular
process slightly longer than articular process, and distinctly
broader, forming an obliquely-set plate directed almost horizon-
tally backward, the extremity slightly hooked upward.
Ti^eth.—In proportion to the size of the skull the teeth are
moderately large, their general
asjject noticeably trenchant. Upper
incisors simple, chisel - shaped,
IXirpendicular, forming a strongly
convex row between canines, their
height equal, but their breadth
diminishing regularly from first to
third ; crowns of first and second
usually in contact, that of third
separated by a slight space from
second and from canine.
Lower incisors projecting
somewhat obli(juelv for-
ward, essentially similar to
the upjier teeth in form,
but smaller and narrower.
Upper canine large, two-
rooted,* the height of the
shaft greater than that of an v
of the othei' teeth, and fully
e(jual to anterior breadth of
palate ; shaft wider ante-
riorly than posteriorly, with
anter-o-internal longitudinal
groo^•e, and highly deve-
loped, slightly concave pos-
terior cuttirrg edge. Lower canine very small, resembling a
fourth incisor, but with shaft conical instead of chisel-shaped,
* The roots of this tooth as well as those of the small premolars are
distinctly visible iu old individuals.
h'lO. 2.
Talpa europtea. Teeth x o.
INSECTIVORA
and set in the jaw at a slightly different angle posterior
surface of shaft with well developed longitudinal ridge. First,
second and third upper premolars two-rooted, small, their
points on level with those of incisors and inner cusps of
molars, their crowns separated from each other as well
as from canine and large premolar by narrow equal spaces ;
crowns rather higher than long, compressed, with slightly
developed posterior cutting edge, narrowly triangular in outline
when viewed from the side, the first more slender than the
others, the second and third with slight though evident postero-
external angle on cutting edge. Fourth premolar essentially
like the other three in form, but much larger and three-rooted,
the length of crown greater in proportion to height ; no secondary
cusps. Lower premolars two-rooted, similar to the upper teeth
in form, the second and third the same size as smaller upper
premolars, the first nearly as large as pm*, but narrower,
owing to absence of third root, the fourth intermediate. Upper
molars with crowns much wider externally than internally,
and completely divided into two unequal sections by a deep
longitudinal groove jjassing between protocone and bases of
paracone and metacone. Protocone large, its posterior com-
missure extending in line parallel to sagittal plane, and ending
abruptly at posterior edge of crown, near which it is slightly
thickened, its anterior commissure similar though shorter, but
usually showing some trace of thickening,* particularly in m-.
Paracone and metacone sub-equal in iir and vi^, the metacone
the larger in the former, the paracone in the latter. In «<' the
metacone is about doulile the size of paracone, and is the
largest and highest cusp of the upper molar series, its posterior
cutting edge and long commissure functioning with similarly
enlarged protoconid of m.,. Styles and outer commissures well
developed in m- and forming a distinct VV-p^-ttern ; mesostyle
entire or with apex slightly notched. In m^ the parastyle is
reduced to a minute though usually evident cusplet on the
cingulum., and the mesostyle to a thickening or angle in
commissure connecting the two main cusps ; metastyle well
developed.f In m^ the parastyle, mesostyle and their commis-
sures are well developed, metastyle and fourth commissure
absent ; mesostyle with apex usually bifid. Middle lower molar
largest, its protoconid the highest cusp in the series. First lower
molar broader posteriorly than anteriorlj-, second and third
slightly broader anteriorly than posteriorly, the two triangles
essentially alike in form. Metaconid of 7/*i low, scarcely more
* The thickenings vary considerably iu different individuals. Occasion-
ally they arc obsolete, but more frequently they are so well developed as
to form an evident protoconule and metaconule, the latter always the
larger of the two.
t In this tooth there is no anterior y, the outer surface of paracone
essentially resembling that of ^ww' except for its smaller size.
than a slight thickening of the cingulum. In the other teeth
it is a well developed cusp nearly equal to entoconid. In all
three teeth the outer cusps are noticeably higher than those of
the inner row.
Menmrements. — Average and extremes of five males from
Borrohol, Sutherland, Scotland: head and body, 148 "4 (145-
152); tail, 26-8 (26-28); hind foot, 18-8 (18-19). Average
and extremes of four females from the same locality : head and
body, 135 (133-138) ; tail, 25-3 (24-28) ; hind foot, 17-5 (17-18).
Average and extremes of five males from Solferino, Landes,
France : head and body, 138 (134-142) ; tail, 26-2 (25-28) ; hind
foot, 17-8 (17-18). Five females from the same locality: head
and bodjs 128-6 (126-133) ; tail, 27-4 (25-29) ; hind foot, 17-2
(17-18). Average and extremes of six males from Lucinges,
Haute-Savoie, France: head and body, 136-6 (132-140); tail,
24-8 (23-26); hind foot, 18-1 (17-19). Average and extremes
of three males from Turin, Italy : head and body, 138-3 (123-
147) ; tail, 31 (27-34) ; hind foot, 19-2 (18-20). Average and
extremes of eight males from the Dehesa de Valencia, Spain :
head and body, 144-3 (135-165) ; tail, 26-6 (26-28); hind foot,
18-1 (18-19). For cranial measurements see Table, p. 12.
Specimens examined. — Three hundred and ninety, from the following
localities : —
Scotland: Borrohol, Sutherland, 9 (Wilson); Black Isle, Cromarty, ;3;
Cromarty, 1 ; Gordonstown, Elgin, 1 ; Grantown-on-Sisey, Elgin, 13 (Wilson) ;
Cortach)-, Forfar, 1 (Wilson) ; Stockbriggs, Lanarkshire, 1.
England : Bowdon, Cheshire, 1 ; Altriucham, Chester, Cheshire, 1 ;
Parsop, Hereford, 1 ; Lavenham, Suffolk, 1 (Wilson) ; Barrow, Suffolk, 1
(U.S.N.M.); Arley, Staffordshire, 1 (Wilson); Rugby, Warwickshire, 2;
Warwickshire, no exact locality, 2 ; Pulbourn, Cambridge, 1 ; HoUoway,
Somersetshire, 2; Somersetshire, no exact locality, 1 ; Banstead, Surrey, 2 ;
Coonibe, Surrey, 1; Cobham, Surrey, 2; Egham, Surrey, 2; Knockholt,
Kent, 2; Bromley, Kent, 2; Devonshire, no exact locality, :3.
Denmark : No exact locality, 1 (U.S.N.M.).
Belgium: Waremrae, Li6ge, 3 (U.S.N.M.).
France : Guines, Pas-de-Calais, 1 ; Pont-de-Briques, Pas-de-Calais, 1
Trinity, Jersey, 2 ; St. Lawrence, Jersey, 1 ; Barbizon, Seine-et-Marne, 2
Melun, Seine-et-Marne, 1 (Mottaz) ; Lignieres, Charente, 1 (Mottaz)
Huelgoat, Brittany, 1 ; Cadillac-sur-Garouue, Gironde, 1 (U.S.N.M.)
Foret de Bouconne, Gers, 13 ; Solferino, Landes, 10 ; Calerille, Haute-
Garonne, 10; Legouvin, Haute-Garoune, 8; Luchon, Haute-Garonne, 3;
Bareges, Hautes-Pyr6n6es, 5; I'Hospitalet, Ariege, 1: Port6, Pyreuees-
Orientales, 3 ; St. Gilles, Gard, 1 ; Valescure, Var, 1 ; Agay, Var, 1 ;
Etupes, Doubs, 3 (Mottaz) ; Lucinges, Haute-Savoie, 9 ; Montaubau,
Haute-Savoie, 15 ; Cranves-Sales, Haute-Savoie, 12.
Spain: Pajares, Leon, 2; Castrillo de la Reina, Burgos, 9; Castafiares,
Burgos, 8; L6rida, 1; Barracas, Castellou, 24; Catarroja, Valencia, 4;
Dehesa de Valencia, Valencia, 17.
Gerjuany : Konigsberg, 4 (U.S.N.IM.) ; Moritzburg, Saxony, 8 (U.S.N.M) ;
Ummerstadt, Thiiiingen, 3; Brunswick, 15 (U.S.N.M.); Aachen, 9
(U.S.N.M.); Bremen, 1 ; Ingelheim, Rheinhessen, 4; Strass, near Burg-
heim, Bavaria, 7; Niesky, Silesia, 1; Kalbe, Saale, 2; Magdeburg,
Saxony, 1 ; Strassburg, 3.
.VusTRi A-HuNGARY : Csallokoz-Somorja, Pressburg, Hungary, 2 ; Hatszeg,
Huuyad, Transylvania, 2.
10
INSECTIVORA
RouMANiA : Bustenari, Prahova, 1 ; Comana, Vlasca, 1 ; Bucharest, 1
(Genoa) .
Bulgaria: Sofia, 1 (Andersen).
Switzerland : Geneva, 15 (U.S.N.M. and Mottaz) ; Lausanne, Vaud, 1
(U.S.N.M.) ; Les Plans, Vaud, 2 (U.S.N.M.) ; Chesieres, Vaud, 1 (Mottaz)
Andermatt, Uri, 2 (U.S.N.M.) ; Miirreu, 1 ; Tliurgau, 1 ; Oberhasli VaUey
1 ; St. Galleu, 3 ; Rheinthal, St. Gallen, 2; Degersheim, St. Gallen, 3
Gossan, St. Gallen, 3; Untervatz. Grisons, 1; Breganzona, Ticino, 1
Cortivallo, Ticino, 6 (B.M. and U.S.N.M.) ; Comano, Ticino, 2 (U.S.N.M.)
Lugano, Ticino, 4 (B.M. and U.S.N.M.); Muzzano, Ticino, 2 (B.M. and
U.S.N.M.) ; Sorengo, 1 (U.S.N.M.) ; Cremignone, Ticino, 1.
Italy : Turin, 4 (U.S.N.M.) ; Certosa di Pesio, Cuneo, 1 (Genoa) ;
Parma, 2 (U.S.N.M.) ; Gozzano, Novara, 3 (Genoa) ; Frugarolo, Ales-
sandria, 4 (Genoa) ; Vaccarezza, 1 (Genoa) ; Perti, Finalborgo, 10 (Genoa) ;
Florence, 10 (U.S.N.M.).
BemarJcs. — With the possible exception of certain bats, the
common mole shows less tendency to vary geographically than
any othei' European mammal of equally wide range.
2 6. Black Isle, Cromarty, Scot-
land.
9. Gordonstowu, Elgin.
6. Stockbriggs, Lanarkshire.
9. Parsop, Herefordshire, Eng-
land.
2 6. Rugby, Warwickshire.
2. Warwickshire.
6. Fulbourn, Cambridgeshire.
2 al. Holloway, Somerset.
2. Somerset. [Hiigcl.)
2 al. Cobham, Surrey.
1 al. 1. Eghani, Surrey.
?. Knockholt, Kent.
2 (J.al. Bromley, Kent.
3 6. Devonshire.
2 6. Trinity, Jersey, Channel
Islands. (/?. H. Bunting.)
?. St. Lawrence, Jersey.
6. Guines, Pas-de-Calais, 80 ft.
France.
i. Pont - de - Briques, Pas - de-
Calais.
6. Huelgoat, Brittany, 600 ft.
2 6,29. Foret de Bouconue, Gers,
2.50 m. {A. Robert.)
2 6,29. Solferino, Landes. (.4. Fw-
hrrt.)
2 6,2 9. Caterille, Haute - Garonne,
9U0-1000in. {A. Bobcrr.)
2 6, 9. Luchon, Haute - Garonne.
(A. Bohcrt.)
2 6,19. Bareges, Hautes-Pyr6nees.
c5. L'Hospitalet, Ariege. (A.
Bobert.)
6. Port6, Pyren6es-0rientales.
2 6 Porte, Pyren6es - Orientales,
1600-1700 m. (.1. Bobert.)
6. Valescure, Var.
W. R. Ogilvie-
Grant (p).
W. R. Ogilvie-
Grant (p).
E. R. Alston (r).
E. A. Denny (p).
El. E. Austen (p).
Tomes Collection.
H. King (p).
E. R. Alston (p).
Dr. Leach (p).
F. Heiss (p).
W. Blackwell (p).
H. E.Rawson (p).
Oxley Grabhani
0. Thomas (p).
Mrs. Power (p).
O. Thomas (c&p).
11. 1. 3. 62-G3.
11. 1. 3. 66.
79. 9. 25. 3.
11. 1. 3. 67.
11. 1. 3. 64-65.
7. 1. 1. 17-18.
11. 1. 3. 71.
58. 4. 22. 1-2.
79. 9. 25. 5-6.
58. 1. 2. 1-2.
11. 1. 3. 61.
81. 4. 2. 1-2.
11. 1. 3. 68-70.
8. 9. 2. 3-4.
8. 12. 17. 1.
94, 6. 6. 19.
O.Thomas (c&p). 98. 1. 9. 3.
0. Thomas (c & p)
O. Thomas (p).
0. Thomas (p).
0. Thomas (p).
O. Thomas (p.)
G. S. Miller ^c).
O. Thomas (p).
G. S. Miller (c).
O. Thomas (p).
92. 9. 5. 1.
G. 4. 1. 28-31.
6. 4. 1. 32-.35.
6. 4. 1. 24-27.
G. 4. 1. 21-23.
8. 8. 4. 129-132.
8. 9. 1. 38.
8. 8. 4. 133.
8. 9. 1. 36-37.
G. S. Miller (c). 8. 8. 4. 134.
TALPA 1 1
6. Agay, Var. G. S. Miller (c). 8. 8. 4. 135.
1. Montauban, Haute - Savoie, A. Eobort (c & p). 97.1.9.1.
1000 m.
3 <i, 1 9. Cranves-Sales, Haute-Savoie, A. Eobert (c & p). 5. 4. 4. 1. 5. 4,
900 m. 9. 3-5.
5 (J, 3 ?. Cranves-Sales, Haute-Savoie, 0. Thomas (p). 5. 11. 18. 1-8.
909-1200 m. France. [A.
Robert.)
6, 9. Pajares, Leon, Spain. {N. O. Thomas (p). 8. 2. 9. 26-27.
Gomalez.)
2. Castrillode la Reina, Burgos, G. S. Miller (c). 8. 8. 4. 20-21.
Spain.
2 (J, 5 9. Gastrillo de la Reina, Burgos. N. Gonzalez (c.) 8.7.7.1-7.
6. L6rida, Spain. (N. Gonzalez.) 0. Thomas (p). 8. 2. 9. 28.
6(J, 6 9. Barraoas, Castellon. {N.Gon- 0. Thomas (p). 8.2.9.29-40.
zalez.)
3 £, 9. Gatarroja, Valencia. {N. Gon- O. Thomas (p). 8. 2. 9. 12-15.
zalez.)
4 (5, 5 9. Dehesa de Valencia, Val- 0. Thomas (p). 8.2.9.17-25.
encia. (N. Gonzalez.)
3 6, 9. Ingelheim,Rheinhessen,Ger- C. Hilgert (c). 8. 11. 2. 11-14.
many.
(J, 9. Ummerstadt, Thiiringeu. Lord Lilford (p). 11.1.1.115-116.
[Schuchardt.)
6. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.137.
(Schuchardt.)
6,S9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.1-4.
(Korbitz.)
2 <i, 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.132-134.
{Kiirbifz.)
6. Niesky, Silesia. (W. Bacr.) Dr. E. Hamilton 97.12.4.18.
9. Kalbe, Saale. (TT'. Bauer.) Lord Lilford (p). 11. 1. 1. 5.
6. Kalbe, Saale. {W. Bauer.) Lord Lilford (p). 11.1.1.136.
i. Magdeburg, Saxony. (IloZ^er- Lord Lilford (p). 11.1.1.135.
storff.)
6, 9. Strassburg, Alsace. (C. Mot- 0. Thomas (p). 8. 8. 10. 10-11.
taz.)
2. Csallokoz-Somorja, Pressburg, Budapest Museum 94. 3. 1. 27-28.
Hungary. (p).
2 6. Hatszeg, Hunyad, Transyl- C. G. Dauford (c). 3.2.2.20-21.
vania, Hungary.
6. Bustenari, Prahova, Rouma- Lord Lilford (p). 4. 4. 6. 13.
nia. (W. Dodson.)
6. Comana, Vlasca, Roumania. Lord Lilford (p). 4. 4. 6. 14.
1. Miirren, Switzerland. W. Gartner (p). 92. 10. 5. 3.
9. Thurgau, 400 m. Switzerland. O. Thomas (p). 4. 4. 5. 26.
(£. H. Zollikofer.)
1. Oberhasli Valley, Switzerland. Tomes Collection. 7.1.1.130.
(Kocscrma7i.)
2 6,9 St. Gallen, 650 m. Switzer- O. Thomas (p). 4. 4. 5. 23-25.
land. {E. H. Zollikofer).
6. Breganzona, Ticino, Switzer- 0. Thomas (p). 2. 8. 4. 17.
land. (£■. H. Zollikofer).
6. Cortivallo, Ticino. (E. H. O. Thomas (p). 2. 8. 4. 15.
Zollikofer).
9. Lugano, Ticino, 300 m. O. Thomas (c & p). 2. 7. 1. 1.
6. Muzzano, Ticino. {E. H. O. Thomas (p). 2. 8. 4. 16.
Zollikofer).
9. Cremignono, Ticino. (E. H. O. Thomas (p). 2. 8. 4. 20.
Zollikofer).
12
INSECTIVOKA
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14
INSECTIVORA
1
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TALPA C.ECA Savi.
1822. Talpa cxca Savi, Nuovo Gioru. de' Letterati, Pisa, i, p. 265.
1857. Talpa csaca Blasius, Siiugethiere Deutschlands, p. 115.
1906. Talpa cxca Camerano, Boll. Mus. Zool. ed Anat. Comp. della K.
Univ. di Torino, xxi, No. 530, p. 1, June 22, 1906.
1910. Talpa cxca Trouessart, Faune INIamm. d'Europe, p. 63.
Type locality. — Mountains near Pisa, Italy.
Geographical distribution. — Mediterranean region, eastward
into Asia Minor ; details of distribution not known.
Diagnosis. — Smaller than Talpa europwa (condylobasal length
of skull, 29-32 mm. instead of 33-37 mm.) and .skull with
narrower rostrum and palate (breadth of rostrum over canines
about 4 mm. instead of 4 • 6-5 mm.) ; temporal fossa shortened, the
distance from its posterior margin to posterior margin of anteor-
bital foramen about equal to distance from latter point to second
premolar ; anteorbital foramen large, its posterior border over
front of last molar, the plate forming outer wall of canal a
slender, terete thread ; teeth small, the length of upper tooth-row
(exclusive of incisors), 11-12 mm. ; mesostyle of m- and m^ with
apex deeply notched (this character not visible in specimens with
much worn teeth).
Measurements. — Average and extremes of five specimens from
Cortivallo, Ticino, Switzerland : head and body, 133 • 8 (125-142) ;
tail, 25-2 (21-30) ; hind foot (dry), 15-6 (15-16). Average and
extremes of three specimens from Reggello, Tuscany, Italv (in
alcohol) : tail, 23-6 (22-25) ; hind foot, 14-7 (14 -4-15 -5). " For
cranial measurements see Table, p. 17.
Specimens examined. — Twenty-five, from the following localities : —
Switzerland : Agmizzo, Ticino, 1 ; Bt-llinzona, Ticino, 1 (U.S.N.M.) ;
Breganzona, Ticino, 1 (U.S.N.M.) ; Cauabbio, Ticino, 1 (U.S.N.M.) ; Corti-
vallo, Ticino, 5 (U.S.N.M.); Lonvico, Ticino, 2 (U.S.N.M.); Lugano,
Ticino, 3 (U.S.N.M.) ; Muzzano, Ticino, 1 (U.S.N.M.) ; Origlio, Ticino 1
(U.S.N.M.) ; Sorengo, Ticino, 1 (U.S.N.M.) ; Stabio, Ticino, 1.
Italy: Regello, Tuscany, 4 (U.S.N.M.); N.S. della Vittoria, Ligurian
Appenines, 1 (Genoa) ; no exact locality, 2.
6. Agmizzo, Ticino, Switzerland. 0. Thomas (p). 2. S 4 18
{E. H. Zollikofer.)
6. Stabio, Ticino. (E. H. Zol- O. Thomas (p). 2. 8. 4. 19.
likofvr.)
c5, 9 al. Italy.' Dr. Riippell (c). 45.7.22.32-3.3.
TALPA OCCIDENTALIS Cabrera.
1907. Talpa cxca occidentalis Cabrera, .•Knn. and Mag. Nat. Hist., 7th ser.,
XX, p. 212, September, 1907. Type in Cabrera collection.
1907. Talpa cxca occidentalis Cabrera, Bol. Real. Soc. Espafi. Hist. Nat.,
Madrid, vii, p. 222, October, 1907. (For date see Cabrera, Ann.
and Mag. Nat. Hist., 8th ser., I, p. 189, February, 1908.)
1910. Talpa cxca occidentalis Trouessart, Faune Mamm. d'Europe, it. 63.
Type locality. — La Granja, Province of Segovia, Spain.
Geographical distrihtiiion. — Iberian Peninsula.
16
IXSECTIVOKA
Ding)tosis. — Like Talpa cseca but skull rathtn- I'obust, the
breadth of rostrum over roots of canines usually more than
4 mm. ; anteorbital foramen contracted, its posterior border ovei'
mesostyle of vi'^, the plate forming outer wall of canal usually as
wide as transverse diameter of foramen ; crowns of molars
appreciably enlarged (length of upper tooth-row, exclusive of
incisors, frequently more than 13 mm.), but dentition otherwise
as in Tnlpa csern.
Skull. — The skull is somewhat larger and more robust than
that of Talpa cseca, a character particularly noticeable in the
greater width of rostrum and palate. In details of structure,
however, it shows no striking peculiari-
ties except in the form and position of
the anteorbital foramen. This foramen
i.s noticeably smaller than in Talpa
cseca, and its posterior boi'der lies over
middle or front of second molar instead
of over front of third. The plate form-
ing outer wall of canal is wider and less
thread-like than in TaJpa cseca. Zygoma
about as long as in Talpa cseca, but
owing to the diflerent position of pos-
tei'ior border of anteorbital foramen, the
distance from this poiiat to posterior
edge of temporal fossa equals that from foramen to canine, or
even in one exceptional instance, to outer incisor.
Teeth. — The upper molars are distinctly larger than those of
Talpa cseca, but otherwise the teeth show no peculiarities.
Measurements. — Type (from Cabrera) : head and body, 102 ;
tail, 24; hind foot, 15 "5. Average and extremes of five
specimens from La Granja, Segovia, Spain (in spirit, body
contracted): head and body, 101*6 (98-107); tail, 25-6
(25-27); hind foot, 16-4 (16-17). Adult male from Galicia :
head and body, 112; tail, 26; hind foot, 15 •4. Two males
from Cintra, Portugal : head and body, 118 and 120; tail, 26
and 26 ; hind foot, 16 and 17. For cranial measurements see
Table, p. 17.
Specimens examined. — Teu, from the following localities : —
Spain : Galicia, 1 ; La Granja, Segovia, 5.
Poetugal: Cintra, 4.
Talpa caeca (upper figure), and
T. occidentalis (lower tigure).
Nat. sizj.
lal.
Galicia, Spain.
Prof.Seoane (c&p).
94. 1. 1.
23.
2^,29al.
La Granja, Segovia,
Spain.
M.delaEscalera(c).
8. 7. 30.
19-22.
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La Granja, Segovia.
M.delaEscalera(c).
6. 11. 4
2.
2<f,lal.
Cintra, Portugal.
0. Thomas (c&p).
98. 2. 2.
8 9,58
1 skull.
59.
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INSECTn^OEA
TALPA ROMANA Thomas.
1902. Talpa romana Thomas, Ann. and Mag. Nat. Hist., 7tli ser., x, p. 517,
December, 1902. Type in British Museum.
1904. Talpa romana Camcrano, Mem. Reale Accad. Sci. di Torino, Qnd ser.,
Liv, p. 81.
1910. Talpa romana Trouessart, Faune Mamm. d'Europe, p. 64.
Type locality. — Ostia, Eome, Italy.
Geographical distribution. — Vicinity of Rome, Italy.
Diagnosis. — Externally similar to Talpa europsea. but a trifle
larger ; skull and dentition more robust than in the related
animal (greatest diameter of m^ about 4 mm. ; length of upper
tooth-row, exclusive of incisors, 14 to 15 mm.) ; mesostyle of all
three upper molars bifid at tip.
Skull and teeth. — Except that it is rather more robust the
skull does not differ appreciably from that of Talpa europsea,
though posterior base of zygoma appears to be usually situated
somewhat further back. Teeth as in Talpa europsea, but larger
throughout, a diffei'ence particularly noticeable in the first upper
molar and in the large lower cheek-teeth (the combined length of
Fig. 4.
Crown of niol.ars in Talpa europsea (a), and T. romana (b).
these four teeth about 10 mm. instead of about 8 mm.). In form
the teeth are .similar to those of the related animal, but the
mesostyle of m^ is relatively as well as actually larger, and its tip
is distinctly bifid, though not so deeply as in the succeeding
teeth. Cingulum between outer bases of main cusps of lower
molars better develojied than in T. europsea, its edge frequently
forming an evident cusp, especially in m.y
Measurements. — Extei'nal measurements of type (from
Thomas) : head and body, 126 ; tail, 29 ; hind foot, 19 (all
measurements from skin). Four males from the vicinity of
Rome (in alcohol) : head and body, 138-3 (130-14-5) ; tail, 28-6
(27-30) ; hind foot, I'J -6 (19-20). Three females from the same
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INSECTIVOEA
locality: head and body, 127-5 (125-130) ; tail, 29-5 (29-30);
hind foot, 18-4 (18-18'6). For cranial measurements see Table,
p. 19.
Specimens examined. — Fourteen, from the vicinity of Rome (B3I. and
U.S.N.M.) ; also about fifty from the same region in Turin Museum.
Remarks. — Talpa romana is a well characterized species,
readily distinguished from other European moles by its unusually
lai-ge teeth.
1. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 8.
{Type of species.)
1. Frascati, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 9.
S, ? al. Rome. Genoa Museum (e). 3. 1. 31. 1-2.
2 6. Rome. [C. Coli.) G. Barrett-Hamilton (p). 11.1.2.1-2.
Sub-Family DESMANIN.'E.
Geographical dii<trihution. — South-western France and northern
half of Iberian Peninsula ; eastern Russia and western Siberia.
Characters. — Teeth in front of molars not differentiated by
form into incisors, canines, and premolars ; anterior upper
incisor greatly enlarged, canine-like, directed downward, its
outer edge highly trenchant, the two anterior lower incisors
slightly elongated, projecting obliquely forward ; external form
less evidently mole-like than in the Talpime, modified for
aquatic life, the much enlarged hind feet with completely webbed
toes, the front feet smaller, not peculiar in form ; tail long ;
muzzle depressed.
SemnrJcs. — The sub-family Desmanlnx contains the genera
Desmana of Russia and .Siberia, and GaJeinys of the Iberian
Peninsula and south-western France, strictly aquatic animals
not distantly related to the American moles. The anterior
teeth present a much higher degree of specialization than that
met with in the TaJphiie.
Genus GALEMYS Kaup.
1829. Galemys Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 118 [Mygale pyrcnaica Geoffroy).
184G. Galoniys Agassiz, Nom. Zool., Index Univ., p. 149 (Emendation of
Galemys) .
1849. Mygalina I. Geofiroy, D'Orbigny's Diet. Univ. d'Hist. Nat., iv, p. 709
(Mygale pyrenaica Geoffroy).
Type species. — Mygale pyrenaica< Geofiroy.
Geographical distribution. — Northern half of Iberian Peninsula,
and Pyrenean region of south-western France.
Characters. — Tail flattened laterally at distal extremity, else-
where terete ; unicuspid teeth slender, their width conspicuously
less than height of crown ; main cusp of large premolar trenchant
anteriorly ; .brain-case without unusual ridges.
GALEMYS 21
Remarks. — The genus Galemys is readily distinguishable from
the Russian and Siberian Dcsmana* in which the tail is flattened
laterally throughout, the unicuspid teeth are low and thick, their
width about equal to height of crown, the main cusp of large
premolar is rounded anteriorly, and the ridges on brain-case are
unusually developed. It contains a single species, peculiar to
south-western Europe.
GALEMYS PYRENAICUS Geoffroy.
(Synonymy under subspecies.)
Geographical disti-ibnfion. — South-western France (Pyrenees
and their immediate neighbourhood) and northern half of Iberian
Peninsula.
Diagnosis. — General characters as in the genus ; head and
body, 110 to 135; tail, 130 to 155; hind foot, 32-5 to 38;
condylobasal length of skull, 33 to 35-5 ; mandible, 22 to 24.
External characters. — Form somewhat intermediate between
that of a mole and rat, the body less elongated than in Talpa,
but the neck almost equally short (so that auditory oritice
appears to be at shoulder) and the muzzle similarly produced ;
legs less shortened than in the moles and front feet not
specially enlarged ; tail rat-like, longer than head and body.
Fur less dei:ise and velvety than in the moles, and of the peculiar
quality characteristic of acjuatic mammals, the hairs of two
kinds, the longer, coarser ones about 12 mm. in length, those of
shorter under fur about half as long. Head conical, as in the
moles, but the much elongated snout (length about 20 mm. from
incisors) greatly flattened instead of terete, its breadth at middle
about 7 mm., dejDth in same region 2 mm. At tip the muzzle
broadens rather abruptly to about 10 mm., the broadened
portion divided by shallow notch at middle of anterior border
into two slightly indicated lobes ; at middle of each lobe and
about 1 mm. back from anterior edge is situated one of
the rather large, transversely elongated nostrils. | The median
* The synonymy of this genus is as follows : —
1777. Dcsmana Gueldenstaedt, " Beschiift. Berliner Gesellsch. Naturforsch.
Freunde, iii, p. 108." (Castor moscliatus Linnaeus.)
1799. Desman Lacepede, Tabl. Mamm., p. 7. Same type.
1800. ikfyjaZc Cuvier, Lee. d'Anat. Comp., I, Tabl. 1. Same type. (Described
in Tabl. E16m. d'Hist. Nat. des Anim., p. 109.)
1815. Desmanus llafinesque, Analyse de la Nature, p. 59 (Emendation of
Desman).
1829. Mi/ogalea Fischer, Synops. ^lamm., p. 250 (Substitute for Mijgalr).
1830. Caprios Wagler, Nat. Syst. Amphibien, p. 14 (Substitute for Mi/galc).
1836. Myogale Brandt, Wiegmauu's Archiv fiir Naturgesch., 1S34, i, p. 17(3.
t The actual narial opening is situated at extreme inner portion of
nostril, and is capable of complete closure bj- the combined action of
antoro-interual border of nostril, a narrow semilunar membrane at upper
edge of inner narial aperture, and a large wart-like thickening of upper
border of nostril.
22
INSECTIVORA
notch is continued backward on upper surface as a narrow
groove still further dividing the two lobes ; on under side
it is continuous with a similar though much longer groove
extending to middle of upper lip. Each lobe is further
marked by a narrow groove extending backward and slightly
inward from near outer edge of nostril. Surface of lobes
very iinely rugose and pitted, that of rest of muzzle coarsely
rugose and without pits. Behind lobes the muzzle is naked
along median region, thinly haired at sides and beneath.
Eye minute, essentially as in Taipa, though probably never
covered by the integument. No external ear, the meatus about
4 mm. in diameter. Front feet rather large and broad, readil}'
turned outward but not perma-
nently in this position, the five
short fingers joined by a narrow
web, and armed with strong,
slightly curved claws 4 to 5 mm.
in length ; fourth digit longest,
tifth and third sub-equal and
slightly shorter, second and first
still shorter. Palms naked, their
surface finely tuberculo - rugo.se^
without trace of larger tubercles,
though the surface is marked
by three deep wrinkles ; balls of
digits projecting conspicuously
beneath bases of claws. Dorsal
surface of fi'ont foot covered
with minute hairs, these lengthen-
ing along edges to form distinct
fringes. Hind foot much larger
than front foot, the toes webbed
to base of claws, the claws
similar to those on front foot
but lai'ger ; fourth digit longest,
third, fifth, second and first suc-
cessively shorter, tlie first extending nearly to end of first
phalanx of second ; surface of sole hke that of palm, the
three longitudinal wrinkles at bases of digits well developed,
the large inner tubercle present in the moles very slightly
developed, its extremity not projecting like a supplemental
digit. Upper surface of hind foot naked, somewhat more
coarsely tuberculate than sole. A fringe of stiffened hairs
along outer edge of outer toe and continuing along foot
nearly to heel. Tail longer than head and body, terete except
at tip, where it is flattened laterally. Scales arranged in some-
what irregular rings, of which there are about nine to the
centimeter at middle of tail. Hairs of tail short, not conceal-
ing scales except on flattened terminal portion, where they form
}"lfi. 5.
Gdlemys i->ijreiiaicits
GALEMYS 23
a rudimentary dorsal and ventral keel. Mamniiv :pl — l,al — 1,
i2-2 = 8.
Colour. — Back and sides dark brown varying somewhat in
exact shade, but never a distinct slaty as in Talpa, the longer
hairs lighter than the under fur and sometimes producing a
slight eliect of coarse " lining," particularly on posterior third
of back and along sides. Under parts buffy in rather strong
contrast with back, but without true line of demarcation.
Hairs of tail and feet buffy. Claws whitish.
Skull. — In general the skull resembles that of Talpa europsea,
but the brain-case is shorter and squarel)' truncate jwsteriorly,
the interorbital region is shorter and narrowed instead of
widened at middle, and the rostrum is longer. Surface of skull
smooth except for the same ridges as in Talpa, those at sides of
brain-case not unusually developed. Brain-case decidedly moi-e
than half as deep as wide, its outline when viewed from behind
vaguely pentagonal, its posterior margin nearly straight, though
with slight median swelling ; condyles not projecting posteriori}^,
completely hidden when viewed from above. Interparietal
projecting further forward than in Talpa, its antero-posterior
diameter nearly equal to its width. Base of brain-case with
deep but broad median furrow, the surface of the bones more
angular and less inflated than in Talpja ; no pits in basioccipital
in front of foramen magnum. Bullse low and flattened, less
perfectly formed than in Talpa, the tympanic bone annular and
retaining its distinctness, though joined with surrounding parts ;
meatus large, occupying about one-half surface of bulla.
Mesopterygoid space short, its length scarcely equal to width of
palate between posterior molars, its width about one-third length,
its anterior border double rounded, encroached on by slight
median spine. Outer pterygoid plate i-educed to a small but
evident ridge. Interorbital region hour-glass shaped, widening
more rapidly posteriorly than anteriorly, its narrowest region
slightly behind middle. Zygoma straight, flattened posteriorly,
compressed anteriorly, its length (measured from posterior border
of orbit to posterior border of anteorbital foramen) equal to
distance from posterior border of anteorbital foramen to front of
canine. Rostrum about as wide as in Talpa, but relatively
longer, the distance from posterior border of anteorbital foramen
to gnathion about equal to greatest breadth of brain-case instead
of much less, its dorsal surface on level with that of interorbital
region, its outer margins nearly parallel to the squarely truncate
anterior extremity. Xares scarcely emarginate posteriorly.
Anterior portion of border of alveolus of large incisor distinctly
thickened, the thickened region terminating laterally in a small
but evident wart-like nodule. Lachrymal foramen over middle
of anteorbital foramen and midille of m^. Posterior border of
anteorbital foramen o^er parastyle of tir. Palate essentially as
in Talpa, but vacuities smaller and incisive foramina large, their
24
IXSECTIVORA
longitudinal diameter about equal to width of palate in same
region. Posterior palatal ridge much as in TaJpa, but the
extremities produced as distinct backward-curved processes.
Mandible lather robust, the ramus neai'ly straight, the angular
I^rocess much below alveolar line Coronoid process high and
narrow, slightly recurved at tip, its height above alveolar line
considerabl)' greater than least breadth of posterior segment of
mandible. Articular process short and robust, the single
articular surface rather large, normal in position. Angular
process longer than articular process, its form essentially as
in Talpa.
Teeth. — Dentition relatively heavier than in Talpa euro-psea,
the teeth, with exception of anterior upper incisors, less trenchant
in general aspect. An-
terior upper incisor much
the largest of all the
teeth, the two together
closing entire front of
palate ; shaft triangular
in cross section, and all
three faces sub-triangular
in outline, the posterior
and antero-external faces
widest, the former slightly
concave, the latter
slightly convex, the two
forming a perpendicular
external cutting edge
about 4 mm. in length
along their line of con-
tact ; a much shorter but
well developed cutting
edge along line of contact
of posterior and antero-
internal faces, extending
from acutely triangular-
pointed apex of tooth to
point of contact with
tooth of opposite side,
a distance of about
■ 6 mm. ; line of contact
between antero-external
and anterointernal faces
marked by a slight thougli evident ridge ; height of shaft
about equal to width of palate ; tirst and second lower incisors
small, chisel-shaped, strongly imbricated, the second about
twice as large as first, their shafts directed forward in line
with upper portion of symphysis menti, the tips of the four
teeth together forminsc a straio;ht transverse cutting edge which
Fig. 6.
Galemys j'urenaicus.
I
acts iu oi^position to combined posterior surface of large ujjper
incisors. Upper unicuspids forming an unbroken row continuous
posteriorly with series of cheek teeth, but separated anteriorly
from large incisoi' by distinct space into which the apex of
second lower incisor tits when jaws are closed. Two anterior
unicuspids (?"" and v') minute, terete, single-rooted, their axes
directed backward and falling in same line with those of two
anterior lower unicuspids when jaws are closed. Third unicuspid
(canine) two-rooted, its crown perpendicular, compressed, larger
than those of first and second combined, and distinctly greater
in height. Fourth unicuspid (pm^) single-rooted, subterete,
scarcely larger than second. Fifth and sixth unicuspids (pnr
and 2>m^) essentially like canine, their crowns distinctly higher
than wide, compressed obliquely to the tooth-row, with slightly
developed anterior and posterior cutting edge. Height of third
and fifth sub-equal, greater than in the others, their tips about
on level with main cusps of molars ; sixth lower than fifth but
with crown longer and posterior ridge better developed. Lower
unicuspids not unlike jj?»^ and jv?«^ but with crowns lower and
longer, slanting a little forward, each with a faintly developed
antero-internal lobule. These teeth are slightl}' imbricated and
their form approximates that of the unicuspids of the Soricidie.
First and second {i^ and ,) higher than third, their form suggest-
ing that of anterior lower incisor, all three single-rooted.
Fourth larger than any of the first three, obscurely two-rooted ;
fifth slightly smaller, single-rooted ; sixth (pm^) largest of the
series, distinctly two-rooted, its cusp nearly on level with
main cusps of molars, its an tei'O external lobule (rudiment of
parastyle) more evident than in the others. Large upper pre-
molar three-rooted, its crown area about equal to that of third
molar, its main cusp with well de^ eloped anterior and posterior
cutting edges, its antero-internal cusp small but evident, its
posterointernal cusp about equal to protocone of m^. Upper
molars with crowns wider and less oblique than in Talpa
europsea, and main cusps not so high. Transverse groove
between bases of main cusps converted into a median pit by
better development of commissures of protocone and larger size
of protoconule and metaconule. Paracone and metacone about
equal in height, the latter slightly the more robust. Styles well
developed, except the reduced parastyle of m^, the mesostyle in
each tooth completely divided into two cusps, the W-pattern thus
changed into two V-shaped figures. Third upper molar with
crown area about two-thirds that of second, its metastyle and
fourth commissure absent. Lower molars essentially as in
TaJpn etirop»a, but contrast in height of outer and innei'
cusps very slight.
26
INSECTIVORA
Galemys pyrenaicus pyrenaicus Geoffroy.
1811. Mijgale pyrenaica GeoSroy, Ann. Mus. d'Hist. Nat., Paris, xvii,
p. 193.
1910. Mijogale pyrenaica Trouessart, Faune Mamm. d'Europe, p. 60.
Type localitj/. — Near Tarbes, Hautes-Pyrenees, France.
Geogrnphical distrihtition. — Pyrenees and adjacent portion of
southern France ; probably also north-eastern Spain to the
Ebro ; Asturias ?
Diagnosis. — Hind foot, 32 -4 to 3-1: '6 mm.; condylobasal
length of skull about 33 to 34 mm.
Colour. — Back and sides intermediate between prouts-bi'own
and seal-brown, the longer hairs not so dark as under fur and
with a conspicuous lustre ; a small ill-defined buffy area around
eye. Underparts varying from ochraceous-buff to a dull light
cream-buli', clouded by slaty under colour. Front feet dull
ochraceous-buft' tinged with dark brown. Hairs of tail and
fringe on hind foot light buffy.
Measurements. — Two males from Ax-les-Thermes, Ariege,
France: head and bodj^, 115 and 130; tail, 134 and 137 ; hind
foot, 34 "6 and 34. Average of three females from the same
locality: head and body, 117 (110-133) ; tail, 131-6 (126-137) ;
hind foot, 32 '9 (32-4-34). For cranial measurements see Table,
p. 27.
Specimens examined. — Fifteen, from the following localities : —
FiiAXCE : Ax-les-Thermes, Ariege, 5 ; Pyrenees, no exact locality, 8
(B.M. and U.S.N.M.).
Spain : Pajdres, Leon, 2.
2 al. Ax-les-Thermes, Ariege, Toulouse Museum 1.7.27.1-2.
720 m. France. (p).
<5, ?. Ax-les-Thermes, Ariege. V. Bullies (p). 8. 3. 27. 4-5.
2. Pyrenees. (Yeircaiix.) Tomes Collection. 7. 1. 1. 15-lC.
1. Unknown. F.Maxwell Lyte (p). 62.1.13.2.
1 al. Pyrenees. (No history.)
1. Pyrenees. Purchased (Parzu- 41. 918.
daki).
1. Pyrenees. Dr. .J. E. Grav (p). 43. 10. 14. 1.
6, ?. Pajares, Leon, Spain. O. Thomas (p). 8. 2. 9. 47-48.
(N. Gonzalez.)
Galemys pyrenaicus rufulus Uracils.
1897. Myogalea riifida Graells, ilom. Real. Acad. Sci., Madrid, xvii, p. 460.
1910. Myogale pyrenaica rufula Trouessart, Faune Mamm. d'Europe, p. 61.
Tyle locality. — Rio Balsain, above the Venta de los Mosquitos,
Sierra de Guadarrama, Segovia, Spain.
Geograpliical distribution. — Central Spain, south of the Ebro
Valley.
Diagnosis. — Hind foot, 36 to ofi mm. ; condylobasal length
27
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28
INSECTIVORA
of skull iibout 34' 5 to 35 • 5 mm. ; colour apparently not so dark
as in the Pyrenean race.
Colour. — Upper parts essentially as in G . pyrenaicus pyrenaicus,
but slightly less dark, the general hue somewhat leaden. Longer
hairs on rump distinctly buffy. Pale area around eye larger and
more noticeable than in the p3'renean foi'ui.
Measurcvienfs. — Three males from Silos, Burgos, Spain : head
and body, 123, 131 and 134 ; tail, 135, 145 and 156 ; hind foot,
36, 38 and 38. For cranial measurements see Table, p. 27.
Specimens examined. — Six, five froin Silos, x'rovince of Burgos, Spain,
and one from Buitrago, province of Madrid (U.S.N.M.).
Hemarhs. — The central Spanish form of Galemys appears to
be well differentiated from true pyrenaicus by its greater size,
a character which is particularly noticeable in the larger,
more massive skull. Two specimens from the Asturias (Xos. 8.
2. 9. 47-48, Pajares, Leon, N. Gonzalez, collector) are apparently
identical with the Pyrenean animal.
When in the water this animal shows much less agility
than the water-rat and watei'-shi-ew, probably because, though in
appearance the most perfectly adapted of the three to aquatic
life, it retains too much of its Talpine inheritance of shortness of
limb and heaviness of gener'al form to be an active swimmer.
Its defective vision, inherited from the same source, would also
tend to a like result.
2 ? sks. Silos, Burgos ; Spain. Eev. S. Gonzalez (c). 8. 7. 7. 8-9.
6, 2 sks. Silos. G. S. Miller (c). 8. 8. 4. 43-45.
Family SORICIDyE.
1821. Soricidx Gray, London Med. Eepos., xv, p. 300, April 1, 1821.
Geographical distribution. — Throughout tropical and temperate
Africa, Europe, Asia (including the Malay Archipelago), North
America, and the extreme northern portion of South America.
Characters. — Skull long and narrow, strongly tapering an-
teriorly', most of the sutures disappearing early in life ; zygomatic
arch incomplete, represented by a slight though usually evident
I'udiment of the zygomatic ^^rocess of maxillary ; floor of brain-
case with median longitudinal bridge of bone and wide lateral
fenestrate area on each side, in which auditory parts are sus-
pended ; tympanic bone annular-, not attached to skull ; basi-
sphenoid without auditory process ; no external pterygoid plate :
mandible with complete double articulation ; anterior teeth not
differentiated by form into incisors, canines and premolars, the
first upper incisor very large, strongly projecting forward, its
tip hooked downward, its base with a secondary lobe, the anterior
lower incisor nearly straight, much i)roduced in axis of mandible,
the other anterior teeth foi'ming a series of small " unicuspids,'
differing from each other chiefly in size : crowns of upper molars
SOREX 29
low, sub-(iuadi'ate in outline (except the much reduced third), the
paracone and metacone near middle of crown, the styles and
commissures well developed and forming an important functional
part of the cutting apparatus ; form mouse-like, but snout always
pointed and much produced beyond incisors, eyes small, and ears
often partly or entirely hidden in the fur.
Remarks. — The members of the family Soricidse are at once
recognizable among European Insectivora by their mouse-like
form, small eyes, and sharply pointed muzzle. They are all of
small size, the largest (Neomys fodiens) not so large as a house-
mouse, while the smallest {Pachyura etrusca) is one of the least
of known mammals. Notwithstanding their manifestly primitive
general structure, the Soricidse present a very high degree of
specialization in the form of the anterior teeth, the absence of
the zygoma, and the remarkable double articulation of the jaw.
About fifteen genera are known. Four of these are represented
in Europe.
KEY TO THE EUROPEAN GENERA OF SOBICIDjE.
Posterior lower molar with five cusps ; teeth pigmented
at tips ; tail without sprinkliug of elongated hairs.
Upper unicuspid teeth 6-5 ; cutting edge of anterior
lower incisor with more than one lobe ; feet not
fringed Sorex, p. 29.
Upper unicuspid teeth 4-4 ; cutting edge of anterior
lower incisor with one lobe; feet fringed. (Water
Shrews) Neomys, p. 65.
Posterior lower molar with four cusps ; teeth white
throughout ; tail with noticeable sprinkliug of
elongated hairs.
Upper unicuspid teeth 4-4 Pachyura, p. 81.
Upper unicuspid teeth 3-3 Crocidura, p. 86.
Genus SOREX Linn.'eus.
1758. Sorex Linnseus, Syst. Nat., i, 10th ed., p. 53.
1829. Oxyrhin Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 119 (Included the undeterminable Sorcx constrictus Hermann
and S. tctragonnriis Hermann ; the latter may be chosen as type).
1835. Amphisorcx Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,
II, p. 28 (hcrmanni = Neo7nys fodiens skull -f Sorex araneus
tetragomrus animal) Part.
1838. C&rsira Gray, Proc. Zool. Soc. London, 1837, p. 123. June 14, 1837
(vulgaris = araneus).
1842. Otisorex De Kay, Zool. of New York, i, Mamm., p. 22 (j)latyr]iinus
= personafus).
1857. Sorcx Blasius, Siiugethiere Deutschlands, p. 124.
1890. Homalurus Schulze, Schriften Naturwiss. Vereins Harzes in Werni-
gerode, v, p. 28 (nlpinus).
Type species. - Sorex araneus Linnanis.
Geographical distribution. — Northern portion of both hemi-
spheres ; in Europe west to Ireland and south to central Spain
and southern Italy.
30
INSECTIVOEA
Characters. — Ujjper unicuspid teeth 5-5 (dental formula :
i ?=?, cti ^Jm'^, iii'^ = 32) ; posterior lobe of anterior upper
incisor fully half as high as main cusp ; anterior lower incisor
with three well developed lobes on cutting edge ; third lower
molar with hypoconid and entoconid small but distinct, so that
the form of the tooth differs from that of first and second molars
in the reduced size of the second triangle only, its crown, like the
others, 5-cusped ; second lower unicuspid with rudimentary
second cusp and commissure ; points of all the teeth pigmented
(the coloured portion wearing away in extreme old age) ; skull
lightly built, with slender weak rostrum and abruptly wider
brain-case ; rudimentary zygomatic process of maxillary evident ;
no special modifications in external form ; tail covered with
hairs of uniform length (except that those of pencil are elon-
gated) ; ear nearly concealed by the fur, the meatus closed by
a large valvular outgrowth from the antitragus supplemented
by a fold on inner surface of conch ; habits terrestrial.
Bemarhs. — This is the most widely distributed genus of
Insectivora. It contains about sixty described forms, fourteen
of svhich occur in Europe.
KEY TO THE EUROPEAN FORMS OP SOBEX.
Anterior mandibular incisor with low, sometimes ill-
defined lobes on cutting edge ; first lower uni-
cuspid two-pointed; lachrymal foramen over
point of contact between m' and m- ; tail about
as long as head and body ; colour uniform darlj
slaty grey. (Alpine Shrews) .S'. alpinus, p. 60.
Condylobasal length of sliuU 19-4 to 20-6 mm.;
upper tooth-row 8 • 6 to 9 ■ mm. (Alps ; east-
ward to Transylvania) S. a. alpinus, p. 62.
Condylobasal length of skull 19 to 19 -6 mm. ; upper
tooth-row 8-2 to 8'6 mm. (Harz Mountains
and Riesengebirge) S. a. hercynicus, p. 63.
Anterior mandiljular incisor with high, distinct lobes
on cutting edge ; first lower unicuspid single-
pointed ; lachrymal foramen in front of point of
contact between ?»' and m" ; tail shorter than
head and body ; colour never uniform dark slaty
grey, the under parts usually much lighter than
back.
First, second and third upper unicuspids sub-equal;
condylobasal length of skull 14 '8 to 16 ■6 mm.;
head and body usually about 60 to 60 mm.
(Pigmy Shrews) S. minutus, p. 53.
Molars and anterior upper incisor normal (Dis-
tribution general) S. m. minutuK, p. 55.
Molars and anterior upper incisor enlarged
(Southern Italy) ,S'. m. hicanius, p. 60.
First and second upper unicuspid much larger than
third; condylobasal length of skull 17-4 to
20 mm. ; head and body usually about 65 to
80 mm. (Common Shrews) S. arancus, p. .31.
SOREX 31
Palate broad anterioi'ly, its width at level of first
unicuspid nearly equal to that of tooth.
Condylobasal length of skull 18--1 to 19-2 mm.;
anterior toeth enlarged (Island of Jersey). . . S. a. fretalis, p. 45.
Condylobasal length of skull about 17 • 5 mm. ;
anterior teeth not enlarged (Jlountains of
central Spain) S. a. granarius, p. 52.
Palate narrow anteriorly, its width at level of
first unicuspid barely more than half that
of tooth.
Condylobasal length of skull usually 19 to 20
mm.; hind foot usually 13 to 14 "4 mm.
(mountain and northern forms).
Hind foot 13-6 to 14*4 mm.; colour in
summer pelage very dark, the tricolor
pattern usually conspicuous (South-
western Norway) S. a. bergensis, p. 41.
Hind foot 13 • to 14 mm. ; colour in summer
pelage frequently light and brownish.
Back frequently blackish in summer pelage
(Alps and neighbouring regions) S. a. tetra^ontirus, -p. 4:2.
Back rarely if ever blackish in summer
pelage (Pyrenees) S. a. pyrenaicus, p. 44.
Condylobasal length of skull usually 17 '4 to
19 mm. ; hind foot usually 11 to 13 mm.
(lowland forms).
Underparts blackish, not contrasted with
back (Charente, Prance) S. a. aantonus, p. 40.
Underparts greyish or brownish, contrasted
with back except when latter is also
brown.
Colour of sides not distinctly contrasted
with that of back ; belly heavily
washed with wood-brown (Plains of
south-western France) S. a. euronotus, p. 41.
Colour of sides usually forming distinct
contrast with that of back ; belly
lightly washed with wood-brown.
Average colour darker, the back ranging
from bister to a deep blackish
brown (Central Europe and Scan-
dinavia, except south - western
Norway) ,S'. a. araneus, p. 35.
Average colour less dark, the back
ranging from hair-brown tinged
with bister to seal-brown (Great
Britain) : s. a. castaneus, p. 37.
SOREX ARANEUS Linnajus.
(Synonymy under subspecies.)
Geographical distribution. — Northern portions of Europe and
Asia from England and Scotland eastward ; exact limits of
range unknown ; in western Europe south to central Spain, and
central Italv, north to northern Scandinavia.
Diaijnosia. — Size medium, head and body usually about
65-80 mm, the tail 20-35 mm. shorter, condylobasal length of
skull, 17 •8-20 mm.; posterior lobe of anterior upper incisor
32
IXSECTIVORA
compressed latei'all\', the length of its base equal to that of base
of anterior cusp ; colour brown or blackish, the underparts
never as dark as back (usually much lighter), and the sides often
contrasted with both.
External characters. — Fur moderately dense, its depth at
middle of back about -i mm. in summer, 8 mm. in winter, its
texture not specially modified ; no elongated hairs on Hanks and
across rump. Eyes small and inconspicuous ; ears nearly
concealed in the fur. Feet not peculiar in form, thinly clothed
with very fine hairs on dorsal sui-face ; fore foot with third and
fourth digits sub-equal and longest, fifth extending just beyond
base of fourth, first barely reaching base of second, the claws
small but well developed ; palm so conspicuously rugose reticulate
that tubercles are not very distinct ; tubercles 6, sub-equal, the
three at bases of main digits well-defined, that at base of thumb
sometimes confluent with that at inner side of wrist (so that the
number is ajjparently reduced to 5), the two wrist-pads separated
from each other in median line by a noticeable space ; extreme
posterior edge of palm covered with ordinary integument. Hind
foot with third and fourth digits sub-equal and longest, second
slightly shorter, fifth reaching base of fourth, and first extending
to base of fifth ; surface of sole as in palm, but tubercles better
defined, four at bases of digits and two situated more posteriorly,
all six about equal in size ; sole finely haired fi-om hinder
tubercles to heel, the middle of which is bare ; claws like those
of fore-foot. Tail terete or somewhat 4-sided, rather more than
half as long as head and body, its hairs minute, ratlier closely
appressed, and nearly concealing the annulation ; pencil usually
well developed, 4-6 mm. in length, but occasionallj^ in aged
individuals much reduced or absent, together with the rest of
the hairy covering of the tail. Caudal annulations rather
indistinct, about 24 to the centimeter at middle. jNIammfe :
/ 3-3 = 6.
Colour. — Dorsal area, extending from base of tail to crown
cheeks and muzzle, brown, the exact shade varying much both
seasonally, racially and individually, but the normal extremes
falling between hair-brown or light bister and a very dark,
blackish seal-brown. Sides wood-brown, usually forming an
evident contrast with dorsal area, though this contrast is
occasionally inconspicuous in dull light specimens in summer
coat, or wholly obliterated in the general darkening of entire
animal in S. araneus sanfonus. Underparts smoky grey washed
with wood-brown, or occasionally sufiused with slaty (particularly
in the dark »S'. a. santonus). Between colour of back and sides
there is usually an evident line of demarcation : between sides and
belly the contrast is less marked and the transition less abrupt.
Feet a dull indefinite light brown, often with a faint dark shade
along outer edge. Tail dark brown above and at tip, light brown
below, especially near base, sometimes bicolor throughout.
SOREX 33
While there is no invariable rule, the colour in winter is
usually darker than in summer, and the tricolor pattern of
dark back, yellowish brown sides and greyish belly is more
pronounced. A trace of tliis pattern is often the most convenient
character by which to recognize shrunken ill-prepared specimens,
which might otherwise be mistaken for Sorex minutus.
SJcuIl~The skull is slender and lightly built, with no special
peculiarities of form as compared with that of other shrews
Brain-case well marked off from interorbital region, its surface
smooth except in extreme old age, its main sutures remainincr
open until late in life. It is sub-circular in °
general outline when viewed from above,
but with antero-external portion of border
noticeably flattened, so that at point of
greatest breadth there is usually an evident
angle ; condyles scarcely visible, causing no
break in posterior outline. Depth of brain-
case at middle slightly more than half
greatest width ; no sagittal crest except in
extreme old age; lambdoid crest at tirst
confined to lateral portions of occiput, rarely
extending to median line. Dorsal profile
usually with evident concavity in inter- ^^''" ""
orbital region (more marked than in Sorex ^"wat S!"'"
minutus and ,S'. alpiniis). Nares broadly
rounded posteriorly, the lateral margin obtusely angled near
middle. Anteorbital foramen moderately large, not very con-
spicuous when skull is viewed from in front. Lachrymal
foramen over middle of m\ Mesopterygoid space nearly parallel-
sided, less than half as wide as long.
Teeth. — Anterior upper incisor with basal lobe relatively
larger than in any other European shrew, the length of its
base nearly equal to diameter of anterior lobe at level of angle
between the two cusps. When tooth is viewed from below the
posterior lobe appears nearly as large as anterior cusp ; in
lateral view it approximates the size and form of first and
second unicuspid, and in height falls a little short of anterior
cusp. The two teeth come in contact anteriorly slightly below
tips, which do not diverge conspicuously. Anterior lower incisor
robust, its shaft very slightly tapering, its cutting edge with
three well-developed lobes, the lengths of bases of which
fhminish regularly from first to third, the first lobe more
distinctly separated from succeeding lobe than from anterior
point of tooth. Upper unicuspids robust, their crowns squarish
in outline when viewed from below, bluntly triangular when
viewed from the side ; height about e.jual to length ; anterior
and posterior borders straight or faintly concave, upper border
convex, more strongly posteriorly than anteriorly. The highest
point of cusp lies slightly in front of middle of crown. Upper
D
34
INSECTIVORA
riG. 8.
Siirex araneits: Anteriur
teeth in prolile. X 5.
and posterior borders sub-equal, slightly longer than anterior
border. Cusp rounded on antero-external face, squarely truncate
postero-internally atong line connecting antero-internal and
postero-external corners of crown. The
region behind this line is occupied by a
flattened or somewhat concave, nearly
horizontal crushing surface opposed
during mastication to second and third
lobes of lower incisor (1st and 2nd upper
unicuspids) and to points of the two
lower unicuspids (3rd and 4th upper
unicuspids). In size the first and second
unicuspids are sub-equal and decidedly
larger than third, which in turn some-
what exceeds fourth. Fifth still smaller
than fourth and closely crowded between it and antero-external
cusp of large premolar, its cusp relatively lower than in the
other teeth of the series. First lower iniicvispid essentially
similar to i3rst and second upper,
but crown longer than broad and
without distinct crushing surface.
Second larger than first, not
conspicuously different from it in
general form when viewed from the
side, but cutting edge much longer
and better developed, distinctly
angled behind middle, the angle
clearly i-epresenting a rudimentary
second cusp, and transverse portion
behind it a second commis-
sure, the tooth thus contain-
ing the modified elements
of one of the triangles of a
molar. Large upper pre-
molar with protocone
smaller than in the first
and second molars, though
well developed and of
essentially the same form ;
paracone smaller than pro-
tocone, therefore much
smaller than in first and
second molars ; hypocone
small but well developed ;
posterior margin of crown
more deeply emarginate
than that of molars.
Crowns of first and second molars squarish in outline, though
somewhat wider posteriorly than anteriorly. Outer re-entrant
angles deeper in second than in first. Pi-otocone long and rather
i'lG.
Sorex aranetis.
SOREX 35
low. Hypocone small but well developed. Third upper molar
with about half the crown area of second, all the elements of the
tooth present except hypocone. First and second lower molars
alike in size and form, the posterior triangle slightly larger than
anterior. Third molar smaller than the others, the relative size
of the triangles reversed, but elements of tooth all present.
Measuretiients. — While there is some variation in size among
the different races the head and body in full-grown individuals
is seldom if ever less than 65 or more than 80 mm. The tail
falls short of head and body by from 20 to 35 ram., being thus
relatively shorter than in either of the other European species.
The hind foot ranges from 11 to about 14 '5 mm. and the
condylobasal length of skull from 17 '8 to 20 mm. Detailed
measurements are given under each of the subspecies.
Bemarlcs. — Sorex araneus, the most widely distributed and
best known of the European shrews, is so easily recognized that
it needs no special comparison with the other species. Specimens
of the dark race from Charente, France, have a superficial
likeness to Sorex alpinus, but are at once distinguishable
externally by their blackish instead of slaty coloration and
relatively short tail. Immature, dull coloured individuals of
the other races may occasionally be confused with S. minutus.
The same is true of shrunken, badly prepared skins. But some
trace of the tricolor pattern can almost invariably be found in a
common shrew no matter what its condition, while a glance at
the size of the teeth and form and proportions of the unicuspids
will always serve to determine the identity of any specimen in
hand.
SoBEX AKANEUS AKANEUS Linnseus.
1758. [Sorex^ araneus Linnseus, Syst. Nat., i, 10th ed., p. 53 (Sweden).
1828. Sorex coronatus Millet, Faune de Maine-et-Loire, i, p. 18 (Blou,
Maine-et-Loire, Prance).
1828. Sorex iiersonatus Millet, Faune de Maine-et-Loire, I, p. 18, foot-
note (Rejected MS. name for coronatua). Not Sorex personatus
I. Geoffroy, 1827.
1832. Sorex concmnus Wagler, Isis, p. 54 (Bavaria).
1832. Sorex rhinolophus Wagler, Isis, p. 54 (Bavaria).
1832. Sorex vielanodon Wagler, Isis, p. 54 (Bavaria).
1838. Sort's; vulgaris Nathusius, Wiegmann's Archiv fiir Naturgesch.
IV, I, p. 45.
1839. Skirex] macrotrichus de S61ys-Longchamps, Etudes de ^licromamm.,
p. 20. (Specimen of S. araneus briefly described as agreeing with
the S. "inacrotrichus Mehlis MSS. No locality given.)
1839. S[orex'] labiosus Jenyns, Ann. Nat. Hist., ii, p. 326, January, 1839.
(Frankfurt a/.\I., Germany.)
1857. Sorex vulgaris Blasius, Siiugethiere Deutschlands, p. 129 (part).
1895. Sorex araneus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,
1895.
1910. Sorex araneus Trouessart, Faune Mamm. d'Europe, p. 51 (part).
T[ipc locality. — Upsala, Sweden.
Geographical distribution. — Western Continental Europe,
36
INSECTIVORA
except Atlantic watershed of south-western Norway, from
Finland to central France, central Germany, and northern Hun-
gary. Exact southern and eastern limits of I'ange not known.
Diagnosis. — Size small (condylohasal length of skull usually
17 "8 to 19 mm., hind foot usually 11 to 13 mm.) ; colour rather
dark, the back ranging from bister to a deep blackish brown ;
sides distinctly lighter than back except in specimens representing
the pallid extreme of colouration ; teeth moderately pigmented,
the hypocone of m^ and m'^ usually white to tip.
Teeth. — The teeth show no special peculiarities of form. In
pigmentation they represent the extreme of restriction of the dark
areas. While the area of j^igoientation on all of the cusps is less
extensive than in the Alpine and Pyrenean races, the ditierences
are best seen in the hypocones of the three lai'ge upper cheek
teeth and }:)rotocone of m^, as these small cusps, unlike the larger
ones, may be completely without brown colour. A comparison
of seventy-five topotypes of Sorex araneus with eighty Swiss
specimens of *S'. araneus tetragonmus gives the following results : —
araneuK. tetragonurug.
Large premolar with pigment on hypocone . -0% ... 38'7 %
First molar with pigment on hypocone. . 22'6 % ... 93"7 %
Second molar with pigment on hypocone . 21'2 % ... 88' 7 %
Third molar with pigment on protocone . 45'.3 % ... 100'0%
None of the small cusps pigmented . . 54 • G % ... " %
All of the small cusps pigmented . . . -0% ... 37'2%
Measurements. — Average and extremes of twenty specimens
from the type locality : head and body, 77 "5 (72-85) ; tail, 40" 1
(38-43); hind foot, 12 '6 (12—13). Average and extremes of
nine specimens from Lillehammer, centi'al Gudbrandsdal, Norway
(dry): hind foot, 12 '3 (12 "2-1 2 '6). Average and extremes of
four specimens from Holaaker, upper Gudbrandsdal, Norway :
head and body, 70-2 (69-71); tail, 36-5 (34-39); hind
foot, 12 "2 (12-13). Average and extremes of twenty specimens
from Brunswick, Germany : head and body, 78 " 9 (68-85) ;
tail, 39 ■ 7 (36-45) ; hind foot, 1 2 • 7 (1 2 • 2-1 3 -^O). Average and
extremes of ten specimens from Waremme, Liege, Belgium : head
and body, 68-9 (66-72); tail, 42-3 (38-47)'; hind' foot (dry),
12 '2 (11 '8-1 2 '8). For cranial measurements see Table, p. 46.
Specimens examined. — Two hundred and ninety-eight, from the following
localities : —
Norway : Molmen, Upper Gudbrandsdal, 2 ; Holaaker, Upper Gud-
brandsdal, 4 ; Lesjevark, Middle Gudbrandsdal, 1 ; Lillehammer, Middle
Gudbrandsdal, 9 (U.S.N.M.) ; Eggedal, Buskerud, 8 (U.S.N.M.) ; Spjosod,
Telemarken, 4 (U.S.N.M.) ; Asker, near Ohristiania, 4 (U.S.N.M.) ; Holme,
Mandal, 7.
Sweden: Upland,!; Upsala, 97 (U.S.N.M.) ; Skaane, 3 (U.S.N.M.).
Denmabk : HillerOd, Zealand, b ; Nystad, LoUand, 3 (U.S.N.M.) ;
Skansen, Lolland, 1 (U.S.N.M.).
Holland: Oosterbeek, Guelderland, G; Leiden, 4 (U.S.N.M.).
Belgium : Hastiere, Namur, 1 ; Waremme, Li6ge, 10 (U.S.N.M.).
Fbance : Guines, Pas-de-Calais, 4 ; Manonville, Meurthe-et-Moselle, 2 ;
Barbizon, Seine-et-Marne, 3.
37
Gebmany: Brunswick, 35 (BM. and U.S.N.M.) ; Bodethal, Harz Mts.,
15 (U.S.N.M.); Mcauseklippe, Harz Mts., 2 (U.S.N.M.); Bahrenberg, Harz
Mts., 9 (U.S.N.il.) ; Tharaud, Saxony, 1 ; Magdeburg, Saxony, 5 ; Moritz-
burg. Saxony, 1 (U.S.N.M.); Ingeiheim, Rheinhessen, 3; Nuremberg,
Bavaria, 10 (U.S.N.M.) ; Marxheim, near Monheim, Bavaria, 15 ; Strass,
near Burgheim, Bavaria, 1 ; Eulengrund, Riesengebirge, Silesia, 2
(U.S.N.M.) ; Wolfshau, near Sneekoppe, Riesengebirge, Silesia, 8 (U.S.N.M.);
Niesky, Silesia, 5 ; near Konigsberg, 6 (U.S.N.M.) ; no exact locality, 1.
AusTRiA-HuKGAEY : Haida, Arva, Bohemia, 9.
6, 9.
2(5,2 9.
6.
6, 5 9.
dal.
9.
4:6,2 9.
4 d,2 9.
9.
3 6,19.
2 6,2 9.
6, 3 9, 2.
9.
5 al.
<5, 2 9.
3 6, 4 9.
2 6.
6.
3 6, 2 9.
2al.
1.
3 6. 4 9.
jNIolmen, Gudbrandsdal.
Norway.
Holaaker, Gudbrands-
dal, 1900 ft.
Lesjevark, Gudbrands-
dal.
Holme, Mandal, 200 ft.
Norway.
Upland, Sweden.
(G. Kolthojf.)
Hillerod, Zealand, 10 m.
Denmark.
Oosterbeek, Guelderland ,
10-15 m. Holland.
Hastiere, Namur, Bel-
gium.
Guines, Pas-de-Calais,
10 m. Prance.
Barbizou, Seine - et-
Marne.
Auerum Forest, Bruns-
wick, Germany.
Tharandt, Saxony.
Magdeburg, Saxony.
Ingelhoim, Rheinhessen.
Marxheim, Bavaria.
Bayreuth, Bavaria.
Strass, Burgheim, Ba-
varia.
Niesky, Silesia, 200 m.
(W. Bacr.)
No exact locality.
Germany.
Haida, Bohemia.
Haida, Bohemia.
R. J. Cuuinghame
R. J. Cuninghame
(P)-
Miller Collection.
K. J. Cuninghame
Lord Lilford (p).
O. Thomas (c & p).
O. Thomas (c & p).
G. A. Boulenger
(c & p).
0. Thomas (c & p).
G. S. Miller (c).
G. Barrett-Hamil-
ton (c & p).
Lord Lilford (p).
Dr. W. Wolterstorff
(P)-
C. Hilgert (c).
Lord Lilford (p).
IMiller Collection.
Lord Lilford (p).
Lord Lilford (p).
Zool. Soc. Collec-
tion.
Stockholm ^Museum
(E).
Lord Lilford (p).
Lord Lilford (p).
98. 5. 2. 1-2.
98. 2. 28. 1-4.
7. 7. 7. 4452.
8. 8. 9. 1-6.
8. 8. 9. 40.
8. 9. 8. 19.
98. 6. 7. 2-7.
98. 2. 1. G-8.
94. 7. 9. 1.
94. G. 6. 4-7.
8. 8. 4. 155-158.
11. 1. 2. 75-80.
99. 1. 9. 14.
92. 12. 1. 3-7.
8. 11. 2. 6-3.
^. 9. 8. 7-13.
7. 7. 7. 2363-
2869.
8. 9. 8. 20.
99. 1. 9. 9-13.
55. 12. 26. 300-
301.
46. 6. 2. 36.
8. 9. 8. 14-18.
8. 9. 8. 21-22.
SOREX ARANEUS CASTANEUS JenyiLS.
1838. S[oc(?a-] tctragonums var. ;8 Slorcx] castaneus Jenyns, Ann. Nat.
Hist., I, p. 424, August, 1838 (Burwell Fen, Cambridgeshire,
England).
1857. Sorex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part)
1910. Sorex araneus Trouessart, Faune Mamm. d'Europe, p. 51 (part).
Type locality. — Burwell Fen, Cambridgeshire, England.
Geographical distribution. — Great Britain. Usually confined
38
INSECTIVORA
to the mainland, though occurring on Bardsey Island, Carnar-
vonshire.
Did'jiwsis. — Similar to Sorex araneus araneiis, but colour in
series of skins averaging less dark, that of dorsal area ranging
from hair-brown slightly tinged with bister to seal-brown, and
seldom if ever attaining the deep blackish brown frequently
seen in the typical race.
Skull mid teeth. — The skull and teeth resemble those of true
Sorex araneus.
Measureiiients.--Avevd.ge and extremes of ten specimens from
Cromarty, Scotland: head and body, 70*6 (62-78); tail, 39 "8
(38-43); hind foot, 12 '9 (12- 5-13). Average and extremes
of eight sjaecimens from Aberia, Merioneth : head and body,
67-5 (58-73); tail, 38-7 (36-41); hind foot, 12 (11-13).
Average and extremes of eight specimens from Grimsby, Lincoln-
shire : head and body, 65-2 (58-72); tail, 42 (38-44); hind
foot, 12 '5 (12-13). Average and extremes of six specimens
from Northlew, Devonshire : head and body, 66 '6 (65-70) ; tail,
36 '8 (35-39); hind foot, 13 (13). For cranial measurements
see Table, p. 48.
Specimens examined. — Two hundred and fifty-two, from the following
localities : —
Scotland: Black Isle, Cromarty, IG ; South Sutor, Cromarty, 5;
Nairn, Morayshire, 4 ; Dunphail, Elgin, 2 ; Gordonstown, Elgin, 3 ; Lhan-
bride, Elgin, 1; Lossiemouth, Elgin, 3; Grantown-on-Spey, Elgin, 26
(Wilson) ; Kennordy, 1 (Wilson) ; Cortachy, Forfar, 6 (Wilson) ; Cromlix,
Stirling, 10 ; Islay, 3 ; Dunkeld, Perthshire, 2 ; Loch Earn Head, Perth-
shire, 1; Stoclibriggs, Lanarkshire, 2; Kirtle Bridge, Dumfriesshire, 4;
Wyseby, Dumfriesshire, 3.
Wales : Aberia, Merionethshire, 8 ; near Bridgend, Glamorganshire, 6
Bardsey Island, Carnarvonshire, 1.
England : Berwick-on-Tweed, Northumberland, 2 ; Riding Mill-ou-
Tyne, Northumberland, 3 ; Newby Bridge, Lake Windermere, Cumber-
land, 1 ; Grimsby, Lincolnshire, 17 ; Whitnash, Warwickshire, 1 ; Rugby,
Warwickshire, 2 ; Filey, Yorkshire, 3 ; Wellersey Hill, Broadway, Wor-
cestershire, 1 ; W^est Cheshire, 1 ; Shropshire, 1 ; StaSordshire, 1 ; Swith-
land, Leicestershire, 10 ; Bishopstoke, Herefordshire, 1 ; Leominster, Here-
fordshire, 1 ; Graftonbury, Herefordshire, 19 ; Lilford, Northamptonshire,
2 ; Drinkstone Park, Bury St. Edmunds, Suffolk, 2 ; Lowestoft, Suffolk, 6 ;
Wormsley, Oxfordshire, 1 ; Stokenchurch, Oxfordshire, 4 ; Cambridge-
shire, 1; Kensington Gardens, London, 1; Hillingdon, Middlesex, 3;
Bletchingley, Surrey, 4; Godalming, Surrey, 2; Merstham, Surrey, 1;
Richmond Park, Surrey, 1; Crowborough, Sussex, 3; St. Leonard's,
Sussex, 1 ; Tunbridge Wells, Sussex, 1 ; Eastwell, Kent, 3 ; Lyndhurst
Road, Hampshire, 2; New Forest, Hampshire, 17; Basingstoke, Hamp-
shire, 1 ; Alum Bay, Isle of Wight, 3 ; Cliftou Bridge, Gloucestershire, 2 ;
Leigh Woods, Clifton, Gloucestershire, 3 ; Blandford, Dorsetshire, 1 ;
Combmartin, Devonshire, 4 ; Chagford, Devonshire, 5 ; Northlew, Devon-
shire, 6 ; no exact locality, 3.
Remarks. — While the British common shrew is an incom-
pletely differentiated form, the average characters of the large
series of specimens examined seem important enough to warrant
the use of Jenyn's name. As compared with the Continental
SOREX 39
race the dark extreme is less dark and less frequent, while the
light extreme is more light and more frequent. The constantly
small size of this animal as compared with the large Continental
races (benjensls, ietragonurus and pi/renaicus), is shown by the
fact that among 102 British specimens measured by many
different collectors the hind foot exceeds 13 '2 mm. in only six
instances.
9. Black Isle, Cromarty, W.R. Ogilvie-Grant 94.10.6.1.
Scotland. (c & p).
9 i. Black Isle, Cromarty. W. R. Ogilvie-Grant 11. 1. .3. 103-
(c & p). 107. 109-112.
2(5. South Sutor, Cromarty. W.R. Ogilvie-Grant 11. 1. 8. 108,
(c & p). 113.
2 (5, 2 ?. Nairn, Morayshire. W.R. Ogilvie-Grant 11. 1. 3. 99-102.
(c & p).
5. Morayshire. E. R. Alston (c & p). 79.9.2.5.79.
9. Dunphail, Elginshire. W. R. Ogilvie-Grant 11.1.3.97.
(c & p).
1. Lhanbride, Elginshire. Miller Collection. 7. 7. 7. 3598.
2 6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.95-96.
shire. (c & p).
6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.98.
shire. (c & p).
3(5,7 9. Cromlix, Stirlingshire. W. R. Ogilvie-Grant 11.1.3.114-128.
(c & p).
3 al. Islay. H. RusseU (c & p). 9. 9. 11. 1-3.
6,9. Stockbriggs, Lanark- E. R. Alston (c & p). 79.9.25.11-12.
shire.
6, 3 9. Kirtle Bridge, Dumfries- Miss D. Bate (c <fep). 11. 1. 3. 88.
shire. 11. 1. 3. 92-94.
3 9. Wyseby, Dumfriesshire. Miss D. Bate (c & p). 11. 1. 3. 89-91.
5 (5, 3 9. Aberia, Merionethshire, G. H. Caton Haigh 11. 1. 3. 156-
Wales. (c & p). 163.
3(5,3 9. Bridgend, Glamorgan- R. I. Pocock (c & p). 11. 1. 3. 164-
shire. 169.
2 al. Bar wick-on- Tweed, J. H. Fryer (c & p). 47.11.11.6-7.
Northumberland,
England.
1 al. Newby Bridge, Cumber- J. Paul (c & p). 94. 9. 3. 2-3.
land .
(5,5 9,2. Grimsby, Lincolnshire. G. H. Caton Haigh 11. 1. 3. 124-
(c & p). 131.
(5,6 9. Swithland, Leicester- P. A. Butler (c & p). 11. 1. 8. 132-
shire. 188.
1 al. Shropshire. T. 0. Eyton (c & p). 68.10.12.4.
9 al. Bishopstoke, Hereford- S. 0. Ridley (c&p). 84. 10. 6. 1.
shire.
8 c?, 8 9. Graftonbury, Hereford- W. de Winton 11.1.3.72-87.
shire. (c & p).
3 St. Graftonbury, Hereford- W. de Winton 96. 4. 28. 17-19.
shire. (c&p).
2al. Bury St. Edmunds, J. H. Powell (c & p). 80.5.22.1-2.
Suffolk.
3 (5, 3 9. Lowestoft, Suffolk. 0. Thomas (c & r). 11. 1. 8. 189-
144.
4 9. Stokcnchurch, Oxford- W.R. Ogilvie-Grant 11. 1. 8. 145-
shire. (c & p). 148.
3 9. Hillingdon, Middlesex. MiUer Collection. 7. 7. 7. 8595-
(O. Thomas.) 3597.
40
IXSECTIVORA
1. Kensington Gardens, N. Churton (c & p). 161. a.
Loudon.
4 f . Bletchingley, Surrey. W.R. Ogilvie-Grant 11. 1. 3. 149-
_(p). 152.
1. Crowborough, Sussex. Miller Collection. 7. 7. 7. 3593.
{W.li.Ogilvic-Grant.)
9 al. Richmond Park, Surrey. Prof. Owen (p). 75.9.17.1.
7 6, 10 9. New Forest, Hampshire. Miller Collection. 7. 7. 7. 2851-
2867. 3023.
9. Basingstoke, Hamp- Miller Collection. 7. 7. 7. 3611.
shire. {W. P. Stark.)
6, 2 9. Alum Bay, Isle of Wight. 0. Thomas (c & p). 11. 1.8.153-155.
9 (albino) Winscombe, Somerset. F. A. Knight (c.&p). 4. 8. 9. 1.
6, 4 9. Nortblew, Devonshire. R. C. Wroughton 11. 1. 3. 170-
(c & p). 174.
2 c5, 3 9. Chagford, Devonshire. Miller Collection. 7. 7. 7. 3606-
(W. P. Stark.) 3610.
1 al. England. Dr. J. E. Gray (p). 46. 5. 2. 7.
80RE.\ ARANEUS SANTONUS Mottaz.
1908. Sorex santonus Mottaz, Bull. Soc. Zool. de Geneve, i, p. 118,
April 30, 1908. Type in Mottaz Collection.
1910. Sorex ,';a)itonus Trouessart, Faune Mamm. d'Europe, p. 54.
Type locality. — Ligiiieres-Sonneville, Charente, France.
Geographical distribution. — Known only from the vicinity of
the type locality.
Diagnosis. — Size as in Sorex araneus araneus and the other
small races ; colour throughout a nearly uniform dark sooty
brown.
Colour. — Upper parts dai'k sepia anteriorly, deepening to
blackish posteriorly, the sides essentially similar to the back.
Underparts a slaty drab washed with wood-brown and forming
no evident contrast with sides and back, though a well-detined
line of demarcation is present. Tail blackish throughout. Feet
scantily clothed with inconspicuous brownish hairs.
Skull and teeth. — The skviU and teeth do not differ appreciably
from those of the other small races.
Measurements. — Type (female), from Mottaz : head and body,
75 ; tail, 42 ; hind foot, 13. For cranial measurements see
Table, p. 51.
Specimens examined. — Ten, all from the type locality (Mottaz).
BemarJcs. — The Charente shrew is so different in aspect from
tlie other European races of Sorex araneus that it needs no
special comparison with any of them. Its jseculiar colour is,
however, almost exactly duplicated by that of a larger, longer -
tailed form from Asia Minor.
41
SOKEX ARANEUS EURONOTUS Miller.
1901. Sorex araneus curonotus Miller, Proc. Biol. Soc, Washington, xiv,
p. 44, April 25, 1901 (IMontrejeau, Haute-Garonne, Prance). Type
iu U.S. National Museum.
1910. Sorcx araneus curonotus Trouessart, Paune Mamm. d'Europe, p. 53.
Ti^je locality. — Montrejeau, Haute-Garonne, France.
Geographical distribution. — Probably the i^lains between the
Pyrenee.s and the Garonne. At present known from the type
locality only.
Diarjnotiis. — Similar to Sorcx araneus araneus, but colour in
.summer (winter pelage not known) more dull, the sides scarcely
if at all contrasted with the back, and underparts more heavily
washed with wood-brown. 8kull with slightly narrower less
elevated brain-case, and teeth just perceptibly smaller than in
true araneus.
Measurements. — External measurements of type specimen :
head and body, 78; tail, 44; hind foot (dry), 12 '6. Average
and extremes of nine specimens from the tyjje locality : head and
body, 71-4 (67-78); tail, 42 (37-44); hind foot (dry), 12-5
(12-2-13'0). For cranial measurements see Table, p. 51.
Specimens examined. — Niue, all from the type locality (U.S.N.M.).
SoREX ARANEUS BERGENSIS Miller.
1909. Sorex araneus bergensis Miller, Ann. and Mag. Nat. Hist., 8th ser..
Ill, p. 416, May, 1909. Type iu U.S. National Museum.
1910. Soi-ex araneus bergensis Trouessart, Paune Mamm. d'Europe, p. 52.
Tijpe locality. — Graven, Hardanger, Norway.
Geographical distribution. — Western Norway, from region of
Bergen, north at least into Nordland.
Diagnosis. — Larger than Sorex araneus araneus (hind foot,
13-6 to 14-4 mm. ; condylobasal length of skull, 19 to 20 mm.),
and colour in summer pelage darker, the dark brown or blackish
dorsal area sharply defined from the yellowish brown of sides.
Colour. — Dorsal area well defined, ranging from a blackish
seal-brown to bister, usually darker on rump and lumbar region
than on head, and almost invariably with sharp lateral line
of demarcation. Sides yellowish isabella-colour. Underparts
greyish, washed with light wood-brown. The contrast between
colour of sides and underparts, though less marked than that
between sides and back, is usually evident. Feet scantily clothed
with inconspicuous Isabella-coloured or dusky hairs. Tail obscurel)*
bicolor, brownish above, dull yellowish below.
Sliull and teeth. — The skull and teeth do not differ appreciably
from those of Sorex araneus araneus, except in their larger size,
as shown by the detailed measurements. The pigmentation of
42 INSECTIVORA
the teeth is scarcely, if at all, more extensive than in the typical
race.
Measurements. — External measurements of type : head and
body, 80-5; tail, 44-5; hind foot, 13-6. Average and extremes
of eleven specimens from the Bergen district : head and body,
78-7 (76-83) ; tail, 49-3 (44-56) ; hind foot, 13-8 (13 -6-14 -4).
For cranial measurements see Table, p. 51.
Specijnens examined. — Twenty-seven, from the following localities: —
Noeway: Vefsen, Nordland, 1; Skjserdal, Nordfjord, 7; Opheim,
Bergen, 4; Graven, Bergen, 8 (U.S.N.M.) ; near city of Bergen, 7 (B.M.
and U.S.N.M.).
Bemarlcs. — This large race of Sorex araneus, closely resembling
the large Alpine and Pyrenean forms, but decidedly darker in
colour, appears to be strictly confined to the Atlantic slope of
western Norway. On the eastern watershed it is replaced by
true araneus, even so far north and west as the upper portion of
the Gudbrandsdal. I have not seen specimens in full winter
pelage, but an adult female taken at Graven on June 10, 1898
(No. 84,663, U.S.N.M.), is moulting, the winter fur remaining on
posterior half of body. Others taken at the same place and
about the same date have completed the change to the short,
velvety summer coat.
1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 2.
way. Waldo (c & p).
6, 4 9. Skjserdal, Nordfjord. C. H. Stephenson 8. 1. ,5. 1-5.
(c & P).
■2 6,29. Opheim, Bergen. Miller CoUection. 7. 7. 7. 44.53-
445G.
6, 2 ?. Bergen. ^Miller Collection. 7. 7. 7. 4457-9.
?. Bergen, 2,700 ft. G. Barrett-Hamil- 8. 9. 21. 1.
ton (p).
SoREX ARANEUS TETRAGONURUS Hermann.
1780 Sorc.v tetragonurus Hermann in Zimmermann, Geogr. Gesch., ii,
p. 383 (Strassburg, Germany).
1792. Sorcx quadricaudatus Kerr, Auim. Kingd., p. 208 (Strassburg,
Germany). (Based on Pennant's account of S. tetragomirus
Hermann.)
1835. Sorcv fodiens Duvernoy, Mem. Soc. du I\Ius. d'Hist. Nat., Stras-
bourg, II, p. 17. Part : skull, not animal.
1835. Sorex hermanni Duvernoy, M6m. Soc. du Mus. d'Hist. Nat., Stras-
bourg, II, p. 23 (Near Strassburg, Germany). Part: animal, not
skull.
1857. Sarex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part).
1868. ? Sorex aranetis, pallidus Fitzinger, Sitzungsber. kais. Akad. Wis-
sensch., Wieu, Math. Naturwiss. Classe, lvii, pt. I, p. 488
(Based on specimen from unknown locality, probably in Italy,
figured by Bonaparte, Iconogr. Faun. Ital., fasc. xxis, pi., fig. 5.)
18G9. [Sorex vulgaris'] var. nitda Fatio, Faune Vert. Suisse, i, p. 127
(Bernese Oberland, Switzerland).
SOEEX 43
1869. ISorcx vulgaris] vkr. 7ugra Fatio, Faune Vert. Suisse, i, p. 127
(Lucerne, Switzerland).
1900. Slorcx} vulgaris var. vel subsp. mollis Fatio, Rev. Suisse de Zool.,
VIII, p. 471 (Substitute for nigra).
1901. Sorex araneus alticola ililler, Proc. Biol. Soc, Washington, xiv,
p. 43, April 25, 1901 (Meiringen, Switzerland). Tj^pe in U.S.
National Museum.
1905. S[orex] vulgaris crassicaudatus Fatio, Arch. Sci. Phys. et Nat..
Geneve, 4th ser., xix, p. 201, February 15, 1905 (Zermatt, Switzer-
land). Cotypes in Geneva Museum.
1905. Crossojjus ou Sorcx ignotus Fatio, Arch. Sci. Phys. et Nat., Geneve,
4th ser., xix, p. 202, February 15, 1905 (Switzerland). Part:
mandible, not skull. Type in Geneva Museum.
1905. Sorex araneus carpathicus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xv, p. 506, May, 1905 (Hatszeg, Hunyad, Hungary).
Type in British Museum.
1910. Sorex araneus tetragonurus and S. aranezis carpathicus Trouessart,
Faune Mamm. d'Europe, pp. 52, 54.
Type locality. — Strassburg, Germany.
Geographical distribution. — Alp.s and neighbouring portions of
Germany, France and Italy ; eastward through Tirol to the
mountains of Transylvania. "
Diagnosis. — Similar to Sorex araneus hergensis, but colour in
summer pelage not so dark, the contrast between back and
sides often not consj^icuous, though back is frequently blackish ;
pigmentation of teeth more extensive than in the northern races,
the hypocone of m^ and m- usually brown at tip (see tabular
comparison with true araneus on p. 36).
Measurements. — Average and extremes of eight specimens
from the type locality : head and body, 75-8 (71-79) ; tail, 47-5
(45-50); hind foot, 13-5 (13 •2-14 -2). Average and extremes
of twenty specimens from Andermatt, Uri, Switzerland : head
and bod}', 73-1 (70-81); tail, 51-1 (50-55); hind foot, 13-3
( 1 3-14). Measurements of an adult female from Zermatt, Valais,
Switzerland (in alcohol) : * head and body, 74-6 ; tail, 51 ; hmd
foot, 13-6. Average and extremes of three specimens from
Vitznau, Switzerland : f head and bodv, 67-3 (62-72); tail,
46-3 (44-49); hind foot, 13-2 (12 -8-13 -6). Type oi Sorex
araneus alticola Miller (Meiringen, Switzerland,' No. 85,930
U.S.N.M.) : head and body, 76 ; tail, 55 ; hind foot, 14. Average
and extremes of ten specimens from Meiringen, Switzerland : t
head and body, 74-0 (70-77); tail, 52-5 (47-57); hind foot,
lo'3 (13-14). Average and extremes of four specimens from
Locarno, Ticino, Switzerland: head and body, 76-7 (76-82):
tail, 48-5 (45-50): hind foot, 13-6 (13-4-13"'-8). For cranial
measurements see Table, p. 49.
* Cotype of Sorex vtdgaris crassicaudatus Fatio.
t Essentially topotypes of Sorex vulgaris nigra and itwllis Fatio.
X Topotypes of Sorer araiieus alticola ililler and essentially topo'types
of Sorex vulgaris nuda Fatio.
44
INSECTIVORA
Specimens examined. — Two hundred and sixty-two, from the following
localities : —
Feance : Etupes, Doubs, 10; Barcelonnette, Basses-Alpes, 8; Cha-
monix, Haute-Savoie, 10 (U.S.N.M.).
Germany : Strassburg, 8.
SwiTZEBLAKD : Geneva, 17 (U.S.N.M. and Mottaz) ; St. Cergues, Vaud,
19 (U.S.N.M. and Mottaz) ; Chesieres, Vaud, 13 (Mottaz) ; Bioux-Dessus,
Vaud, 4 (Mottaz) ; Les Plans, Vaud, 4 (U.S.N.M.) ; Zermatt, Valais, 7 (B.M.,
U.S.N.M. and Geneva) ; Stalden, Valais, 2 (Geneva) ; Grindelwald, Bern, 4
(U.S.N.M.) ; Briinig, Bern, 9 (U.S.N.M.) ; Meiringen, Bern, 17 (U.S.N.M.) ;
Vitznau, Lucerne, 6 (B.M. and U.S.N.M.) ; Goschenen, Uri, 5 (U.S.N.M.) ;
Audermatt, Uri, 49 (U.S.N.M.) ; Hospenthal, Uri, 1 (U.S.N.M.) ; Ziirich, 1 ;
Murgsee region, St. Gallen, 16 (U.S.N.M.); Degersheim, St. Gallen, 3
(U.S.N.M.); Uzwil, St. Gallen, 1 (U.S.N.M.); Sitterwald, St. GaUen, 5
(U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 (U.S.N.M.) ; Wildlnrchli, Appen-
zell, 1 (U.S.N.M.) ; Albulapass, Grisous, 1 (U.S.N.M.) ; Untervatz, Grisons,
5 (U.S.N.M.); Grisons, no exact locality, 4 (U.S.N.M.); Faido, Tioino, 3
(B.M. and U.S.N.M.); Lugano, Ticino, 1 (U.S.N.M.); Gentilino, Ticino, 1
(U.S.N.M.) ; Locarno, Ticino, 4 ; no exact locality, 1.
AusTKiA-HuNGARY : Hatszeg, Hunyad, Transylvania, 10 ; Csalloku'z-
Somorja, Pressburg, Hungary, 3 ; Schwaz, Tirol, 2 (U.S.N.M.).
Italy : Near Turin, 4 (Turin) ; Unerzio, Cuneo, 1 ; Vallombrosa, near
Florence, 2.
8.^,2 9.
Etupes, Doubs, 350 m.
France. (C. Mottaz).
0. Thomas (p).
8. 8. 10. 14-24.
3 6.
Barcelonnette, Basses-
Alpes. (C. Mottaz.)
0. Thomas (v).
8. 8. 10. 25-27.
4 6, 1 9.
Strassburg, Alsace.
(C. Muttaz.)
0. Thomas (p).
8. 8. 10. 28-32.
lal.
Zermatt, Valais, Swit-
Dr. J. Anderson
91. 10. 15. 2G.
zerland.
(c & P).
3 6.
Vitznau, Lake Lucerne.
0. Thomas (c & p).
5. 8. 2. 82-34.
1 al.
Zurich.
C. Mosch (p).
89. 11. 8. 3
9.
Faido, Ticino.
0. Thomas {c & p).
5. 8. 2. 19.
2 i5' 2 9.
Locarno, Ticino.-
0. Thomas (c & p).
5. 8. 2. 2-5.
9.
Switzerland.
E. E. Alston (p).
79. 9. 25. 10.
8 6, 1 9.
Hatszeg, Hunyad, Tran-
C. G. Dauford (c).
3. 2. 2. 8-10.
sylvania, 1500-2000£t.
3. 11. 8. 15.
Hungary.
3.
Csallokdz-Somorja,
Budapest ^Museum
94. 3. 1. 22-25,
Pressburg, 400 ft.
(E).
Hungary.
2al.
Vallombrosa, Florence,
Dr. G. Cecconi
1. 8. 2. 2-3.
Italy.
(c & P).
80REX ARANEUs pYREXAicus Miller.
1909. Sorex araneus injrenaicns Miller, Ann. and JIag. Nat. Hist., 8th ser.,
Ill, p. 416, May, 1909. Type in British ]Museum.
1910. Sorex araneus pyrenaicus Trouessart, Faune Mamm. d'Europe, p. 53.
Type localiti/. — L'Hospitalet, Ariege, France. Altitude
4,700 feet.
Geographical disirihution. — Pyrenees. At present known
from France only, though occurring on the south slope of the
mountains in the Department of Pyrenees-Orientales.
Diagnosis. — Very similar to Sorex araneus tetragonurus, but
distinguishable by the duller, less evidently tricolored summer
SOREX 45
pelage (winter coat not known), in which the back rarely if
ever assumes the blackish-brown tints often seen in the Alpine
form.
Colour. — The colour scarcely needs detailed description. As
compared with that of the other large races it is characterised
by extreme dulness and lack of noticeable contrast between the
dorsal area and sides. In none of the thirty-two skins examined
is the back so dark as in the dark individuals frequently found
in Switzerland in summer.
Skull and teeth. — As in S. araneus tetragonurus.
Measurements. — External measurements of type : head and
bod\', 72; tail, 51 ; hind foot, 14. Average and extremes of six
specimens from the type locality : head and body, 70*6 (69-72) ;
tail, 47 (44-4-51); hind foot, 13-3 (13-14). Average and
extremes of nine specimens from Bareges, Hautes-Pyrenees : head
and body, 72-5 (69-75); tail, 45 (42-49); hind foot, 13-3
(13-14). For cranial measurements see Table, p. 50.
Specimens examined. — Thirty-two, from the following localities in the
French Pyrenees : —
Porte, Pyren^es-Orientales (Spanish watershed), 9 ; I'Hospitalet, Ariege,
12 ; As-les-Thermes, Ariege, 2 ; Bareges, Hautes-Pyr6n6es, 9.
Bemarhs. — In its dull colour the Pyrenean shrew differs from
the other large members of the group in much the same way as
the small Sorex araneus euronotus of the neighbouring lowlands
differs from true araneus. It thus represents the opposite
extreme from the dark Norwegian form.
c$, ?. Port6, Pyr6n6e8-Orien- G. S. Miller (c). 8. 8. 4. 142-143.
tales, 1600m. France.
2 i, 9. Porte, Pyrenees-Oricn- 0. Thomas (p). 8. 9. 1. 49-51
tales, lGOO-1700 m.
{A. Bobcrt).
6,4:9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.151-154.
4700 ft. 301.
(8. 8. 4. 301. Type of subspecies.)
?. Ax-les-Thermes, Ariege, G. S. Miller (c). 8. 8. 4. 141.
2600 ft.
6, 6 9. Bareges, Hautes-Pyr6- G. S. Miller (c). 8. 8. 4. 144-150.
uees, 1300-1500 m.
SOREX ARANEUS FRETALIS Miller.
1909. Sorex araneus frctalis Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,
p. 416, IMay, 1909. Type in British Museum.
1910. Sorex araneus fretalis Trouessart, Faune Mamm. d'Europe, p. 52.
Type locality. — Trinity, Jerse\', Channel Islands.
Geographical distribution. — Known only from the island of
Jersey.
Diagnosis. — Like Sorex araneus araneus, but skull with
rostral portion shortened, broadened and deepened, and anterior
46
INSECTIVOKA
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48
INSECTIVORA
3
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Germany : near Sneekoppe, Riesen
gebirge .
S. araneus castaneus.
Scotland : Black Isle, Cromarty
Cromlix
Morayshire .
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Swithland, Leicestershir
49
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52
INSECTIVOEA
teeth (incisors both above and below, and first and second ujjper
unicuspids) enlarged.
Colour. — Siiramer pelage about like that of S. araneus araneus,
but upper parts perhaps not so dark. Sides apparently averaging
lighter than in true araneus, and underparts in some specimens
a very pale, almost whitish, bufiy grey, decidedly lighter than in
any skins of the other races yet examined.
SJchU. — In size and general form the skull does not difier
appreciably from that of true Sorex araneus, but on comparison
of specimens the rostral portion in front of large premolar is
seen to be relatively shorter, broader and deeper. The greater
bi'eatlth is especially noticeable from below, the space between
the anterior unicuspids about equalling diameter of these teeth,
while in ail the other races (except (S'. o. granarius) it is evidently
less.
Teeth. — While in general the teeth resemble those of Sorex
araneus araneus, the large incisors both above and below are
appreciably more robust, and the first and second upper uni-
cuspids are wider. The difference is particularly noticeable in
the lower incisor, the shaft of which is thickened and the lobes
on cutting edge distinctly enlarged.
Measurements. — External measurements of tyjje (adult female) :
head and body, 63; tail, 48-2; hind foot, 13. For cranial
measurements see Table, p. 51.
Specimens exmnined.— Five, all from the island of Jersey.
Bemarhs. — The Jersey shrew is easily distinguishable fi'om
the other races by its enlarged anterioi' teeth. It is probable
that a sufficient series of skins will show that there is an average
difference in colour as well.
9,6. Jersey, Channel Islands. 0. Thomas (p). 8.9.2.1-2.
{I!. H. Bunting.) (8. 9. 2. 1. Type of subspecies.)
Sorex araneus oranarius Miller.
1910. Sorex araneus granarius Miller, Ann. and Mag. Nat. Hist., 8th ser.,
VI, p. 459, November, 1910. Type in British Museum.
Ti/pc localiiij. — La Gi'anja, Segovia, Spain.
Geographical dislrihuiion. — At present known from the type
locality only, and probably confined to the mountains of central
Spain.
Diagnosis. — Smallest known European race of Sorex araneus,
the condylobasal length of skull onlj^ about 17 "5 mm., upper
tooth-row about 7 ■ 5 mm. ; palate wide anteriorly as in S. araneus
fretalis, but anterior teeth not enlarged.
External characters and colour. — Externally the animal shows
no special peculiarities, though the ear and the plantar tubercles
SOREX 53
seem to be relatively smaller than usual. Colour of specimens
in alcohol apparently as in true aranctts.
Skull and teeth. — Apart from its small size the skull does not
differ noticeably from that of the other races, except in the
relative shortness of the rostral j^ortion and tooth-row as com-
pared with the breadth of palate. Palatal breadth between
anterior unicuspids as great in proportion to width of teeth as
in S. araneus f fetalis. Teeth small, normal in form, the pig-
mentation apparently less extensive than usual.
Measurements. — Type (adult male), and an older male, also
from La Granja : head and body, 62 and 66 ; tail, 36 and 37 ;
hind foot, 11 '6 and 11*6; ear from meatus, 6'6 and 7. For
cranial measurements see Table, p. 51.
Specimens examined. — ^Two, both from the tj^pe locality.
Hemarlcs. — ^In its broad palate the (Tuadarrama shrew bears
a remarkable likeness to the form inhabiting the island of
Jersey. It is readily distinguishable from the Jersey animal
by its small size, and by the absence of all tendency to enlarge-
ment of the anterior teeth.
2 al. La Granja, Segovia, M.dela Escalera(c). 6.11.4.3-4.
Spain. (6.11.4.4. TyjJC of subspecies.)
SOREX MINUTUS Linnajus.
(Synonymy under subspecies.)
Geographical distribution. — Northern portion of Eurasia from
Ireland eastward (exact eastern limits of range not known). In
Europe south to the Pyrenees and southern Italy.
Diagnosis. — Size small, head and body usually about
50-60 mm., the tail 10-15 mm. shorter, condylobasal length
of skull 14 '8-16 "6 mm.; posterior lobe of anterior upper
incisor sub-terete, the length of its base about half that of
anterior cusp ; colour brown, the underparts always lighter
than back, and sides never specially contrasted.
E.iternal characters. — In general external characters Sorex
minutus agrees with S. araneus, except for its smaller size and
relatively longer tail.
Colour. — The colour rather closely resembles that of Sorex
araneus in dull summer pelage, except that the back usually has
a peculiar greyish cast not easy to describe, but by which it is
possible to recognise skins with much certainty. There is never
any indication of a specially differentiated colour area along sides.
U})per parts between sejjia and wotxl-brown in summei', more
nearly hair-brown in winter, the hairs slate-grey at ba.se and
with faint .silvery sub-terminal annulations more visible in some
lights than in others, and giving rise to the grej'ish effect already
alluded to. Underparts smoke-grey of varying depth, some-
54
INSECTIVORA
times almost whitish, the line of demarcation along sides usually
evident though not very conspicuous. Feet pale wood-brown
with a silvery gloss or suffused with drab, sometimes noticeably
lighter than back. Tail concolor with back above, not so dark
below.
SlcuU. — Apart from its conspicuously smaller size (condj'lo-
basal length, 14 '8 to 16 mm. instead of 17 "8
to 20 mm.) the skull differs notably from
that of Sorex araneus and S. nlpinus in the
narrower, more elongate brain-case, the outline
of which when viewed from above is distinctl)'
oval instead of sub-circular. Antero-external
jwrtion of border flattened, though less notice-
abh' than in Sorex araneus. Owing to its
different form the brain-case is less abruptly
Fio. 10. marked off" from interorbital region than in
Sorex minutus. *^^® other European species. Depth of brain-case
Nat. size. usually less relatively to breadth than in
S. araneus. Lachrymal foramen opening over
posterior half of first molar.
Teeth. — Aside from their very much smaller size (upper
tooth-row 6 to 7 mm. instead of 8 to 9-6 mm.) the teeth of
Sorex minutits differ in numerous details of form from those of
S. araneus. Anterior upper incisor with basal lobe nearly as
high as anterior cusp, so that the two points of the tooth are
essentially in line with the tips of the first three unicuspids.
AVhen viewed from below the posterior lobe appears, howe\er,
distinctly smaller than anterior cusp, while from the side its
outline is conspicuously higher and narrower than that of first
unicuspid. Anterior lower incisor with
first lobe on cutting edge as well defined
from anterior point of tooth as from second
lobe, and lengths of bases of all three lobes
approximately equal. Upper unicuspids not
essentially different in form from those of
Sorex araneus, but crowns perceptibly longer
than wide and less abruptly rounded off'
anteriorly, the resulting form less nearly ^"^" "'
T 1 J- 1 • J.1 J.T ■ -L Sorex miniitus. Anterior
square. In lateral view the outline is cjuite teetii in prutiie. x 5.
as in the larger animal. The relative size
of the unicuspids differs markedly from that in the larger
animal. The first, second and third are sub-equal, with the
second usually a trifle smaller than the third, and the first with
distinctly the greatest crown area. Fourth slightly smaller than
third, and fifth equally smaller than fourth'"', between which and
large premolar it is tightly crowded. First lower unicuspid
differing from that of S. araneus in its greater length along
cingulum and less height of cusjj, its form when viewed from
* Sometimes equal to fourth or slightly larger.
SOREX S5
the side thus noticeably different from that of first upper
unicusisid. Second unicuspid, together with other mandibular
teeth, essentially as in S. aranem. Maxillary cheek-teeth as in
S. araneus, except that hypocones are less developed, that on
large premolar obsolete.
Bemarks. — Sorex minutus is at once distinguishable from
S. araneus by its smaller size and relatively longer tail, as well
as by the more technical characters of the skull and teeth.
Immature individuals of araneus might sometimes be mistaken
for minutus, but their larger feet will serve to indicate their
identity ; while if the skull and teeth can be examined, a positive
identification is easily obtained.
Sorex minutus minutus Linnaeus.
1766. [Sorex] mimUus Linnaeus, Syst. Nat., i, 12th ed., p. 73 (Siberia).
1769. Swcx injgmxus Laxmann, Sibirische Briefe, p. 72 (Barnaul, Tomsk,
Siberia).
1789. [6'orea-] exilis Gmelin, Syst. Nat., i, 13th ed., p. 115 (Yenesei River,
Siberia).
1806. Soi'ex canaliculatus Ljungh, Kongl. Vetensk. Akad. Nya Handl.,
XXV3I, p. ,263 (Lommaryd Vicarage, northern Vedbo district,
Jornkoping, Sweden).
1811. Sorex pygtnx us Pallas, Zoogr. Rosso-Asiat., i, p. 134 (Ob and Yenesei
Rivers, Siberia).
1811. Sorex minimus Geoffrey, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 186
(Accidental renaming of minutus).
1832. Swex puviilio Wagler, Isis, p. 54 (Bavaria).
1838. S[orex] rusticus Jenyns, Ann. Nat. Hist., i, p. 423, August, 1838
"(England).
1838. Siorex] rusticus var. S S{orex} hihernicus Jenyns, Ann. Nat. Hist., i,
p. 423, August, 1838 (Dublin, Ireland).
1844. Sorex immilus Nilsson. Ofversigt af Kongl. Vetensk.-Akad. Forhandl.,
Stockholm, I, p. 33, March 20, 1844 (North-eastern Skaane,
Sweden).
1857. Sorex pygmmus Blasius, Siiugethiere Deutschlands, p. 133.
1895. Sorex minutus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,
1895.
1910. Sorex minutus Trouessart, Faune Mamm. d'Europe, p. 55.
Type locaJiiij. — Vicinity of the Yenesei River, Siberia.
Geoijraphiral distribution. — The entire European range of the
species, except southern Italy.
Diagnosis. — Teeth normal in size, the molars and anterior
upper incisor not enlarged.
Measurements. — .Average and extremes of thirteen specimens
from Grantown-on-Spey, Elgin, Scotland : head and body, 52 • 3
(49-55); tail, 36 (32 -5-39 -5); hind foot, 10-4 (10-11).
Average and extremes of eight specimens from the Isle of ^lan :
head and body, 59-6 (52-64); tail, 40-2 (36-43): hind foot,
11-1 (10-12). Average and extremes of five specimens from
Ariege, France: head and body, 55-8 (51-62); tail, 44*2
(42-46): hind foot, 11-4 (11-12). Average and extremes of
56
IN'SECTIVORA
four specimens from the Harz Mountains, Germany : head and
body, 57-3 (51-63) ; tail, -40 -5 (39-42) ; hind foot, 11-5 (11-12).
Average and extremes of four specimens from Stalden, Valais>
Switzerland: head and body, 55-6 (50-61); tail, 42-6 (42-43);
hind foot, 11 "0 (10 "8-11 '2). Average and extremes of seven
specimens from Hatszeg, Hunyad, Transylvania : head and body,
51-3 (47-55): tail, 40-6 (32-46); hind foot, 11-2 (10-12).
For cranial measurements see Table, p. 58.
Specimens examined. — One hundred and twenty-five, from the following
localities ; —
Scotland: Lossiemouth, Elgin, 2; Gordonstown, Elgin, 1 ; Lhanbryde,
Elgin, 2; Grantown-on-Spey, Elgin, 13; Cromlix, Dunblane, 1; Aber-
deen, 1; Dunvegan, Skye, 1; Stornoway, Lewis, Hebrides, 1; Newton,
North Uist, Hebrides, 1 ; Barra Island, Hebrides, 1 ; Stockbriggs, Lanark-
shire, 1 ; Kirtle Bridge, Dumfriesshire, 1 ; Wyseby, Dumfriesshire, 2.
Wales : Aberia, Merionethshire, 2 ; near I3ridgend, Glamorganshire, 1
no exact locality, 1.
England : Spurn Head, Yorkshire, 1 ; Grimsby, Lincolnshire, 3
Waltham, Lincolnshire, 2 ; Thornhaugh, Northants, 2 ; Swithland, LeiceS'
tershire, 1 ; Graftonbury, Herefordshire, 2 ; Crippetts, Gloucestershire, S
Clifton, Gloucestershire, 1 ; Loughton, Essex, 2 ; Wilbraham, Cambridge-
shire, 1 ; Dartford, Kent, 1 ; New Forest, Hampshire, 1 ; Alum Bay, Isle
of Wight, 1 ; Chagford, Devonshire, 2 ; Combmartin, Devonshire, 1 ; South
Molton, Devonshire, 1 ; Lundy Island, Devonshire, 1 ; Isle of Man, 8.
Ireland : Clonbroch, Co. Galway, 1 ; Caragh Lake, Co. Kerry, 1 ; Co.
Longford, 1 ; Knock, Co. Down, 1 ; North Esk, Dunkettle, 2 ; Duncannon,
3 ; Kilkenny, 1 ; no exact locality, 2.
Norway : Holaaker, upper Gudbraudsdal, 1 ; Graven, Hardanger, 1
(U.S.N.M.).
Sweden : Upsala, 1 (U.S.N.M.) ; near Stockholm, 1 ; Svarta, Orebro, 1
(U.S.N.M.).
France ; Barbizon, Seine-et-^Iarne, 1 ; Montr^jeau, Haute-Garonne, 1
(U.S.N.M.); Ax-les-Thermes, Ariege, 2; I'Hospitalet, Ariege, 3; Porte,
ipyren^^es-Orientales, 1.
Germany: Brunswick, 3 (B.M. and U.S.N.M.) ; Bodethal, Harz Mts., 3
(U.S.N.M.); Mauseklippe, Bodethal, Harz Mts., 1 (U.S.N.M.); Frankfort-
on-Main, Hessen-Nassau, 1 ; Niesky, Silesia, 1 ; Marxheim, near Mannheim,
Bavaria, 2 ; near Konigsberg, 2 (U.S.N.3M.).
Austria-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 1 ; Hatszeg,
Hunyad, Transylvania, 7.
Switzerland : St. Cergues, Vaud, 2 (U.S.N.M.) ; Stalden, Valais, 4
(Geneva) ; Untervatz, Grisons, 5 (U.S.N.M. and Mottaz) ; St. Gothard,
Uri, 1.
Italy: Vallombrosa, near Florence, 1.
1. Lossiemouth, Jloray- G. Denson (c & p). 8. 9. 17. 1.
shire, Scotland.
9, 1. Lhanbryde, Elgin, 200ft. W. Taylor (c & p). 8. 9. 3. 1-2.
9. Dunblane, Stirlingshire. W. R. Ogilvie-Grant 8.9.0.1.
(c & p).
1. Aberdeen. Dr. Macgillivray 52. 7. 10. 13.
(c & p).
6. Dunvegan, Skye. J. S. Elliott (c & p). 8. 9. 4. 1.
6. Stornoway, Lewis. R. M. D. Hawker 8. 9. 18. 1.
(0 & P).
1 al. North Uist, Hebrides. J. A. Harvie Brown 79. 9. 18. 1.
(c & p).
1. Barra Island, Hebrides. W. E. de Winton 8. 9. 20. 1.
(c & P).
57
6. Stockbriggs, Lanark-
shire.
1. Bridgend, Glamorgan-
shire, Wales.
1 al. Wales.
9. Grimsby, Lincolnshire,
England.
2. Waltham, Lincolnshire.
2 st. Graftonbury, Hereford-
shire.
9. Wilbraham, Cambridge-
shire. (S. F.Harmcr.)
1. Clifton, Gloucestershire.
(R. I. Pocock.)
6. Lundy Island, Devon-
shire.
2 6. Chagford, Devonshire,
85Gft. (W.P.Stark.)
6, 4 9. Ramsay, Isle of Man.
6. Sulby Glen, Isle of Man.
9. Clonbroch, Galway, Ire-
land.
1 al. Longford.
6 St. Knock, Down.
2 al. Dunkettle, Cork.
2 6,19. Duncannon, Wexford.
(Rev. Dr. Martin.)
1 al. Ireland.
6. Holaaker, Gudbrandsdal,
1900 ft. Norway.
9. Barbizon, Seine-et-
Marne, France.
2 9. Ax-les-Thermes, Ari^ge.
6. V Hospitalet, Ariege,
4700 m.
6. r Hospitalet, Ariege,
1450 m. (A. Robert.)
9. Port6, Pyr6n6es-Orien-
tales, 1600 m.
2. Querum Forest, Bruns-
wick, Germany.
1 al. Frankfort - on - ]\Iaiu,
Nassau.
9. Niesky, Silesia, 181 m.
(W. Baer.)
6. IMarxheim, Bavaria.
1. Csall6k6z-Somorja, Press-
burg, 400 ft. Hungary.
1. Hatszeg, Hunyad, Tran-
sylvania, 2000 -
6500 ft.
3. Hatszeg, Transvlvania,
2000-6500 ft."
3 6. Hatszeg, Hunyad, 1600-
2000 ft.
E. R.Alston (c&p).
R. I. Pocock (c & p).
G. Stokes (c & p).
G. H. Caton Haigh
(c & p).
G. H. Caton Haigh
(c & p).
W. de W i n t o n
(c & p).
Miller Collection.
Miller Collection.
N. H. Joy (c & p).
Miller Collection.
C. H. B. Grant
(c & p).
C. H. B. Grant
(c & p).
H. P. Witherby
(c & p).
Dr. G. E. Dobson
(c & p).
Hon. N. C. Roths-
child (c&p).
Miss O'Keefe (c&p).
G. Barrett-Hamilton
Zoological Society's
Collection.
R. J. Cuuinghame
(c & p).
G. S. Miller (c).
V. Builles (c & p).
G. S. Miller (c).
O. Thomas (p).
G. S. Miller (c).
G. Barrett-Hamilton
Tomes Collection.
Lord Lilford (p).
Lord Lilford (p).
Budapest Museum
(e).
C. G. Danford (c).
C. G. Danford (c).
C. G. Danford (c).
79. 9. 25. 13.
8. 9. 6. 1.
48. 9. 24. 3.
8. 9. 16. 1.
11. 1. 2. 81-82.
96. 4. 28. 20-21.
7. 7. 7. 2850.
7. 7. 7. 3604.
6. 5. 16. 1.
7. 7. 7. 4472,
4474.
8. 9. 7. 1-5.
8. 9. 8. 4.
8. 9. 19. 1.
80. 12. 14. 6.
I. 9. 3. 7.
80. 11. 20. 4.
81. 6. 29. 1.
II. 1. 2. 83-85.
52. 9. 13. 288.
98. 2. 28. 5.
8. 8. 4. 160.
8. 3. 27. 2-3.
8. 8. 4. 159.
8. 9. 1. 45.
8. 8. 4. 161.
11. 1. 2. 86-87.
7. 1. 1. 46.
99. 1. 9. 15.
11. 1. 1. 146.
94. 3. 1. 25.
3. 2. 2. 11.
3. 2. 2. 13-15.
3. 11. 8. 16-18.
58
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60
IXSECTIVOKA
1. St. Gothard, Uri, Swit- Purchased (Brandt). 46. 2. 13. 14.
zerland.
1 al. Vallombrosa, Florence, Dr. G. Cecconi (p.) 1. 8. 2. 4.
Italy.
SoREx MixuTUS LUCANius Miller.
1909. Sorex niinutus Incaniits Miller, Ann. and Mag. Nat. Hist., 8tli ser.,
Ill, p. 417, May, 1909. Type in British ]Museum.
1910. Sorcx minutiis lucanms Trouessart, Fauna Mamm. d'Europe, p. 55.
Type locaUty. — Monte Sirino, Lagonegro, Italy.
Geographical distribution. — At present known from the type
locality only.
Diagnosis. — Similar to Sorex minutus minufus, but with molars
and anterior upper incisor noticeably enlarged
Teeth. — As compared with specimens of Sorex minutus minutus
from a wide range of European localities and also with specimens
from Scalipa and Sumela, Asia Minor, the type of S. minutus
Iwanius is at once recognizable by its enlarged teeth The actual
size is so small that it is impossible to express the differences by
measurements, but to the e3'e the greatei- area of the molar
crowns and the larger more projecting anterior upper incisor
and more robust unicuspids and anterior lower incisor are at
once apparent.
Measurements. — Tail, 42; hind foot, 10 '4. For cranial
measurements see Table, p. 59.
Specimen examined. — The type.
1. IMonteSirino, Lagonegro, 0. Thomas (p). 8. 9. 1. 5.
S. Italy. (.4. Robert.) (Ti/pe of subspecies.)
SOREX ALPINUS Schinz.
(.Synonymy under subspecies.)
Geographical (Jistrihution. — Pyrenees, Alps, Harz Mountains,
Riesengebirge and Carpathians.
Diagnosis. — Size essentially as in Sorex aranvus (head and
body usually 65 to 70 mm.), but tail about equal to head and
body ; colour uniform dark slaty grey, the underparts nearly as
dark as back ; basal lobe of anterior upper incisor sub-terete, the
length of its base about half that of antei'ior lobe.
External characters. — Except for the dark, slaty colour, and
the much greater relative and actual length of the tail, Sorex
alpinus does not differ markedly from Sorex araneus in general
external characters. The fur of the back varies from 5 mm. to
8 mm. in de^^th according to season, and the tail may either be
well covered with closely appressed hairs and provided with a
pencil 4 to 5 mm. long, or practically naked at tip and ver}-
SOREX 61
scantily haired elsewhere, a variation not wholly dependent on
season.* At middle of tail there are about 18 annulations to the
centimeter. Feet relatively larger than in Sorex araneus, but
not peculiar in form.
Colour. — Entire upper parts a deep slaty grey produced by
the combination of blackish-slate under colour and short seal-
brown tips, lightened to a varying degree by silvery sub-terminal
annulations. Underparts slightly washed with sepia, but the
difference in colour of the two surfaces not conspicuous, and sides
entirely without line of demarcation. Feet scantily clothed with
silvery hairs. Tail sharply bicolor, nearly black above and at
tip, buffy whitish below, the light area always less extensive than
the dark, and sometimes reduced on distal half of tail to a
narrow median line.
SJcidl. — The skull, though of the same general dimensions as
that of Sorex araneus, is recognizable by its
less elevated brain-case, the outline of which
when viewed from above is more nearly
circular, owing to the very slight flattening
of antero-external portion of border. Inter-
orbital region less tapering than in the
related species, a character due to the
slightly though appreciably greater lachrymal
breadth. The diameter of the tympanic ring
is usually though not constantly less than in
Sorex araneus. Anteorbital foramen slightl}'
larger than in S. araneus. Lachrymal fora-
men opening over point of contact between .
first and second molars.
Teeth. — Kelatively to size of skull the teeth
are noticeably smaller than in Sorex araneus; this is particularly
evident in the anterior upper incisor when viewed from the side,
and the upper unicuspids when viewed from below. Anterior
upper incisor with basal lobe even smaller than in S. miniitus, its
height slightly less than that of first unicuspid and much less than
that of anterior lobe of incisor. Viewed from below it appears
to occupy decidedly less than half of tooth, while from side it
appears scarcely half as large as first unicuspid, from which it
further diflers in its simple peg-like form. The two teeth come
in contact at about the same level as in S. araneus, but their tips
diverge more noticeably. Anterior lower incisor with shaft
distinctly tapering, its cutting edge with three low, sometimes
ill-defined lobes. Upper unicuspids less robust than in Sorex
araneus, their crowns distinctly longer than broad. The cusp
occupies more than half area of crown, at expense of crushing
area, most of which, except in fifth unicuspid, lies obliquely
instead of horizontally. Viewed from the side the unicuspids
* A specimen in full winter coat, but with the tail bare at tip (B.M.
3. 2. 2. 2), was taken at Hatszeg, Hunyad, Hungary, on December 12.
62
INSECTIVOEA
are all longer in proportion to their height than in Sorex
armieus, and the posterior bordei is more concave. In size there
is a gradual and regular diminution from first unicuspid to third ;
fourth somewhat more abruptly smaller ; "■ fifth slightly smaller
than fourth when viewed from the side, but with larger crown
area due to the presence of a well developed postero-internal
crushing surface. This tooth is rela-
tively larger and more functional than
in Sorex araneus, showing no tendency
to become subordinate to paracone of
large premolar. Lower unicuspids
narrower and more trenchant than in
Sorex araneus. Except for this general
tendency the second shows no special
, . ■ ' . peculiarities of form. The first, how-
Sorex ahnnus. Anterior ^ . ..... ,.,,, , i. , i
teeth. X 5. ever, IS strikingly dinerent irom the
corresponding tooth in S. araneus.
Its general outline when viewed from side is irregularly
elliptical, with longest axis parallel to that of mandible, the
upper edge with two low, rounded cusps, the anterior of which
is about as large as lobes on cutting edge of incisor, the posterior
smaller. Upper cheek-teeth not essentially difl^erent from those
of Sorex araneus, except that hypocones are less developed.
Lower cheek-teeth as in the related species. Pigmentation of
teeth slightly less extensive than usual in Sorex araneus.
Sorex alpinus alpinus Schinz.
1837. Sorex alpinus Schinz, Neue Denkschr. Allgem. Scbweiz. Gesellsch.
Naturwiss., Neuchatel, i, p. 13 (St. Gothard Pass, Switzerland).
1840. ? Sorex antinorii Bonaparte, Iconogr. Faun. Ital., i, fasc. 29 (No exact
localit}-, and probably not a European species).
1857. Sorex alpinus Blasius, Saugetbiere Deutscblands, p. 126.
1870. ? Sorex intermedins Cornalia, Catal. Descrit. IMamm. Ital., p. 27 (Hills
of Brianza, Uomo, Italy). Part : body ; see Sordelli, Atti Soc. Ital.
Sci. Nat. e del Mus. Civ. Stor. Nat., Milano, xxxviii, p. 364, 1899.
1899. ? Sorex alpinus var. longobarda Sordelli, Atti Soc. Ital. Sci. Nat. e del
Mus. Civ. Stor. Nat., Milano, xx^viii, p. 363 (MS. .synonym of
intermedius) .
1910. Sorex alpinus Trouessart, Faune Mamm. d'Europe, p. 50. ^
Type locality. — St. Gothard Pass, Uri, Switzerland.
Geoijrapliical distrihution. — From the Jura and Alps through
Tirol to Transylvania ; Pyrenees. f
* In an adult female from Briinig, Switzerland (No. 85830, U.S.N.M.),
tbe tbird and fourtb unicuspids on rigbt side are fused into a single
2-cusped tootb, wbile tbose on left side are normal.
t I bave seen no Pyrenean specimens of Sorex alpinus. For record of
its occurrence see Trutat, Bull. Soc. Hist. Nat., Toulouse, xii, p. 100, 1878
("massif de la IMaladetta '"). Tbe animal is probably less common in tbe
Pyrenees tban in tbe Alps, as I was unable to find it in several localities
resembling tbose wbere it regularly occurs in Switzerland.
SOREX 63
Diarjnosis. — Skull and teeth of maximum size for the species
(condylobasal length of skull, 19 '4 to 20' 6; upper tooth-row,
8 • 6 to 9 • ; lower tooth-row, 8 • to 8 • 4).
Measurements. — Average and extremes of five specimens from
Briinig, Switzerland: head and body, 74-6 (72-77); tail, 73
(70-75); hind foot, 16. Average and extremes of three speci-
mens from Vitznau, Switzerland : head and body, 72 '3 (69-75) ;
tail, 74-0 (72-75); hind foot, 15-5 (15-16). Adult male from
Vitznau, Switzerland : head and body, 69 ; tail, 68 ; hind foot,
15 '3. Adult female from Chamonix, Haute-Savoie, France:
head and body, 73; tail, 74; hind foot, 15 "4. For cranial
measurements see Table, p. 64.
Specimens examined. — Thirty-one, from the following localities : —
France: Chamonix, Haute-Savoie, 1 (U.S.N.M.).
Switzerland : St. Cergues, Vaud (Jura), 4 (U.S.N.M. and Mottaz) ;
Briinig, 5 (U.S.N.M.); Vitznau, 4 (B.M. and U.S.N.M.); Andermatt, 1
(U.S.N.M.) ; St. Gothard, Uri, 6 (B.M. and U.S.N.M.) ; Untervatz, Grisons,
1 (U.S.N.M.) ; Murgthal, St. Gallen, 1 (U.S.N.M.) ; Santis, St. Gallen, 1
(Mottaz) ; no exact locality, 3.
Austria-Hungary : Hatszeg, Hunyad, Transylvania, 4.
6. Vitznau, 440 m. Swit- O. Thomas (c & p). 5. 8. 3. 15.
zerland.
1. St. Gothard, Uri. Baron E. de Selys- 45. 7. 5. 1.
Longchamps (p).
1, 1 al. St. Gothard, Uri. Purchased (Brandt). 46. 2. 13. 18.
46. 2. 13. 14.
1. St. Gothard, Uri. Purchased (Stock- 46. 6. 2. 32.
holm Museum).
1. Switzerland. E. R. Alston (p). 79. 9. 25. 9.
1. Switzerland. Purchased (Ger- 76. 9. 18. 13.
rard).
9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 2. 2. 2.
sylvania, 2000 ft.
Hungary.
i, 2 9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 12-14.
sylvania, 1500-2000 ft.
SoREX ALPINUS HERCYNICUS Miller.
1909. Sorc.x: alpinus heixynicus Miller, Ann. and Mag. Nat. Hist., Sth ser.,
HI, p. 417, May, 1909. Type in U.S. National Museum.
1910. Sorex alpinus hercynicus Trouessart, Faune Mamm. d'Europe, p. 51.
Tjipe localitij. — Miiuseklippe, Bode Valley, Harz Mountains,
Germany.
(Teograpliical distrihntion. — Harz Mountains and Riesen-
gebirge.
Diagnosis. — Skull and teeth not so large as in typical
SorcK alpinus (condylobasal length of skull, 19 to 19 "6 mm.;
upper tooth-row, 8"2 to 8*6 mm. ; lower tooth-row, 7-8 to 8'0 mm.).
Measurements. — ■External measurements of type : head and
body, 71, tail, 67 ; hind foot, 15*4. Average and extremes of
six specimens from the Harz Mountains : head and body, 73 • 4
64
INSECTIVORA
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,, not worn.
„ slightly worn.
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•q}Sn9[
TBSBqoiiCpuoa
0-HC<I0C-10':tHCD0-*0-« C-l-*Cq-*COCD(MCO
O O O O O O O O O CT O C3 C5 03 0> C5 03 O C5 03
0'llMC<IC^<M(MOq<MCMiHOqi-l T-I.Hi-lTHr-tr-(,Ht-l
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O+O^CH-OOO+CH-OO+OOO* 000-CH-'-0*-o'0*XD
&
inrHCMOOi-H^_r/-i'-,r-)^c-i lOCOOC-l^t-COOl
o lO o in CO CO & S S S ?o ^ 0-1 fM CO on CI 0-1 c<i c-1
-iH -H -41 -* -^H -* S S S S S S C-1 01 CM CM C-1 C-1 C-1 0-1
S33?13?3wSS§SS ;:1 ;=! S ^; ;il ;:J ^1 ;:j
Locality.
^ . ... .2
S. alpinus alpinus.
France : Chamonix
Switzerland : St. Ccrgues, Vai
Vitznan
Briiuig
Andermatt
S. alpinus hercynicus.
Germany : Wolfshau, Silosia
Eulengrund .
Bahrcnberg, Harz
Miiuseklippe ,
65
(72-77); tail, 66-6 (6c-68) ; hind foot, 15-1 (14-S-I5-8)
Average and extremes of five specimens from the Riesen-ebir-e •
ui i'lf.R^r\P'i ^''~"\' ^^^^ ^'^'^ ^•^^■'^^) ' '"^d fo^^t,"
1* J (14 a-lo-U). ior cranial measurements see Table, p. 64.
Spechnens examined. -Eleven (all in U.S.N.M.), from the foUowin-
localities m Germany: Bahienberg, Harz Mts., 2; Bodethal, Harz
w iVi' ^^^^^'^'^l'PP'r.HarzMts.,3; Eulengrund, Riesengebirge, Silesia 3
Wolfshau, Riesengebirge, Silesia, 2. 6 ^^ 'e«. '^uebid, o ,
BemarJc^— In external measurements the Alpine shrew of
the Harz MountaiiLs and Riesengebirge agrees with the Swiss
animal ; but the differences in length of skull and of tooth-rows
seem enough to warrant the recognition of the two forms as
distinct.
Genus NEOMYS Kaup.
1829. Keomus Kanp, Entw.-Gesch u Natiirl. Syst. Europ. Thierwelt, i,
p. 117 (boicv daiibcntomi Erxleben).
1829. LeiccorhyncMcsK^ni,, Entw.-Gesch. u. Natiirl. Syst. Europ. Thier-
Th!, fi'' f • I" ('^•^'•^'^' ^^«^«i"^ Geoffroy and ,S'. Icucodon Hermann •
p lotMatHsol).*""' '" Thomas, The Zoologist, 4th ser., n',
1829. Hydrogalc K^n^, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt i
p. 119 {Sorcx remifer Geof!roy). ^ J-nierweu;, i,
1832. Crossopus Wagler, Isis, p. 275 [Sorex fodiens Schreber)
1835. i7y^ro.or.r Duvernoy, M6m Soc. du Mus. d'Hist. Nat., Strasbourg,
II, p. 19 (&ore.i:/ofZjcws Pallas, see p. 17). °
1835. ^«y./»sor.,r Duvernoy, M^m. Soc. du Mus. d'Hist. Nat., Strasbourg
animal).^ P^r'" " ^ ' ^^""^^ "*" ^°'''' '"''"'''"' tetragonurits
1838. Pinalia Gray, Proc Zool. Soc, London (1837), p. 12G, June 14, 1838
(MS. synonym of Crossop^ts).
1848. Galeviijs Pomel, Archiv Sci. Phys. et Nat., Geneve, ix p 249
November 1848 part, included £V«cfe,/4o,-,,, Duvern^y, cX'ssopus
Wagler, and Pachyiua do Selvs-Lougchamps) ^'o.sopus
1854. Myosidis Pomel Catal. M^.th. Vert. Foss. Bassin de la Loire p 14
/S.)°'' "" ''°' distinguished specifically from Nelmys
1857. Crossopus Blasius, Saugethiere Deutschlands, p. 119
1898. Neomys Thomas, Zoologist, 4th ser., ii, p. 100, March 15, 1898.
Ti/pc species.—Sorex daubentonii Erxlehen = S. fodiens Schreber
(geographical distribntion.—I'ali^.iivctic region north of the
Mediterranean from Spain and England to the Caucasus and
Asia Minor, north to northern Scandinavia.
Characters.— Uppev unicuspid teeth 4-1, their form more pre-
Uensory than in Sorex; dental formula : / ;'=i\ t- tj, jjih --=?, vi ?== = 30 •
posterior lobe of anterior upper incisor 'less 'than' lillf 'as hi-h
as hrst ; anterior lower incisor with one ill-defined lobe on cuttin-
edge near middle of tooth ; third lower molar and second lowe?
unicuspid as in Sorev ; points of all the teeth pigmented; fur
unusua ly soft and den.se, entirely concealing ears ; feet with a
noticeable fringe of elongated hairs at edges of soles and toes
F
66
INSECTIVORA
the liind feet enlarged and turned somewhat outward ; in one
species a longitudinal ridge of stiiit'ened hairs along under surface
of tail ; ear small, completely hidden in the fur, the meatus
closed by two valves, one of which lies on inner surface of
antitragus, the other on inner surface of conch ; habits aquatic.
Semarlcs. — In its external form, slight reduction in the
number of teeth, and in the noticeably prehensory adaptation
of the upper incisors and unicuspids, the members of the genus
Neomys are distinctly more specialized than the species of Soreoc.
They retain, however, the primitive form of the third lower
molar and lower premolar characteristic of Sorcx, these teeth
showing no tendency to the more highly modified structure
found in Crocidura. The members of the genus Neomi/s are the
only Europenri shrews specially adapted to aquatic life. They
are at once recognizable Iw their dense, velvety fur, and large,
distinctly fringed hind feet. In the commonest and most widely
distributed species the tail is provided with a keel of stiffened
hairs, a character not found in any other Eurojiean shrew.
KEY TO THE EUROPEAN FORMS OF NEOMYS.
Tail with median keel of stiffened hairs extending
entire length of under surface N. fodicns, p. 66.
Uuderparts whitish or yellowish, occasionally
brownish (Continental Europe except Iberian
Peninsula) S.f.fodienx, p. 69.
Underparts usually brownish (England and Scot-
land) y.f- Mcoloi; p. 73.
Tail without keel.
Length of tail 47 to 53 mm. ; hind foot 16 to
17 mm. (Alps and Pyrenees) N. milleri, p. 78.
Length of tail 55 to 61 mm.; hind foot 16-8 to
18 mm. (Northern and central Spain) K. anomalus, p. 81.
NEOMYS FODIENS Schreber.
(Synonymy under subspecies.)
Gco[irapMral (listribiition. — From Norway and England to the
Pyrenees, northern Italy and western Siberia. ISTorthern and
eastern limits of range not known.
Diagmi^is. — Tail with keel always present ; feet Aery con-
spicuously fringed ; hind foot usually more than 1 7 mm. ; lachrj'-
mal foramen (.)pening over ^^osterior half of m^ ; anterior upper
incisor robust.
E.iternal cliar(tcfert<. — Fur very dense, its depth at middle of
back about 6 inm. in summer, 9 mm. in winter, its texture soft
and velvety, this ji'^i'ti^-'idarly noticeable on underparts : a
few longer hairs usually present on flanks and rump. Eyes
small and inconspicuous ; ears completely hidden by the fur.
Muzzle less slender than in Sorea- araneiis, the median ridge on
upper lip with distinct wart-like outgrowth which tits between
points of ujjper incisors when mouth is closed. Feet larger than
NEOMYS
67
more widely spaced ardbette;- defined Tf ■'" T''7' ^"'
pencil evident, utally ab™ 3 to°f ° ''™'''"f'='- 1' ■"^l" ^
median li„e «t tail bcLt frL extrameTaie'l r°"'\ "^'""^
and on head mssin<- ilf-Lr ''f ""'"j''""" everywhere evident,
size''!:ht^Iw t"'' ^'^'" ^™^" ^'^^^ «^ ^-- — '"'^ in larger
size leldtnely larger, more inflated brain- "
case (depth decidedly more than half breadth
and lower, more flattened rostrum and
interorbital region. The brain-case is sub-
circular m outhne, though distinctly flattened
antero-externally and postero-exteLallv-in
rSnl-'' ""^"^"'^ °^ '"^^ interJrbital
region by a conspicuous, well deflned anc^le
lu ateral view the proflle of brain-case^ is
S . '''''''''^f posteriorly, low anteriorly
o s ly abrupt curve into the nearly flat dJrsal
V ."'l> iT*"'""'^'*"' '"^Sion and rostrum.
Anteorbital foramen relatively larger and more
Midelv open than in the European species of
hnlTnf ^^"\7"^^^ f^'-'-^n^en over posterior
half of u\ i^Jesopterygoid space decidedly more than half ..
-Ida as long; hamular processes shorter than in^W arteTs
KlQ. 14.
yeomys fodiens.
-Nat. size.
68 INSECTIVORA
and more abruptly turned outward. Foramen ovale appearing
rather conspicuously in region between hamular and postglenoid
process. Rudimentary zygomatic process of maxillary well
developed, essentially as in Sorex araneus.
Teeth. — Though in general not strikingly different from those
of Sorex araneus the teeth show a tendency towards greater
elevation of the cusps as well as a more pronounced hooking
forward of the anterior upper incisor, peculiarities which impart
to the entire dentition, though j^articularly to the teeth lying
in front of the large premolars, an unmistakably prehensive
character. Anterior upper incisor with main hook more projecting
than in Sorex araneus ; posterior talon low though rather long,
its outline when viewed from the side somewhat triangular, its
height less than half that of main cusp and much less than that
of first unicuspid. Lower incisor more slender than that of
Sorex araneus, its cutting edge with only a single low, ill-defined
lobe near middle. Upper unicuspids large and strong, their
crowns when viewed from below decidedly longer than wide,
truncate posteriorly, narrowed an-
terioi'ly and terminating in a slight
though evident point. Jn lateral
view the cusps appear to be more
slender and set further forward
than in Sorex araneus, and the
posterior part of crown is more
produced backward, the cingulum
often rising to a distinct postero-
internal angle or rudimentary cusp,
particularly in first tooth. Spaces
Neomus fodiens. Anterior teeth t , ,. n . . , ,
in profile, x 5. between tips ot unicuspids greater
than in the European species of
Sorex. Each cusp is nearly terete, but with a small postero-
internal concavity continuous with the concave crushing area
which occupies posterointernal fourth of crown. The con-
cave area of cusja and crown is bounded externally by a
low but distinct ridge extending to middle of posterior border
of crown. First and second unicuspids sub-equal ; third about
half as large as first or second though quite similar to
them in form ; fourth not half as large as third, in tooth-
row or crowded somewhat inward, always distinctly visible
from outer side, in contact posteriorly with large premolar.
Lower unicuspids relatively longer and lower than in Sorex
araneus, their form when viewed from the side much as in
Sorex alpinus, except that the first has a very rudimentary
posterior cusp. First somewhat more than half as large as
second, its general outline much the same as in corres-
ponding upper teeth. Second essentially as in Sorex araneus.
Larger cheek-teeth as in Sorex araneus, but main cusps higher
and hypocones better developed. Pigmentation of teeth about
NEOMYS 69
as in Sorex nraneus araneits, the hypocones usually if not always
white.*
Measurements. — In external measurements this species is
unusally variable, the head and body ranging from 72 to 96 mm.,
tail from 47 to 77 mm., and hind foot from 16 to 20 mm. These
differences appear to be for the most part strictly individual.
The skull is more constant : condylobasal length, 1 9 '6 to 22 • 2 mm.;
upper tooth-row, 9*6 to 11 mm. Here the variation is to a
certain extent geographic, as the skull of the British race
averages smaller than that of the Continental form.
SemnrJcs. — Neomys fodiens is the common and widely dis-
tributed water-shrew of Europe. It may always be recognized
by its large size, fringed feet and keeled tail. While the keel
varies greatlj' in depth and distinctness, some trace of it is
always present on basal half of tail, the region in which the
median hairs are never modified in the round-tailed members
of the genus.
Neomys fodiens fodiens Schreber.
1776. Sorex aquaticus P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band,
p. 36 (Prance ; based on Buffon, viii, pi. 11). Not Sorex aquaticus
Linnseus, 175S.
1777. Sorex fodiens Schreber, Saugthiere, iii, p. 571 (Berlin, Germany).
1777. [Sorex] dauhentonii Erxlebeu, Syst. Regni Amm., i, p. 124 (Burgundy,
France ; based primarily on the Musaraigne d'eau of Daubenton,
Hist, de I'Acad. Roy. des Sci., Paris, 1756, p. 42).
1780. Sorex carinatus Hermann in Zimmermann, Geogr. Gesch., ii, p. 382
(Strassburg, Germany).
1792. Sorex liricaudatus Kerr, Auim. Kingd., p. 208 (Strassburg, Germany ;
based on Pennant's account of Sorex carinatus Hermann).
1793. [Sort\r] fluviatilis Bechstein, Gemeinn. Naturgesch. Deutschlands,
III, p. 746 (Suggested but not adopted as preferable to fodiens).
1793. Sorex eremita Meyer, Zool. Annalen, i, p. 323 (Thiiringen, Germany).
1793. Sorex fluviatilis Meyer, Zool. Annalen, i, p. 323 (Published as a
synomym of Sorex eremita, perhaps from Bechstein MS.).
1800. S[orcx] flodiens'] alhus Bechstein, Thomas Pennant's Allgem. Uebers.
vierfiiss. Thiere, ii, p. 723.
1811. Sorex hydrophilus Pallas, Zoogr. Rosso-Asiat., p. 130 (Berlin,
Germany).
1811. Sorex lineatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 181
(Paris, France). Type in Paris Museum.
1811. Sorex rcmifer Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 182
(Abbeville, Somme, France). Tjpe in Paris Museum.
1818. Sorer collaris Desmarest, Nouv. Diet. d'Hist. Nat., xxii, p. 65
(Holland : islands at mouth of Escaut and Jleuse). Described
but not named bv Geoffroy, M^m. Mus. d'Hist. Nat., Paris, I,
p. 309, 1815.
* In an adult male from Geneva, Switzerland (No. 104G Mottaz), the
right upper incisor and first unicuspid and right lower incisor are white
throughout. All other teeth normal.
70
INSECTIVOKA
1822. Sorcx inacrourus Lehmanu, Observ. Zoologies in fauoam Hamburg-
ensem, i, p. 5 (Sachseuwald, near Friedrichsruh, Schles\Yig-Holstein,
Germany).
1826. Sorcx aniphibins Brehm, Ornis, ii, p. 44 (Renthenclorf, Thiiringen,
Germany).
1826. Siyrex nutans Brehm, Ornis, ir, p. 44 (Renthenclorf, Thiiringen,
Germany).
1826. Sorex stagnatilis Brehm, Ornis, ri, p. 47 (Renthenclorf, Thiiringen,
Germany) .
1830. Sorex rivalis Brehm, Isis, p. 1128 (Renthenclorf, Thiiringen, Germany).
1832. Sorex musculus Wagler, Isis, p. 54 (Bavaria, Germany).
1882. Sorex psilurus Wagler, Isis, p. 54 (Bavaria, Germany).
1834. S[ore,r] nigripcs JMelchior, Den Danske Stats og Norges Pattedyr,
]}. 68 (Sielland, Denmark).
1835. Sorex fodu'iis Duvernoy, ilem. Soc. du ilus. d'Hist. Nat., Strasbourg,
II, p. 17. Part : animal, not skull.
1835. Sonw hcrmanni Duvernoy, !\Iem. Soc. du ilus. d'Hist. Nat., Stras-
bourg, II, p. 23 (Strassburg, Germany). Part : skull, not animal.
1838. AmpJiisorc.v linneana Gray, Ann. Nat. Hist., ii, p. 287, December,
1838 (North Bothnia, Sweden).
1839. \ Sorcx fodiens] var. Icucotis de Selys-Longchamps, Etudes de Micro-
mamm., p. 142 (described on p. 25) (St. Gervais, at foot of Mt.
Blanc, Haute-Savoie, France).
1839. ? [Sorcx fodiens] var. albiventris de Selys-Longchamps, Ktudes de
Micromamm., p. 142 (nomen nudum).
1845. ? [Sorcx fodiens] var. nigricans Nilsson, Atti della sesta Riunione
degli Scienziati Italiani, Torino, 1844, p. 357 (Sweden). Nomen
nudum.
1857. Crossopus fodiens Blasius, Saugethiere Deutschlands, p. 120 (part).
1868. Sorex timbriatiis and Crossop)us fimbriaius Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, ]NIath.-Naturwiss. Clas>"e, lvii, pt. 1,
p. 610 (Synonyms of " Cvossopns daubcntonii nigripcs," wrongly
attributed to Giebel, Saugethiere, p. 899). Not Sorex jinibriatus
Wagler, 1832.
1868. Crossopiiis ciliatiis, griscogularis Fitzinger, Sitzungsber. kais. .\kad.
Wissensch. Wien, IMath.-Naturwiss. Classe, lvii, pt. 1, p. 623
(Chartres, Eure-et-Loire, France).
1870. Sorer intermedins Cornalia, Catal. Descrit. IMamm. Ital., p. 27 (HiUs
of Brianza, Como, Italy). Part: tail only (see Sordelli, Atti
Soc. Ital. Sci. Nat. e del 3Ius. Civ. Stor. Nat., Milano, sxxviii,
p. 364, 1899).
1895. Neoniys fodiens Thomas, The Zoologist, 4th ser., ii, p. 100, March,
1908.
1899. Sorcx alpiniis var. longobardus Sordelli, Atti Soc. Ital. Sci. Nat. e
del Mus. Civ. Stor. Nat., Milano, xxxviii, p. 363 (MS. synonym
of intermedius).
1901. Ncomys fodiens minor Miller, Proc. Biol. Soc, Washington, xiv,
p. 45, April 25, 1901 (Moutrejeau, Haute-Garonne, France). Type
in U.S. National Museum.
1905. Crossopus ou Sorcx ignotiis Fatio, Arch. Sci. Phys. et Nat., Genfeve,
4th ser., xix, p. 202, February 15, 1905 (Switzerland). Part:
skull, not mandible. Type in Geneva Museum.
1905. Ncoviys fodiens naias Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., XV, p. 507, May, 1905 (Hatszeg, Hunyad, Hungary). Type
in British Museum.
NEOJIYS 71
1906. Ncomys fodiens nanus Lydekker, Zoological Record, xlii (1905),
Mamm., p. 34, August, 190C (Accidental renaming of naias).
1910. Neornys fodiens, N. fodiens naias and N. fodiens minor Trouessart,
Paune l\Iamm. d'Europe, pp. 56, 57.
Type localiti/. — Berlin, Germany.
Oeographical distribution. — Continental Europe from Norway
to Italy and the Pyrenees.
Diagnosis. — Underparts buify whitish, occasionally tinged with
a stronger shade of yellow or suffused with salmon-colour, and
sometimes though rarely washed with wood-brown, especially on
chest and along median line of belly. Condylobasal length of
skull usually more than 21 mm.
Measurements. — Average and extremes of ten specimens from
Brunswick, Germany : head and body, 86 • 6 (83-92) ; tail, 65 " 5
(58-77)] hind foot, IS '5 (17-19). Average of three specimens
from Montrejeau, Haute-Garonne, France : * head and body, 83
(82-85); tail, 52 (-50-60); hind foot, 17-1 (16-18). Average
and extremes of seven specimens from Luchon, Haute-Garonne,
France: head and body, 75*7 (72-80) ; tail, 53-1 (47-58) ; hind
foot, 16 '4 (16-17). Average and extremes of five specimens
from Porte, Pyrenees-Orientales, France : head and body, 93
(83-96) ; tail, 6''l (57-65) ; hind foot, 18-2(18-19). Average and
extremes of nine specimens from Meiringen, Bern, Switzerland :
head and body, 82-7 (80-86); tail, 66"- 2 (62-71); hind foot,
19 "3 (18—20). Average and extremes of ten specimens from
Hatszeg, Hunj^ad, Transylvania : f head and body, 77 • 7 (72-82) ;
tail, 60 (56-66) ; hind foot, 18-5 (18-19). For cranial measure-
ments see Table, p. 75.
Specimens examined. — One hundred and thirty-four, from the following
localities: —
Norway: Vefsen, Nordland, 1; Kvikne, Hedemarken, 1; Brekke-
bygden, Trondhjem, 1.
Sweden : Bothnia, 1 ; Upland, 1 ; Medstugan, Jemtland, 1 ; Upsala, 1
(U.S.N.M.) ; no exact locality, 1.
Belgium: Waremme, Liege, 3 (U.S.N.M.).
France : Guines, Pas-de-Calais, 2 ; Abbeville, Soiime, 2 (B.!M. and Paris ;
the latter type of remifer Geoffrey) ; near Paris, 1 (Paris; type of lineatus
Gooffroy) ; Dinan, C6tes-du-Nord, 1 ; Cadillac-sur-Garonne, Gironde, 2
(U.S.N.M.); Montrejeau, Haute-Garonne, 3 (U.S.N.M.); Porte, Pyrenees-
Orientales, 5 ; I'Hospitalet, Ariege, 2; Luchon, Haute-Garonne, 7 ; liareges,
Hautes-Pyrenees, 3 ; Barcelounette, Basses-Alpes, 1 ; Charaonix, Haute-
Savoie, 1 (U.S.N.il.) ; Cranves-Sales, Haute-Savoie, 1 : Scientrier, Haute-
Savoie, 1 (Mottaz) ; fitupes, Doubs, 6 (Mottaz) ; no exact locality, 1.
Germany : Brunswick, 10 (U.S.N.M.) ; Saxony, no exact locality, 1
(U.S.N.:\r.); Bahrenberg, Harz Mts., 1 (U.S.N.M.): Frankfort, Wies-
baden, 1 ; Niesky, Silesia, 1 : Strassburg, 1.
Austria-Hungary : Haida, Arva, Bohemia, 3 ; Hatszeg, Hunyad,
Transylvania, 11.
Switzerland : Geneva, 4 (Mottaz) ; Chesieres, Vaud, 4 (Mottaz) ; Les
Plans, Vaud, 3 (U.S.N.M.) ; Meiringen, Bern, 9 (U.S.N.:M.) ; GrindelwrJd,
* " Neomys fodiens minor ^' Miller.
t " Ncomys fodiens naias" Barrett-Hamilton.
72
INSECTIVORA
Bern, 1 (U.S.N.M.) ; Miirren, 2 ; Goschenen, Uri, 2 (U.S.N.M.) ; St. Got-
hard, Uri, 1 (U.S.N.M.) ; Ziiberwangen, St. GaUen, 8 (B.M. and U.S.N.M.) ;
St. Fiden, St. GaUen, 2 (U.S.N.M.) ; Murgsee region, St. Gallen, 5
(U.S.N.M.) ; Au, St. Gallen, 1 (U.S.N.M.) ; Uzwil, St. Gallen, 1 (U.S.N.M.) ;
Sitterwald, St. Gallen, 1 (U.S.N.M.) ; Faido, Ticino, 1 ; Gordola, Locarno,
Ticino, 3 ; Porlezza, Ticino, 1 (Mottaz) ; Muzzano, Ticino, 1 (Mottaz) ; no
exact locality, 1.
Italy : Busalla, Liguria, 3 ; Vallombrosa, Florence, 1.
Remarks. — There is much individual variation in colour as
well as in external measurements and proportions,* though the
size of the skull and teeth, as may be seen from the Tables,
remains very constant. The degree of development of the fringes
on feet and keel on tail is also variable, the depth of the keel in
some individuals being about equal to diameter of tail, while in
others it is too slight to be measured. This is partly, if not
entirely, due to season, as the deepest keels are found in winter
specimens. With regard to the variations in external measure-
ments shown by the averages and extremes in a preceding
paragraph, it is probable that they are much exaggerated by
different methods in taking the measurements. This is well
illustrated by the two series of Pyrenean specimens measured
by A. Robert at an interval of six years. AVhile much
material has been examined it is not wholly satisfactory, con-
sisting chietly of small lots taken by many diflferent collectors.
Eventually it may be necessary to recognize certain Continental
forms such as minor, naias, and the Scandinavian linneana, as
distinct from true fodiens ; but for the present, in view of the
uncertainties concerning external measurements and the striking
similarity of skulls from the entire range of the animal, there
seems to be no other alternative than to regard all the Con-
tinental water-shrews with keeled tail as belonging to a single
1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 1.
way. Waldo (c & p).
6. Kvikne, Hedemarken, G. Barrett-Hamilton 11. 1. 2. 88.
2200 ft. (A'. F. Ticc- (p).
hurst.)
9. Brekkebygden, Trond- G. Barrett-Hamilton 11. 1. 2. 89.
hjem (A'. F. Ticehurst.) (p).
1. Bothnia, Sweden. Purchased. (Wahl- 38. 9. 24. 14.
berg.)
?. Medstugan, Jemtland, Lord Lilford (p). 11. 1. 1. 147.
550 m. (G.Kolthoff.)
2 9. Gaines, Pas-de-Caiais, 0. Thomas (c \- p). 94. G. 6. 1-2.
France.
6, 2 ?. Porte, Pyr6n6es-Urien- 0. Thomas (p). 8. 9. 1. 39-41.
tales, 1600-1700 m.
France. (A. Robert.)
* This tendency to variation, coupled with the animal's peculiarly
attractive and interesting appearance, has led to the creation of the most
formidable synonymy (33 distinct names) yet applied to a European
mammal.
73
9.
6,2.39.
2 6.
2 9.
6 6,i9.
2.
4:9.
9.
3 6.
i.
(5,2 9.
1 al.
L'Hospitalet, Ariege,
1450 m. (.4. Robert.)
L'Hospitalet, Ariege.
Luclion.Haute-Garonue,
G00-900m. (A.Robert.)
Bareges, Hautes-Pyr6-
n6es, 1300-1500 m.
Barcelonnette, Basses-
Alpes. (C. Mottaz.)
Cranves-Sales, Haute-
Savoie.
Abbeville, Somme.
Frankfort, Wiesbaden,
Germany.
Niesky, Silesia, 181 m.
(ir. Baer.)
Strassburg, Alsace.
(C. Mottaz.)
Haida, Bohemia,
Austria.
Hatszeg, Hunyad, Tran-
sylvania, 1500 ft.
Hungary.
Hatszeg, Transylvania,
1500-2000 ft. "
Miirren, Bern, Switzer-
land.
Ziiberwangen, St.Gallen.
[E. Zollikofer.)
Faido, Ticino.
Locarno, Ticino.
Switzerland.
Busalla, Liguria, Italy.
Vallombrosa, Florence.
0. Thomas (p).
G. S. Miller (c).
O. Thomas (p).
G. S. MiUer (c).
0. Thomas (p).
A. Robert (p).
V. BaiUon (c).
Dr. Dieffenbach (p).
Lord Lilford (p).
O. Thomas (p).
Lord Lilford (p).
C' G. Danford (c).
C. G. Danford (c).
\V. Gurtner (p).
0. Thomas (p).
0. Thomas (c & p).
0. Thomas (c & p).
Purchased (Parreys).
O. Thomas (c & p).
Dr. G. Cecconi (p).
8. 9. 1. 42.
8. 8. 4. 139.
G. 4. 1. 12. 14-
18.
8. 8. 4. 13G-
137.
8. 8. 10. 13.
5. 4. 9. 2.
54. A.
47. 1. 8. 46.
99. 1. 9. 8.
8. 8. 10. 12.
11. 1. 1. 148-
149.
3. 2. 2. 1.
3. 11. 8. 2-11.
92. 10. 5. 1-2.
4. 4, 5. 48-51.
5. 8. 2. 20.
5. 8. 2. 8-10.
46. 6. 15. 61.
5. 8. 3. 16-18.
1. 8. 2. 1.
Neomys fodiens bicolor Shaw.
1791. Sorex bicolor Shaw, Naturalists' Miscellany, ii, pi. 55, named in
index (Oxford, England).
1805. Sorex ciliatus Sowerby, British Miscellany, xlix, p. 103 (Norfolk,
England).
1837. AmpMsorcx pennantiC^T&y , Proc. Zool. Soc, London, p. 125 (England).
1840. Crossopus soiverbyi iBonaparte, Iconogr. Faun. Ital., i, fasc. 29, in
text under C. fodiens.
1857. Crossopus fodietis Blasius, Saugethiere Deutschlands, p. 120 (part).
1905. Nemnys fodiens ciliatus Barrett-Hamilton, Ann. and Mag. Nat. ffist.,
7th ser., xv, p. 508, May, 1905.
1910. Neotnys fodiens ciliatus Trouessart, Faune Slamm. d'Europe, p. 56.
Ty2)e locality. — Oxford, England.
Geofjraplncal distribution. — Great Britain.
Diagnosis. — Similar to Neomijs fodiens fodiens, but underparts
usually washed with wood-brown. Condylobasal length of skull
usually less than 21 mm.
Measurements. — External measurements of two adult females
from Scotland (Grantown-on-Spey and Cortachy, Forfar) : head
74
INSECTIVOEA
and body, 83 and 71 ; tail, 57-5 and 53 '5 ; hind foot, IS and 17.
Adult male from Wellington, Somerset : head and body, 75 ;
tail, 48 ; hind foot, 17. Adult male from Yalding, Kent: head
and body, 82 : tail, 52 ; hind foot, 17. For cranial measurements
see Table, p. 77.
Specimens examined. — Forty-six, from the following localities :—
Scotland : Gordonstown, Elgin, 1 ; Dunphail, Elgin, 1 ; Grantown-on-
Spey, Elgin, 1 (Wilson) ; Cortachy, Elgin, 1 (Wilson) ; Cromlis, Stirling-
shire, 1 ; Knibtuck, Stirlingshire, 1 ; Aberdeen, 1 ; Penkill, Ayrshire, 1.
Wales : Fishguard, Pembrokeshire, 1 ; no exact locality, 1.
England : ilill-on-Tyne, Northumberland, 1 ; Leeds, Yorkshire, 1 ;
Wellington, Somerset, 1; Great Grimsby, Lincolnshire, 2; Shaftesbury,
Dorset, 1; Halesworth, Suffolk, 1; Camliridgeshire, 2; Thornhaugh,
Northampton, 1 ; Podingtou, Wellingborough, Northamptonshire, 1 ;
Henley-on-Thames, Oxfordshire, 2; Drinkstone Park, Bury St. Edmunds,
Suffolk, 1 ; Woolpit, Bury St. Edmunds, Suffolk, 3 ; Halesworth, Suffolk, 1 ;
Cambridgeshire, 2 ; Epping, Essex, 1 ; Banstead, Surrey, 3 ; Esher, Surrey,
1 ; Garratt Park Lake, Surrey, 1 ; Godalming, Surrey, 1 ; Tillingbourne
Stream, Surrey, 1 ; New Forest, Hampshire, 4 ; IBasingstoke, Hampshire, 2 ;
Hampshire, no exact locality, 2; Yalding, Kent, 1.
6. Dunphail, Elgin.
1 al. 9 Aberdeen, Scotland.
1. Penkill, Ayrshire.
5. Fishguard, Pembroke-
shire, Wales.
1. Wales.
9. Mill-on-Tyne, North-
umberland, England.
1 al. Wellingborough, ISTorth-
amptonshire.
1. Halesworth, Suffolk.
9. Cambridgeshire.
1. Cambridgeshire.
(J. Baker.)
1. Epping, Essex.
2 al. Henley, Oxfordshire.
1 al. Bury St. Edmunds,
Suffolk.
Sal. Bury St. Edmunds,
Suffolk.
i al. Esher, Surrey.
9 al. Crodalming, Surrey.
S, 3 ?. New Forest, Hampshire.
2 ?. Basingstoke, Hampshire.
2 al. Hampshire.
INIiller Collection.
Hon. N. C. Roths-
child (p).
E. R. Alston (p).
Y. H. Mills (c A- p).
S. Stokes (c & p).
Rev. H. H. Slater
(c & p).
R. R.Orlebar (c&p).
Rev. W. R. Tate
(c & p).
J. Baker (c & p).
E. R. Alston (p).
J. Baker (p).
W. Royal Dawson
(c & p).
.J. H.Powell (c&p).
Duncan Parker (p).
E. Dalgleish(c&p).
W. T. Blanford
(c & p).
Miller Collection.
Miller Collection.
E. Bartlett (p).
7. 7. 7. 2Si8.
10. 11. 28. 1.
79. 'J. 25. 8.
11. 1. 3. 376.
48. 9. 24. 4.
11. 1. 3. 375.
84. 4. 21. 1.
87. C. 7. 1.
39. 9. 29. 26.
79. 9. 25. 7.
40. 4. 11. 9.
11. 1. 3. 377.
83. 8. S. 1.
84. 5. 16. 1-3.
11. 1. 3. 378.
94. 8. 5. 2.
7. 7. 7. 2845-
2849.
7. 7. 7. 4470-
4471.
74. 11. 24. 1-2.
75
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76
INSECTIVORA
Observations.
go
c
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o
d
worn.
toly worn,
worn,
vorn.
o " ^ -
fl y m
,ja
o
, sligbtlj
, modera
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, much \
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o
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4. 4. 5. 49
4. 4. 5. 50
4. 4. 5. 51
86497
124575
85788
123618
Locality.
to
n •
IS
. '^
' - ::
d -
>
• • • g
France : "Etupes, Doub
Germany: Brunswick
Strassburg
Switzerland : Geneva
9
:2 "
o
St. Galle
Giischene
Anderma
Upper M
77
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o . . . . o ^ . . .^^ .2 . . a r r - s r
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78
IXSECTIYORA
NEOMYS MILLERI Mottaz.
1907. Ncomys milleri Mottaz, !Mtm. Soc. Zool. de France, xx, p. 22,
September 20, 1907.
1910. Neomys milleri Trouessart, Fauna Mamm. d'Europe, p. 58.
Tyjw localitij. — Chesieres, Alpes A^audoises, Switzerland.
Altitude, 1,230 m.
Geographical (listrihitlon. — Pyrenees and Aljjs ; Hungary.
Details of distribution not j^et known.
Diagnosis. — Slightly smaller than Neomys fodiens : tail with
keel absent, oi* represented by a slight lengthening of hairs on
under side of terminal third only ; fringes on sides of feet not
conspicuously developed; hind foot usually less than 17 mm.;
tail usually less than 55 mm. ; lachrymal foramen opening over
point of contact between m^ and vf ; anterior upper incisor
slendei".
External characters. — Similar to Neomys fodiens, except that
the feet are less fringed and the tail lacks the definite keel on
under side. When unworn the hairs on ventral surface of tail
are slightly longer than those on upper side, and on terminal
third this elongation is occasionally sufficient to produce a
rudimentary keel ; but there is never any trace of a well-
defined ridge extending to base of tail. Caudal annulations
slightly less distinct than in Neomi/s fodiens. Mamma^ : a 1-1,
i 4_4 = 10.
Colour. — The colour is similar to that of typical specimens of
Neomys fodiens fodiens. Among the skins examined none shows
any noticeable wash of lirown or suffusion of buff on underparts.
Skull and Teeth. — The skull closely resembles that of Neomys
fodiens, but may be distinguished by its smaller general size and
relatively lower, more slender rostrum ; lachrymal foramen over
point of contact of m^ and nr. Teeth smaller throughout, a
difference especially noticeable in the anterior upper incisor.
Upper unicuspids with crowns longer and narrower than in
Neomys fodiens, the long posterior portion especially noticeable
in lateral view by comparison with height of cusp ; cingula
not so well developed as in the related species, and seldom, if
ever, forming a postero-external cusp.
Measurements. — External measurements of type: head and
body, 7G ; tail, 59 ; hind foot, 16 ; hind foot, including claws,
17. Average and extremes of ten specimens from the tyjje
locality: head and body, 77-4 (71-87); tail, 53-7 (50-59);
hind foot, 15 "8 (15-16 '2). Average and extremes of six
.specimens from Untervatz, Grisons, Switzerland : head and body,
79-3 (75-82); tail, 46-8(45-51); hind foot, 14-16 (14-15-4).
Two adult males from Locarno, Ticino, Switzerland : head and
body, 86 ; tail, 50 and 53 ; hind foot, 16 and 17. Average and
extremes of three adult females from 1' Hospitalet, Ariege,
NEOMYS 79
iooc 10 t (10 4-16). Adult male from Barec^es Hanfp«
Pyrenees: head and body, 76; tail, 56; hind frtf 16-4 For'
cranical measurements see Table, p. 80.
Specimens c.ra»M»ie(Z.— Thirtv-thrpp frnm fv,^ j^^n •
Switzerland: Near Geneva i V ^ *^^ f°H?'^i?S ^"''^^''^^s :-
Chesieres, Alpes Vaudoises 10 m M T^T'Jf' y^^*^' ^ (U.S.N.M.) ;
(U.S.N.M.); Untervl^SSois Su's N M f f 'f ^ = Meiring^n, Bern/i
(B.M. and V.S.N.M.) ; Sorengo Ti ino fm S N An .\""^*^"' ^t- Gallen, 3
Italy: Porlezza, Como. 1 (Mottaz) ^^■'^•^•^^'^•) ' L°°amo, Ticino, 2.
Fr™ ■^"^S-"^''-'/,^''^"'''^'-'' Northern Hungary 1
HaSe^^Tren^efr'*^'''' ^"'^^' '^ ^ ^-^°-' Haufe-Garonne, 1 ; Bareges,
complete absence T t;^ trie keetl^ the'Tr'T, '^-^
elongation of the hairs in'^meTan.^giorwi ^^^'
ThTsir r r;r- ^^ ^^- p--^^- s Uiroftj
S^sf^faji^hr^^^^^
Sa'S^'a^ttMr^™ l^^^r^^^ a^l^t:;; of
Their bnC. 1 " ^ specialized than iV^eo,««s /bf?/e„«
of the c 1^;^^^^^^^^^^^^^^^ t-; thoTe
locaHties where ^^^i^:^^^^^' ^^^ '^^^^ ^^^^^her in
''• "^Fratt^'^*' ^"'°^' G-S-^IillerCc). 8.8.4.138,140.
?. Chesjeres Alpes Vau- C. Mottaz (p) r, q r a
doises,1200 m. Swit- ^^ °- ^- "• ^•
zerland.
^ (^- -^^ Zollikofer.)
'^Sn\^"''^°' ^"^*- 0- Thomas (c&P). 5.8.2.6-7.
1. Zubere,-., Hungary. Budapest Museum 94 3 1 ^G
(e).
Ha^eiy^ILSf Srerif ituu^d^Trrtwi'^'^^P'^^^*' ^^''^ ^^ ^^-^oCS.
were taken alternateh in the sameTr°urs'^t n^H^'^T *°^^ ^^'^
streams. In the original descrint^on nif^ ■ ^\^ """^^^ °^ mountain
observes (p. 23) that according to^i°"llr.!^ \^ov,eyer, Mr. Mottaz
is the more inclined trwander awav fiom f hi ' *^^- ^.'^"'^d-tailed animal
The habits of .V. «.w,„a? fsin SI CenTral ^n"'"'''^'^'" vicinity of water,
aquatic as those of N. fodiens ^^'^ ^^P^'"' *° ''^ »« «t"ctlv
so
INSECTIVOEA
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PACHYURA
81
NEOMYS ANOMALUS Cabrera.
1907. Neomys anomalus Cabrera, Ann. and Mag. Nat. Hist., 7th ser., xx,
p. 214, September 1, 1907.
1907. Neomys anomalus Cabrera, Bol. Real Soc. Espaii. Hist. Nat., Madrid,
VII, p. 224. Published November, 1907.
1910. Neomys fodiens anomalus Trouessart, Faune Mamm. d'Europe, p. 57.
Type locality. — San Martin de la Vega, Madrid, Spain.
Oeographical distribution. — Northern and central Spain.
Diagnosis. — Similar to Neomys milleri, but with longer tail
(usually over 55 mm.), and larger hind foot (16 "8 to 18 mm.).
MammiB : a 1 — 1, i 4 — 4 = 10.
Measurements. — Type specimen, male (from Cabrera) : head
and body, 73 ; tail, 60 ; hind foot, 17 "5. Average and extremes
of nine sj^ecimens from Silos, Burgos, Spain : head and body,
83-2 (76-88); tail, 60 (56-61); hind foot, 17-4 (17-18).
Adult male and adult female from Barracas, Castellon, Spain :
head and body, 99 and 100; tail, 53 and 52; hind foot, 17.
For cranial measurements see Table, p. 82.
Specimens examined. — Tweuty-two, from the following localities in
Spaia: Silos, Burgos, 9; La Granja, Segovia, 2; Barracas, Castellon, 2 ;
L^rida, 9.
Remarlcs. — The Spanish water-shrew is well differentiated
from Neomys milleri by its longer tail and larger hind foot. As
yet no member of the keel-tailed group has been found in the
Iberian Peninsula, though it is probable that N. fodiens occurs in
the region north of the Ebro, and perhaps also in the Asturias.
2 (5, 7 9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 22-30.
2 al. La Granja, Segovia. M. dela Escalera (c). 8. 7. 30. 6-7.
6,9. Barracas, Castellon. O. Thomas (p). 8.2.9.41-42.
(N. Gonzalez.)
Genus PACHYURA de Selys-Longchamps.
1889. Pachyura de S^lys-Longchamps, Etudes de Micromamm., p. 32
(Sorex ctruscus Savi). Sub-genus of Crocidura.
1857. Pachyura Blasius, Saugethiere Deutschlands, p. 147 (Sub-genus of
Crocidura).
1897. Pkrodus Schulze, Mamm. Europ., in Helios, Abhandl. u. Vortraga
Gesammtb. Naturwiss., xiv, p. 90 (Crocidura suaveolens Blasius
= Sorex ctruscus Savi).
Type species. — Sorex etruscus Savi.
Geographical distribution. — ^Africa and warmer portions of
Asia ; in Europe confined to the Mediterranean region.
Diagnosis. — Like Crocidura (p. 86), but with upper uni-
cuspids 4-4, the dental formula : i '"*, c ^i, pm ^3^, m -^"^ = 30.
Remarlcs. — Although perhaps not forming a natural group,
the 30-toothed species of Crocidura may for convenience be
G
82
INSECTIVORA
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PACHYURA
83
treated as ineinbers of a distinct genus. The only European
species is immediately recognizable by its excessively small size.
PACHYURA ETRUSCA Savi.
1822. Sorex ctrnscus Savi, Nuovo Giorn. de' Letterati, Pisa, i, p. 60 (Pisa,
Italy).
1835. [Sorex] pachyunis Kiister, Isis, p. 77 (Cagliari, Sardinia).
1841. Pachyura etrusca Bonaparte, Iconogr. Faun. ItaL, i, Indice disfcrib.
1857. Crocidura suaveolens Blasius, Siiugethiere Deutschlands, p. 147.
1910. Pachyura etrusca Trouessart, Faune Mamm. d'Europe, p. 43.
Tfipe locality. — Pisa, Italy.
Geographical distribution. — Italy and adjoining portions of the
Mediterranean region.* Limits of range not known.
Diagnosis. — Much smaller than Crocidura russula (head and
body, 3.5-40 mm. ; hind foot, 7 ' 6-8 mm.), but with tail (25-
30 mm.) relatively longer, its ratio to head and body about
70 ; skull excessively small, its cond3'lobasal length only about
12 '8 mm., the dorsal profile nearly straight from front of nasal
to back of parietals, usually a little concave in interorbital
region, the brain-case narrow but even more flattened than in
Crocidura leucodon and C. mimula ; teeth not conspicuously
difiei'ent from those of the smaller European species of Crocidura,
except for the presence of the fourth upper unicuspid, and their
much smaller size (maxillary tooth-row about 6 mm.).
External characters. — ^In general external form Pachi/ura
I'trusca does not differ notably from Crocidura rus.mla except in
the relatively longer tail. The weight of a full grown individual
is, however, probably not more than one-fourth or one-third that
of an adult C. russula, and the head and feet are equally small
in proportion, a character by which Pachyura etrusca mav be
easily distinguished from young individuals of the larger animal.
Mamma;, i 3 — 3 = 6.
Colour. — Upper parts a uniform slaty brown, perhaps best
described as drab-grey washed with light bister, the indi^■idual
hairs slate-grey at base. Underparts faintly contrasted, rather
pale drab-grey with silvery reflections in certain lights. Feet
dull light slaty grey not noticeably contrasted with back. Tail
like back above, usually becoming somewhat darker toward tip,
lighter and more like belly below, but with no evident contrast
between the colours of the two surfaces.
Skull. — Apart from the small size the skull differs from that
of its European allies in .several important characters. Most
noticeable among these is the extreme flatness of the dorsal
profile, which is essentialh' straight from nares Lo occipital,
usually a little concave in interorbital region, while in the
European foims of Crocidura it is usually a little convex
throughout, and never evidently concave in interorbital region.
* Spain. See footnote, p. 8G.— O. T.
84 INSECTIVORA
The l)rain-case is excessively flattened, relatively more so than in
Crocifhira miviula, so that there is less contrast between its depth
and that of rostrum. This flattening is not accompanied by any
increase in breadth of brain-case, the ratio of
^^^T^ which to condylobasal length is about 46,
J\^^;^a^ slightly less than in Crocidura mhimla. Ante-
^^^ orbital foramen relatively large, its position
^ « normal. Lachrymal foramen over posterior
f\ i\ ^^^^^ "^^ '"'■ P'^.late relatively short and wide.
/ s M^-\ Mesopterygoid space relatively longer than in
( I ) W%] ^^^® European forms of Crocidura, its anterior
\gi>/ Xjdy border on line with jjosterior edge of third
Fig. 16. niolar instead of distinctly behind it, its lateral
Nat. size. borders nearly f)arallel, though slightly con-
verging posteriorly : hamular excessively deli-
cate, bowed outward and upward, its length relatively greater
than in C. russula, and about equal to greatest breadth of fossa.
Mandible not peculiar except for its small size (length about
7 mm. or less), and very delicate structure.
Tei'th. — While in general, and aside from the presence of
the fourth unicuspid, the teeth do not differ materially from
those of the smaller European Crocidurse except in size, they
show several jjeculiarities in form. The anterior upper incisor
projects more strongly forward, and has the anterior cusp rather
shorter than in the species of Crocidura, while the posterior cup
is more distinctly separated from the cingulum, these two
characters together imparting to the tooth a form somewhat
suggesting that assumed in Sorex. First unicusjaid relatively
larger and higher than in Crocidura rus.mJa, its cingulum less
curved, and its width distinctly exceeding that of palate instead
of barely ecjualling it. Contrast in both height and crown area
of first and second unicuspids greater than in C. runi<n1a, the
area of second about one-third that of first instead of evidently
more than one-third. Third unicuspid slightly larger than second.
Fourth about half as large as third, crowded inward from the
tooth-row, but visible from the outside through space separating
third unicuspid from large premolar, the width of this space
rather more than half diameter of crown of fourth unicuspid.
Large premolar as in C. Jeucodon, except that cutting blade is
shorter and even higher, and posterior border of crown is less
concave. Upper molars with hypocones less distinct than in
C. ruamla, but otherwise not showing any tangible peculiarities.
Mandibular teeth essentially as in Crocidura russula and C. leu-
codon, but posterior section of third molar even more reduced.
Measurements. — Average and extremes of six specimens from
Florence, Italy: head and body, 38-5 (36-42); tail, 27-1
(26-28) ; hind foot, 7-7 (7 • 6-7 • 8). Adult male and female
from near Turin, Italy : head and body, 42 and 40 ; tail, 28 and
27 ; hind foot, 7 • 8 and 7 • 4. Adult male and female from near
PACHYDRA
85
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86
INSECTIVOEA
Genoa, Italj- : head and body, 41 and 42 : tail, 26 and 28 ; hind
foot, 7 • 6 and 8 • 0. Adult female from Hassari, Sardinia : head
and body, 41 ; tail, 29 ; hind foot, 7*8. For cranial measure-
ments see Table, p. 85.
Specimens examined. — Twenty-nine, from the following localities : —
Italy : Near Turin, 2 (Genoa) ; Perti, Finalborgo, 2 (Genoa) ; near
Genoa, 5 (U.S.N.M. and Genoa) ; Pisa, 5 (B.il. and Mottaz) ; Florence, G
(Li .S.N.M. and JMottaz) ; Tuscany, 2 ; Rome, 1 (Genoa) ; no exact locality, 2.
Sicily : No exact locality, 2.
Saedinia: Assuni, 1 (U.S.N.M.). No exact locality. 1.
Greece : Lamia, 1.
lal.
Pisa, Italy.
Marquis G. Doria (p).
G9, 3. 4. 1.
2al.
Tuscany.
(No hist
Dry).
lal.
Italy.
Zoological Society's
Collection.
55. 12. 26. 295.
Skeleton.
Italy.
Purchased.
58. 10. 21. 13.
6, ? al.
Sicily.
Purchased (Parzu-
daki).
62. 2. 26. 22-23
lal.
Sardinia.
Hon. N. G. Roths-
child (1').
11. 10. 1. 1.
1 al.*
Lamia, Greece.
E. MacDonell (p).
8. 7. 22. 1.
Genus CROCIDURA Wagler.
1832. Crocidiira Wagler, Isis, p. 275 (Soivx leucudun Hermann).
1857. CrocidiDxi Blasius, Saugethiere Deutschlands, p. 137.
1869. Leucodan Fatio, Faune Vert. Suisse, i, p. 132 (substitute for
Crocidura).
1897. Paurodus Schulze, Mamm. Europ., in Helios, Abhandl. u. Vortrage
Gesammtb. Naturwiss., xiv, p. 90 (Sorcx Icucodon Hermann and
iS'. arnncus Schreber = .S'. russulus Hermann).
Ti/pe species. — Sorex leucodon Hermann.
GeograjjJiical disirihufion. — Africa and warmer portions of
Europe and Asia, including the Malay Archijjelago ; in Europe,
north to northern Holland and central (Germany, west to the
Atlantic coast and the Channel Islands. Ab.sent from Great
Britain and Ireland.
Characters. — Upper uuicusjjid teeth o-o (dental formula :
' ^ '» ^ il'' i"'* ^' ™ %Er — '^^ '■> posterior lobe of upper incisor
noticeably less than half as high as main cusp ; anterior lower
incisor without lobes on cutting edge ; third lower molar with hypo-
conid and entoconid coalesced, so that form of tooth is strikingly
different from that of other molars, its crown 4-cusped instead
of 5-cusped ; second lower unicuspid without I'udimentary second
cusj) and commissure ; teeth white throughout ; skull lieavier and
more robust than in Sore.v and Neomijs, with conspicuously deeper
rostrum and less contrast between width of brain-case and that
of anterior portion ; rudimentary zygomatic process of maxillary
obsolete; external form rather heavy; ear rising noticeably
* <^, 2 9. Marismas, Lower Guadal- A. Chapman (c & p). 11.12.19.1-3.
quivir, Spain.
Received December, 1911. — 0. T.
CROCIDURA
87
above, t'ur, the meatus closed by two valves as in Neomys ; tail
somewhat thickened, its surface covered with short hairs, among
M^iich are sprinkled numerous longer ones ; habits terrestrial.
Beiiutrkn. — Though not highly modified in external pecu-
liarities the genus Crocidura is the least primitive group of
shrews occurring in Europe. This is indicated by the reduced
number of upper unicuspids, but is more clearly shown by the
highly modified form of the third lower molar and the completely
unicuspid character of the lower premolar. Among the European
members of the family Soricidse the species of Crocidura may
at once be recognized by their moderate size, entirely white teeth,
large ears, and by the presence of long, loosely spreading hairs
scattered over the surface of the tail. Badly prepared specimens,
in which the true characters are obscured, may often be detected
among skins of Sorcx by the noticeable silvery reflections on
hairs of back. Eight species are now known to occur in western
Europe, while the number of forms thus far described from other
portions of the range of the genus is not far from one hundred.
KEY TO THE EUROPEAN FORMS OF CROCIDURA.
Tail decidedly more than half as long as head and
body, the ratio normally varying from 70 to 80.
Hind foot about 14 mm. (Sicilv) C. caudata, p. 110.
Hind foot 11 to 12-5 mm.
Upper tooth-row about 8'6 mm. (Corsica) C. ci/rnensis, p. 111.
Upper tooth-row about 8 mm. (Balearic Islands)... C. balearica, p. 112.
Tail about half as long as head and body, the ratio
normally varying from 40 to 60.
Condylobasal length of skull 16 to 17 '6 mm.
Second upper unicuspid with crown area about
equal to that of third (Crete) C. canem, p. 109.
Second u^sper unicuspid with crown area distinctly
less than that of third C. mimiila, p. 94.
Colour tending to be pale and greyish (Basque
Provinces, Spain) C. m. cantabra, p. 99.
Colour tending to be dark and brownish.
Brain-case not decidedly flattened (Charente,
France) C. »«. kiUisma, p. 98,
Brain-case decidedly flattened (entire range
of species, western portion excepted)... C. 711. mimula, p. 95.
Condylobasal length of skull 17-6 to '20 '4 mm.
(usuallv more than 18 mm.).
Brain-case about half as high as wide or less.
Animal conspicuously bicolor ; condylobasal
length of skull usually more than 19 mm.
(Central Europe, south into Italy) C. h'ucodon, p. 88.
Animal obscurely bicolor ; condylobasal length
of skull usually 18 to 19 mm. (Sicily) C. sicula, p. 108.
Brain-case more than half ashigb as wide C. ritsstila, p. 99.
Condylobasal length of skull usually 19 to
20-4 ram. (Central Europe, south into
Italy) C. r. russiila, p. 101.
Condylobasal length of skull usually 18 to 19 mm.
Colour a dark, coppery-brown (Portugal) C. r. eintrte, p. 108.
Colour a pale drab-ljrown (Spain and south-
western France) C. r. jmichra, p. 103.
88
INSECTIVORA
CROCIDURA LEUCODON Hermann.
1780. Soirx leucodon Hermanu in Zimniermann, Geogr. Gesch., ii,
p. 382 Cvicinity of Strassburg, Germany).
1781. Sorcx leucodon Hermann in Schreber, Saugtbiere, pi. clix.
1782. Sor[ex] leucodon Hermann, Tabula Affinitatum Animalium, p. 79
(footnote).
1792. Sorcx alhipes Kerr, Anim. Kingd., p. 208 (based on Pennant's account
of S. Ieucodo7i Hermann).
1832. Croc[idura] leucodon Wagler, Isis, p. 275.
1857. Crocidura leucodon Blasius, Siiugetbiere Deutscblands, p. 140.
1869. Leucodon niicrourus Fatio, Faune Vert. Suisse, i, p. 173 (Substitute
for leucodon).
1897. [Crocidurci] leucodus Schulze, Mammalia Europsea, p. 18 (Substitute
for leucodon).
1910. Crocidura russula leucodon Trouessart, Faune Mamm. d'Europe, p. 44.
TyjK' locallfj/. — Vicinity of Strassburg, Germany.
Oeograpliical distribution. — Central Europe, from Belgium to
Hungary ; south into Italy. Not known from the Iberian
Peninsula.
Diagnosis. — Size large (among the European species) ; hind
foot, 1 1 to 13 mm. ; condylobasal length of skull, 18-4 to 20 mm. ;
upper tooth-row, 8 "8 to 9*0 mm. ; tail short, its actual length
28 to 38 mm., its ratio to head and body usually ranging from
38 to 43 ; skull with brain-case noticeably depressed, its height
less than half its width ; large upper premolar with antero-
external cusp rather large, its height usually greater than that
of first unicuspid ; colour of underparts whitish, strongly con-
trasted with rather slaty brown of back, the line of demarcation
along sides well defined.
External characters. — Fur shorter and more dense than in
Sorcc araneus, the length of hairs at middle of back about
3 • 5 mm. in summer, 5 mm. in winter, its texture not .specially
modified ; a few slightly elongated hairs (8 mm.) on flanks and
across rump. Eyes small and inconspicuous ; ears small but
rising conspicuously above fur, the two well developed valves a
conspicuous feature of the ear in freshly killed specimens. Feet
not peculiar in form ; less slender than in Sorcx araneus, finely
pubescent on dorsal surface and on lateral portions of posterior
half of sole ; fingers proportioned as in S. araneus, but graduation
less, especially in hind foot ; pads 6-6, those on palm distinct
though somewhat crowded, the surface of palms and soles finely
rugose between the pads. Tail less slender than in Sorcx and
Neomi/s, nearly terete or with under side somewhat flattened,
its length equal to about half that of head and body, its hairs of
two kinds : (a) finely appressed hairs less than 1 mm. in length,
nearly concealing the annulations and forming a very slight
pencil ; and (h) loosely spreading hairs about 5 mm. long, rather
thickly sprinkled among the others ; annulation ill-defined,
about 35 to the centimeter at middle. Mamma?, i 3 — 3 = 6.
CROCIDURA g9
Colour.-Uppev parts varying from a slaty drab to dull
usse , the hairs with a slight znetallic gloss and with silve.y
reflections which in certain lights produc'e an evident effect of
fane speck ing Underparts and inner surface of limbs buf^
white, dulled to a varying degree by the slaty under colour th^
chin and throat often suffused with cream-buff S'e of
demarcation sharply defined, extending just below ear and eye
and alongside of muzzle to middle of pad' The two extremes of
colour probably represent ill-defined dichromatic phases Tlost
?U duirXSh 'ofr^'' "", '? ^^^"^ degree ^termediate
^eet dull whitish often irregularly clouded with drab Tail
sharply bicolor, whitish below, concolor with back above tbe
longer hairs silvery grey. ^^^ auo>e, uie
Skull -The skull is slightly larger than that of Sorex araneus
and noticeably more heavily built, particularly that portion
feCd To ^it ;r""'' i^ ^' ;^'^^^^ '^ both VoadenL Ls
S S- ? } i ^T'^'-'^^ ^"*^"^^ ^^P^^« less conspicuously
fiom behind forward, whether skull is viewed from above or
legion than in the European species of Sorex and Neomvs its
main sutures closing early in life. Brain-case ^ '
slightly longer than broad, its posterior out- ^ --__
Ime rounded but broken by the slightly (L^=:^^X
projecting points of the condyles, its antero- ^^^T""^'''^'^
external border straight, sharply angled in
front ; sagittal crest low but evident in adult
skulls, meeting the complete lambdoid crests
posteriorly. Depth of brain-case at middle
slightly but constantly less than half greatest
breadth (see tig. 20, page 1 00). Dorsal profile
with a slight concavity at front of brain-case
and slight convexity over middle of rostrum
-Nares squarely truncate posteriorly, the ^"'- "•
lateral wall abruptly angled near middle. "^""Nri^r""'-
Anteorbital foramen relatively smaller than in
Sorex aranens, and region between it and edge of alveolus dis-
tinctly wider. Lachrymal foramen over meta^tyle of J PlaJe
forming outer wall of anteorbital canal nearly thi-ee times as wMe
ZoU^T ^'""''T- ^"^"^'•^^" '^'^^-^^'^ ^^gi"" over posterior
molars broader and more prominent than in th'e European si.ecies
of t^orex, but rudimentary zygomatic process of maxillary reduced
floor ofT" """;• /^^-P*-yg-d fo-a as in SoreLranT;
floor of bra n-case between tympanic bones narrower and with
distinct median ridge.
snecieT'if"^!?'"^'^'^'! noticeably heavier than in the European
species of .STor^'.r and Neamy,, the difierence in general asnect
.TttT' A^''" ^^'^^"^^^' brown colouring matter'n jli
ot cusps. Anterior upper incisor with main cusp lon< and
slender, abruptly hooked downward; basal lobe low and
90
INSECTIVOEA
Fig. is.
Crocidura leucodon. Anterior
teeth in profile, x 5.
triangular, its height less than half that of main cusp and only
a little more than half that of first unicusjiid tooth. General
form of first incisor not unlike that of
Neomys fodiens, but with even more
contrast between height of the two
cu.sps. Anterior lower incisor simple,
rather robust, the shaft slightly taj^er-
ing, the point slightly bent upwai'd, the
cutting edge without lobes. Upper
unicuspids robust, strongly contrasted
in size. First about double the height
of second and third, the points of which
are nearly in line with that of basal
lobe of anterior incisor. Its crown is
.somewhat longer than broad, with main
axis slightly oblifiue to that of tooth-row. In lateral view it is
nearly triangular, with anterior-border about half as long as upper
and lower. Cutting edge essentially
as in Neomys, but better develo2)ed
and reaching posterior border of
tooth somewhat outside of middle.
Crushing surface well developed,
occujaying about one-third area of
crown. Second and third unicus-
pids approximately equal, in both
height and crown area, to basal
lobe of anterior incisor, their crowns
distinctly narrower as well as much
shorter than that of first
unicuspid, their cutting
ridge and crushing sur-
face less well diflferen-
tiated than in first.
Second smaller than
third, its crown about
as broad as long. Third
sepai'ated from large pre-
molar b}^ a slight space,
its crown longer than
broad, its cusp about
efiual in height to para-
cone of large premolar.
Lower unicusjiids essen-
tially alike in form, the
second differing from that
of Sorex and Nrontys in
the complete absence of the rudimentary second cusp. First low,
narrow and long ; second high, its crown about as broad as long.
Upper cheek-teeth with crowns narrower than in Sor-ex and
FlC ID
CrocitJuj'd leucodon.
CROCIDURA 91
Neomys, the emargination of posterior border deeper, particularly
in jm^, and hypocones more distinct. Large upper premolar
with antero-external cusp (paracone) well developed, its height
usually about equal to that of third unicuspid or somewhat
more, the distance from its point to well-defined angle in
cingulum over anterior root of tooth distinctly more than half
length of anterior border of main cusp ; posterior cutting blade
high, the angle formed between its edge and the moderately
projecting point of main cusp ill-defined and obtuse. Third
upper molar actually as well as relatively smaller than in Sorcx
araneus, but containing the same elements. Lower cheek-teeth
essentially as in Sorex aranetis except for the pecuharities of m^
characteristic of the genus.
Measurements. — Average and extremes of nine specimens
from Etupes, Doubs, France: head and body, 78-3 (70-87)-
tail, 35-3 (32-39); hind foot, 12-3 (12-13). Average and
extremes of nine specimens from Untervatz, Orisons, Switzerland :
head and body, 81 (77-85); tail, 31-6 (29-34); hind foot,
12-5 (11-8-13). Average and extremes of four specimens from
Lugano, Ticino, Switzerland: head and body, 80 (77-81); tail,
32 • 2 (31-36) ; hind foot, 1 2 • 5 (1 1 • 8-13). Average and extremes
of three specimens from Florence, Italy (in alcohol, bodies
contracted): head and body, 63 '3 (63-64); tail, 34-3 (32-36) ;
hind foot, 12-3 (11-6-13). " For cranial measurements see Table,
p. 92.
Specimens examined. — Sixty-six, from the following localities :—
Belgium: Esneux, Li6ge, 2; Waremme, Li6ge, 1 (U.S.N.M.).
France: Dinau, Brittany, 1; Paris, 1 (U.S.N.M.); Manouville,
Meurthe-et-Moselle, 1 ; Etupes, Doubs, 14 (B.M. and Mottaz).
Germany : Brunswick, 1 (U.S.N.M.) ; Nussberg, Brunswick, 1 ; Frank-
furt, Hessen-Nassau, 1 ; Taucha, Saxony, 1 (U.S.N.M.) ; Marxheim,
Bavaria, 4 ; Ummerstadt, Thiiringen, 1 ; no exact locality, 1. '
Austria-Hungary : Hatszeg, Hunyad, Transylvania, 1.
Switzerland : Geneva, 3 (Mottaz) ; Briinig, Bern, 1 (U.S N.M.) ; Mei-
rmgen, Bern, 4 (U.S.N.M.); Untervatz, Grisons, 15 (B.M. and U.S.N.M.);
Grisons, no exact localitv, 1 ; Lugano, Ticino, 4 (U.S.N M ) • Ziiber-
wangen, St. Gallen, 1. "
Italy: Porlezza, Como, 1 (Mottaz); Boccadassa, Genoa, 1 (Genoa) -
Pisa, 1 (Mottaz) ; Florence, 3 (U.S.N.M. and Mottaz) ; Rome, 1.
BeniarJcs. — This species is recognizable among the European
members of the genus by its sharply bicolor pattern of coloura-
tion and by the low, flattened brain-case. From Crocidura
riissnla, the only species with which it is likely to be confused,
it is further distinguished by the peculiarities of the large upper
premolar, a tooth whose higher cutting edge, better developed
paracone and more prominent cingulum indicate a degree of
efficiency superior to that of the corresponding tooth in the'related
92
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9 4 IXSECTIVORA
■2 ?, 1. Esneux, Liege, Belgium Lord Lilford (p). 'Jo. 1. 1. 1-2.
[H. Griinvold.)
i. Dinan, Brittany, Prance. G. Barrett-Hamilton 11. 1. 2. 158.
i. Manonville,Meurthe-et- Lord Lilford (p). 8. 9. 8. 1.
Moselle.
5 .5, 4 ?. Etupes, Doubs, 350 m. O. Thomas (p). 8. 8. 10. 33-41.
(C. Mottaz.)
2 9. Marxheim, PSavaria, Lord Lilford (p). 8.9.8.2-8.
Germany.
1. Frankfort, Hesseu- Dr. Dieffenbacli (p). 47.1.8.50.
Nassau.
6. Nussberg, Brunswick. G.Barrett-Hamilton 11. 1. 2. 90.
1 al. Germany. Tomes Collection. 7. 1. 1. 45.
9. Hatszeg, Transylvania, G. G. Danford (c). 3. 2. 2. 10.
1500 ft. Hungary.
3<:!, 4 9. Untervatz, Grisons. O. Thomas (p). 4.4.5.21-22.
Switzerland. 10. 8. 16. 9-13.
{E. H. Zollikofer.)
? sk. Grisons, 550 m. O. Thomas (p). 4. 4. 5. 20.
(E. H. Zollikofer.)
9. Lugano, Ticino. ' O. Thomas (p). 10. 8. 16. 15.
(E. II. ZoUil-ofcr.)
?. Ziiberwangen,St.Gallen. 0. Thomas (p). 10. 8. IG. 14.
(E. H. Zollikofer.)
6. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1 2. 103.
(P).
CROCIDURA MIMULA Miller.
(Synonymy under subspecies.)
Geographical distribution. — Central Europe from north-
western Spain through France, Switzerland, central Germany
and northern Hungary to Roumania and Jiulgaria, south into
Italy and Greece.
Diagnosis. — Size less than in Crocidura russida and C. leurodon
(hind foot, 10 to 12 mm. ; condylobasal length of skull, IG to
17 '6 mm.) ; skull and teeth resembling those of C. leucodon, the
brain-case nearly as much depressed, its height seldom more than
half width, the third unicuspid similarly low as compared with
small anterior cusp of large premolar ; colour not very different
from that of C. aranea, the upper and lower surfaces of body not
strongly- contrasted, and no line of demarcation along sides.
Colour. — Ti^pper pai'ts varying from a dull russet tinged with
sepia to a dark hair-brown with a tinge of drab, the liaii-s with
metallic gloss and silver}' reflections ; underparts varying from
dull ochraceous-buff" to greyish cream-buff", never sharply defined
from colour of back and sides. Feet bufly whitish, usually with
some dark clouding. Tail greyish or brownish, very obscurely
bicolor.
Skull and teeth. — Except for their noticeably smaller size the
skull and teeth closely resemble those of Crocidura leucodon,
though the brain-case is less constantly flattened, sometimes
assuming a form essentially like that in C. ru.^sula. Plate foT-ming
CROCIDURA
95
outer wall of aiiteorbital canal narrower than in C. Jeucodon, its
width scarcely twice that of lachrj-mal foramen. Large upper
premolar with high cutting blade and well developed paracone,
the form and size of which relatively to main cusp and to third
unicuspid are exactly as in C. Icucodon.
Measurements. — According to measurements made by various
collectors, the head and body varies from 55 to 72 mm., tail '
from 28 to 40 mm., and hind foot from 10 to 12 mm. The
condylobasal length of skull ranges between 16 '0 and 17 '6 mm.,
and length of upper tooth-row between 7 • 2 and 8 • mm.
Remarhs. — Ciocidnra mimiihi differs from the other continental
European members of the genus, in its small size, a character in
which it is approached by the small races of C. russulu, though
not sufficiently to cause any confusion. In addition to the
typical form two geographical races have been described, one from
south-western France, the other from northern Spain, the
status of neither of which is clearl}' understood.
Crocidura mimula mimula Miller.
1839. ?? [Crocidura aranea] var. minor de S61ys-Longchamps, Etudes de
Micromamm., p. 35 (Silesia).
1901. Crocidura mimula JMiller, Proc. Biol. Soc, Washington, xiv, p. 95,
June 27, 1901 (Ziiberwangeu, St. Gallen, Switzerland). Type in
U.S. National Museum.
1901. Crocidura antipse Matschie, Sitz.-Ber. Gesellsch. Naturforsch.
Freunde, Berlin, p. 228, November, 1901 (Siulnita and Barza,
Roumania).
1902. [Crocidura"} minuta Lydekker, Zool. Record, xxxviii (1901), Mamm.,
p. 27 (Accidental renaming of mimula).
1910. Crocidura mimula and C.antipai Trouessart, Fauue Mamm. d'Europe,
pp. 46, 48.
Tijpe localitji. — Ziiberwangen, St. Gallen, Switzerland.
Geographical distribution. — Range of the species from the
Rhone Valley eastward.
Diagnosis. — Size maximum for the species ; skull with brain-
case tending to be strictly of the flattened type : colour usually
dark.
Measurements. — External measurements of type : head and
body, 72; tail, 33; hind foot, 11. A male and female from
Untervatz, Grisons, Switzerland: head and body, 71 and 65;
tail, 35 and 35 ; hind foot, 10 and 11. A male and female from
Marxheim, Bavaria : head and body, 71 and 58 ; tail, 36 and 30 ;
hind foot, 10 and 10. Average and extremes of four specimens
from Haida, Arva, Bohemia : head and body, 59 • 7 (55-65) ;
tail, 30 (28-32) ; hind foot, 11-5 (11-1 2). A male from Gageni,
Roumania, and female from Bustenari, Roumania : head and
biuly, 64 and 66 ; tail, 31 and 34; hind foot, 11 and 11. Two
adult males from Agay, Var, France : head and body, 64 and 66 :
tail, 35 and 34 ; hind foot, 1 1 and 1 1 . Aflult female from
Viareggio, Italy : head and body. 70 : tail, 38 ; hind foot, 11 "6.
96
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98 INSECTIVORA
Adult male and female from Corfu, Greece : head and body, 74
and 75 ; tail, 44 and 44 ; hind foot, 11 "8 and 11 '4. For cranial
measurements see Table, p. 96.
Si^ecimens exaynined. — Thirty-seven, from the following localities : —
Phance : Abbeville, Somme, 1 ; Agay, Var, 2.
Germany : Marxheim, Bavaria, 2.
Austria-Hungary : Haida, Arva, Bohemia, 4 ; Hatszeg, Hunyad,
Transylvania, 1; Tatra Mts., Hungary, 1.
RouMANiA : Gageni, Prahova, 1 ; Bustenari, Prahova, 1.
Bulgaria : Sofia, 1 (Andersen) ; Varna, 1 (Andersen).
Switzerland: Ziiberwangen, St. Gallon, 3 (U.S.N.M. and Mottaz) ;
Untervatz, Grisons, 2 ; Faido, Ticino, 1 ; Santa Margherita, Ticino, 3
(Mottaz) ; Davesco, Ticino, 1 (U.S.N.M.) ; Lugano, Ticino, 2 (B.M. and
Mottaz) ; Locarno, Ticino, 1.
Italy: Porlezza, Como, 4 (Mottaz); Viareggio, Lucca, 2; Rome, 1.
Greece : Corfu, 2.
1. Abbeville, Somme, Baillon Collection. 56. b.
Prance.
2 (5. Agay, Var. G. S. Miller (c). 8. 8. 4. 162-168.
6. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.4.
Germany. ( Wolter-
stoif.)
6,9. Haida, Bohemia, Lord Lilford (p). 8. 9. 8. 5-6.
Austria.
?. Hatszeg, Hunyad, Tran- C.G. Danford (c&p). 3.2.2.12.
sylvania, Hungary.
1 al. Tatra Mcmtains. Dr. R. Collett (p). 91. 1. 21. 2.
?. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 11.
mania. {W. Dodson.)
6. Bustenari, Prahova, Lord Lilford (p). 4. 4. 6. 12.
840 m. (W. Dodson.)
9. Faido, Ticino, Switzer- 0. Thomas (c & p). 5. 8. 2. 18.
land.
6. Locarno, Ticino. O. Thomas (c & p). 5. 8. 2. 1.
6,9. Untervatz, Grisons. O. Thomas (p). 4.4.5.21-22.
{E. H. Zollikofcr.)
9. Lugano, Ticino. 0. Thomas (p). 4.4.5.57.
[E. H. Zollikofcr.)
9. Viareggio, Lucca, 5 m. 0. Thomas (c & p). 6. 8. 2. 21.
Italy.
6 Rome. (Coli.) G.Barrett-Hamilton 11.1.2.97.
(P).
6. Corfu, 50 m. Greece. J. I. S.Whitaker (p). 8.10.1.8.
(C. Mottaz.)
Crocidura mimula iculisma Mottaz.
1908. Crocidura nmmila icvMsma Mottaz, BuU. Soc. Zool. de Geneve, i,
p. 119, April 30, 1908. Type in Mottaz Collection.
1910. Crocidura mimula iculisma Trouessart, Paune Mamm. d'Europe, p. 47.
Type locality. — Lignieres-Sonneville, Charente, France.
Geographical clistribution. — Known from the type locality
onl}'.
Diagnosis. — Size as in C. tiiimnla mimula or slightly smaller
(hind foot, 10; condyloba.sal length of skull, 16); brain-case
deep, nearly as in C. russida.
CROCIDURA 99
Measuremenh. — Type (from Mottaz) : head and body, 60 ■ 5 ;
tail, 38*5; hind foot, 10 •2. For cranial measurements see
Table, p. 97.
Specimens examined. — Three, all from the type locality (Mottaz).
BemarJcs. — While this race appears to be distinct from true
miniuJa, the material seen is insufficient to form the basis of any
final opinion as to its status.
Cbocidura mimula cantabra Cabrera.
1908. Crocidura cantabra Cabrera, Bol. Real Soc. Espan. Hist. Nat., viii,
p. 239, May, 1908. Type in Madrid Museum.
1910. Crocidura cantabra Trouessart, Faune Mamm. d'Europe, p. 46.
Ti/jje localiiy. — Bastjue Provinces, Spain, exact locality not
known.
Geographical distribution. — Basque Provinces, Spain.
Diagnosis. — Colour paler and more grey than in the other
races ; size small.
Measurements. — Type (from Cabrera) : head and body, 55 ;
tail, 24 ; hind foot, 10 ; ear, 6*5 ; upper tooth-row, 7*2. (Cranial
dimensions not known.)
Bemarks. — I have not seen this animal, but from the original
description, as well as from information received from Mr. Cabrera,
it appears to be paler than the typical form, to which it bears
much the same relation as C. russula pulehra to true russula.
CROCIDURA RUSSULA Hermann.
(Synonymy under subspecies.)
Geographical (lislrihution. — Central and southern Europe,
from the Mediterranean coast to Holland and central Germany.
Not found in the British Islands.
Diagnosis — Size rather large (among the European forms) :
hind foot, 11 to 14 mm. ; condylobasal length of skull, 18 to
20 '4 mm.; upper tooth-row, 8-2 to 9 mm.; tail rather short,
its actual length 33 to 45 mm., its ratio to head and body
varying from 45 to 55 ; skull with brain-case not noticeably
depiessed, its height always at least half greatest width and
usually more ; large upper premolar with autero-external cusp
small, its height usualh' less than that of third unicuspid ;
colour of underparts not strongl}' contrasted with that of back,
the line of demarcation along side vaguely defined.
E.vtcrnal chararteis. — In external characters, aside from the
relatively longer tail, Crocidura russula agrees with C. Icucodon.
Depth of fur at middle of back about 5 mm. in summer, 8 mm.
in winter.
H 2
100 INSECTIVORA
Colour. — Upper parts varying from a dark hair-brown,
tinged with bister to a light drab with or without a shade of
wood-brown, the darker colour more frequent in winter pelage,
the light apparently pecuhar to summer. The pelage has the
usual metallic gloss, and the individual hairs show strong silvery
reflections in certain lights, particularly in the long full winter
coat. Underparts usually a dull buffy grey or ecru-drab, but
sometimes almost whitish, rarely tinged with a bright yellowish
brown,* never strongly constrasted with back, the line of
demarcation along sides always vague. Feet dull buffy grey or
light drab Tail obscurely bicolor, like back above, like belly
below.
Shtll and teeth. — The skull resembles that of Crocidura
leucodon, except that the brain-case is noticeably less flattened,
its depth at middle always exceeding one half greatest width, a
character readily appreciable to the eye when skulls of the two
animals are viewed from beliind. Teeth essentially as in the
related species, but large upper premolar with antero-external
Fig. 20. Fig. 21.
Posterior view of skull of Crocidura rus$itla. Anterior
Crocidura leucodon (upper teeth in profile. X ."i.
figure), and C. russu!a (lower
figure). X li.
cusp (paracone) low, its height often much less than that of
third unicuspid, the distance from its point to ill-detined angle
in cingulum over anterior root of tooth about half length of
anterior border of main cusp ; posterior cutting blade not so
high as in Crocidura leucodon, the angle formed between its
edge and conspicuously projecting point of main cusp well
defined and less obtuse than in the related species.
Measurements. — In the diflerent races the head and body
ranges from 04 to 95 mm., tail from 33 to 46 mm., hind foot
from 10 '8 to 14 mm., condylobasal length of skull from
18 to 20*4 mm. The unusual apparent variability in length
of head and body is probably in great part due to diflerences in
method of taking the measurement and to differences in the
condition of the specimens measured.
* Such specimens evidently formed the basis of Savi's Sorex tltoracicns
and Dehne's S. chrysothorax.
CEOCIDCRA 101
Crocidura russula russula Hermann.
1777. Sorex araneus Schreber, Siiugthiere, iii, p. 573 (not of Linnaeus, 1758).
1780. Sorex russulics Hermann in Zimmermann, Geogr. Gesch., ii, p. 382
(vicinity of Strassburg, Germany).
1780. ?? Sorex constrictns Hermann in Zimmerinann, Geogr. Gesch., ir,
p. 383 (vicinity of Strassburg, Germany). Based on young in nest.
1792. ■?•? Sorex unicolor Kerr, Anim. Kingd., p. 208 (Strassburg, Germany.
Based on Pennant's account of Sorex constrictus).
1793. Sorex musaraneus Cuvier, Tabl. Til6m. de I'Hist. Nat. des Anim.,
p. 109 (France).
1800. ?? Sorex Icncurus Shaw, Gen. Zool., I, pt. 2, p. 538 (Strassburg,
Germany. Based on Schreber, pi. clixc, S. constrictns Hermann).
1801. S[prex^ alraneiis'] cincreus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
ISOl. S[orcx\ a[raneus'] candidus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
1832. Sorex fimbriat us Wagler, Isis, p. 54 (Bavaria, Germany).
1832. Croc[;idu7-d] moschafa Wagler, Isis, p. 275 (Substitute for Sorex
fimbriatus).
1832. C\^rocidiira] major Wagler, Isis, p. 1218 (Bavaria, Germany).
1832. Crocidura rufa Wagler, Isis, p. 1218 (banks of the Rhine, Germany).
1832. Crocidura poliogaster Wagler, Isis, p. 1218 (banks of the Bhine,
Germany).
1832. Sorex thoracicus Savi, Nuovo Giorn. de' Letterati, Pisa, xxiv, p. 52
(near Pisa, Italy).
1839. ? Sorex inodorus de S61ys-Longchamps, Etudes de Micromamm.,
p. 34 (Savi cited as authority, but name apparently published here
for the first time as synonym of aranea (= russula)).
1839. ? [Crocidura ara7iea] var. ininor de Selys-Longchamps, Etudes de
Micromamm., p. 35 (Silesia).
1839. [Crocidura aranea] var. albivcntris de S61ys-Longchamps, Etudes de
Micromamm., p. 36. (No locality given.)
1839. ? Crocidura hydruntina Costa, Fauna del Ragno di Napoli, Mamm.,
p. 6 (Otrauto, Calabria, Italy).
1855. Sorex chnjsothorax Dehne, Allg. deutsche Xaturhist. Zeitung, Neue
Folge, I, p. 241 (Wilsdurf, near Dresden, Germany).
1857. Crocidura araneus Blasius, Saugethiere, Deutschlands, p. 144.
1895. Crocidura russula Thomas, The Zoologist, 3rd ser., xix, p. 63,
February, 1895.
1910. Crocidura russula Trouessart, Panne Mamm. d'Europe, p. 43.
Tyjje locality. — Vicinity of Strassburg, Germany.
Geographical distribution. — Central Europe, from Holland and
central Germany to the valley of the Garonne and the coast of
south-eastern France (Var) ; Italy ; Sardinia 1 * ; Guernsey and
Alderney, Channel Islands.
Diagruh<<is. — Size rather large (hind foot, 11 "7 to 14, condy-
lobasal lengtli of skull, 19 to 20-4), and colour usually dark,
seldom, if ever, becoming a light drab except in rather worn
summer pelage.
Measurements. — Average and extremes of four specimens from
Oosterbeek, Gueldei'land, Holland : head and body, 78 (76-81) ;
tail, 41 (37-45); hind foot, 12-8 (12 -2-13 -9). "Average and
* I have seen five Sardinian specimens in the Genoa Museum
resembling the typical form of Crocidura russula and differing widely
from the Corsicau C. cyrnensis (see pp. 111-112).
102
INSECTIVORA
extremes of nine specimens from Esneux, Liege, Belgium : head
and body, 77 (72-85); tail, 35-6 (33-38); hind foot, 12-4
(11' 7-13). Average of ten specimens fi-om Pas-de-Calais, France :
head and body, 74-5 (71-80) ; tail, 40-9 (38-46) ; hind foot, 13-1
(12 •5-13' 5). Average and extremes of six specimens from St.
Cergues, Vaud, Switzerland : head and body, 87 (83-95) : tail,
38-1 (35-41); hind foot, 12-8 (12-4-13-2). For cranial
measurements see Table, p. 104.
Spemyiens examined. — One hundred and two, from the following
localities : —
Holland : Oosterbeek, Guelderland, 4.
Belgium : Esneux, Liege, 9 ; Waremme, Liege, 8 (U.S.N.IM.) ; no exact
locality, 1.
France : Boulogue-sur-Mer, Pas-de-Calais, 8 ; Guines, Pas-de-Calais, 2 ;
Abbeville, Somme, 2 (B.M. and Mottaz) ; Cxuernsey, Channel Islands, 4;
Alderney, Channel Islands, 1 ; St. Briac, Brittany, 1 ; Lignieres,
Charente, 1 (Mottaz) ; Nancy, iIeurthe-et-]Moselle, 1 (Merriam) ; Etupes,
Doubs, 9 (Mottaz) ; Montauban, Haute-Savoie, 4 ; Valescure, Var, 2 ; Ax-
les-Thermes, Ariege, 1 ; Luchon, Haute-Garonne, 2.
Germany : Ummerstadt, Thiiringen, 2 ; Ingelheim, Rheinhessen, 2 ;
Strassburg, 1.
Switzerland : Geneva, 10 (LT.S.N.INI. and jNIottaz) ; St. Cergnies, Vaud, 9
(U.S.N.M. and Mottaz); Cxrosjoan, Vaud, 1 (^Mottaz) ; Chesieres, Vaud, 1
(Mottaz) ; Lucerne, 1 ; Vitznau, Lake of Lucerne, 3 ; Thurgau, Roggwil, 1 ;
St. Gallon, 3 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 (B.M. and U.S.N.M.) ;
Degersheim, St. Gallen, 5 (B.M. and U.S.N.M.) ; Engelberg, Unterwalden, 1.
Italy: Ceresole d'Alba, Turin, 2 (Turin).
3 <?, 1 ?. Oosterbeek, Guelderland, O. Thomas (c & p). 98. 2. 1. 9-12.
50 m. Holland.
6, ?. Oosterbeek, CTuelderland. Miller Collection. 7. 7. 7. 3851-
(0. Thomas.)
4 <5, 49,1. Esneux, Liege, Belgium.
Belgium. (H. Grunvohl.)
2(5, ?,1. Guernsey, Channel Is-
lands. (R.H. Bunting.)
Alderney.
Boulogne, Pas-de-Calais,
10 m. Prance.
Guines, Pas-de-Calais.
Abbeville, Somme.
St. Briac, Brittany.
Montauban, Haute-
Savoie.
IMontauban, Haute-
Savoie, 900 m.
(.4. Robert.)
Valescure, Var.
Ax-les-Thermes, Ariege.
Luchon, Haute-Garonne,
600 m. [A. Robert.)
Ingelheim, Rheinhessen,
Germany.
Strassburg, Alsace.
(C. Mottaz.)
Vitznau, Lake of
Lucerne, 500 m.
Switzerland.
Engelberg, 3300 ft.
Switzerland.
9.
5(5,3 9.
2 6.
1.
1.
2rf.
<5, 9.
9.
2 9.
3 9.
1 al.
Miller Collection.
Lord Lilford (p).
Tomes Collection.
0. Thomas (p).
W.Eagle Clarke (p).
O. Thomas (c & p).
0. Thomas (c & p).
Baillon Collection.
W. M. Daly(c & p).
A. Robert (c & p).
0. Thomas (p).
G. S. Miller (c).
V. Builles (c & r).
O. Thomas (r).
C. Hilgert (c).
0. Thomas (p).
0. Thomas (c & p).
7. 7. 7.
3852.
95. 1. 1.
7. 1. 1. 30.
8. 9. 2. 18-21.
-11.
9. 3. 28.
98. 1. 9.
1.
4-11.
94. 6. 6. 8-9.
56. A.
94. 10. 3. 1.
97. 1. 9. 2-3.
6. 4. 2. 2-3.
8.8. 4.164-165.
8. 3. 27. 1.
6. 4. 1. 19-20.
8. 11. 2. 9-10.
8. 8. 10. 43.
5. 8. 3. 9-11.
Dr. J. Anderson (p). 99. 7. 17. ] .
CROCIDDRA 103
Crocidura russula pulchra Cabrera.
1907. Crocidura russula ptilchra Cabrera, Ann. and Mag. Nat. Hist.,
7th ser., xx, p. 213, September 1, 1907. Type in Cabrera col-
lection.
1907. Crocidura russula pulchra Cabrera, Bol. Real Soc. Espafi. Hist.
Nat., Madrid, vii, p. 223, October, 1907. (For date see Cabrera,
Ann. and Mag. Nat. Hist., 8th ser., i, p. 189, February, 1908.)
1910. Crocidura russula pulchra Trouessart, Faune Mamm. d'Europe, p. 45.
Type locality. — Valencia, Spain.
Geographical distribution. — Central and southern Spain ; low-
lands of France south of the Gironde.
Diagnosis. — Smaller than Crocidura russula russula (hind foot,
10" 8 to 13, condylobasal length of skull, 18 to 19 '4), and jjaler
in colour, the back a light drab brown tinged with sepia or dull
russet.
Measurements. — External measurements of type, male (from
Cabrera): head and body, 71; tail, 41*5; hind foot, 12.
Average and extremes of seven specimens from 'Silos, Burgos,
Spain: head and body, 72-7 (67-78); tail, 36 (34-37); hind
foot, 11 "9 (10 •8-12 -4). Average and extremes of .seven
specimens from Ciranada, Spain : head and body, 69 * 6 (68—74) ;
tail, 38-2 (36-40); hind foot, 12-2 (12-12-8). Average and
extremes of ten specimens from Cadillac-sur-Garonne, Gironde,
France: head and body, 75-2 (70-81); tail, 37 (33-40); hind
foot, 11*6 (11-12). For cranial measurements see Table, p. 106.
Specimens examined. — Sixty-five, from the following localities ; —
Feance : Cadillac-sur-Garonne, Gironde, 19 (U.S.N.M.) ; Moutr6jeau,
Haute-Garonne, 3 (U.S.N.M.) ; St. Genies, near Nlmes, Gard, 2.*
Spain : Silos, Burgos, 18 ; Dehesa de Valencia, Valencia, 1 ; Alcoy,
Alicante, 8 ; Elche, Alicante, 2 ; Venta del Baul, Granada, 2 ; Granada, 8 ;
Barracas, Castellon, 1.
Portugal: Sierra de Gerez, 1 (in alcohol; perhaps referable to cintrai).
9. St. Genies, Gard, 102 m. 0. Thomas (p). 8. 8. 10. 42.
France. (C. Mottaz.)
4 6. Silos, Burgos, 980 m. G. S. MiUer (c). 8. 8. 4. 38-41.
Spain.
9. Dehesa de Valencia. O. Thomas (p). 8. 2. 9. 46.
(N. Gonzalez.)
2 ,i, 1 9. Alcoy, Alicante. 0. Thomas (p). 8. 2. 9. 43-45.
(N. Gonzalez.)
6,9. Venta del Baul, Granada. G. S. MiUer (c). 8.8.4.31-32.
4 rf, 1 9. Granada. G. S. Miller (c). 8. 8. 4. 33-37.
<5al. Sierra deGerez,Portugal. Dr. H. Gadow 87.3.28.1.
(c & p).
* Intermediate between pulchra and true russtila.
104
INSF.CTIVOKA
5
J
O
Teeth slightly worn.
,, not worn.
,, slightly worn.
,, not worn.
,, slightly worn.
,, not worn.
,, slightly worn.
„ moderately worn.
,, slightly worn.
,, much worn.
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Number.
98. 2. 1. 11
98. 2. 1. 12
95. 1. 1. 3
95. 1. 1. 4
95. 1. 1. 5
95. 1. 1. 6
95. 1. 1. 10
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86054
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86058
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86060
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T* — '
ID
fc ■
3 . S .
C. russula russ
Holland : Oosterbeek
Belgium : Esneux .
Warcmme
France : near Boulogi
Calais .
CROCIDURA
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INSECTIVOEA
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8. 8. 4. 39
8. 8. 4. 40
8.8.4.41
8.8.4.42
172114
8. 7. 7. 44
1
C. russula russula {continued).
Switzerland: Ziiberwangen, St. \
Gallen . . ./
Degersheim
Lugano, Ticino .
Italy : Ceresole d'Alba .
Pisa
C. russula pulchra.
Spain : Silos, Burgos
CUOCIDURA
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108 INSECTIVORA
Crocidura russula ciNTRyE Miller.
1907. Crocidura russula cintrx Miller, Ann. and Mag. Nat. Hist., 7th ser.,
XX, p. 390, November, 1907. Type in British Museum.
1910. Crocidura russula cintrx Trouessart, Faune Mamm. d'Europe, p. 45.
Type locality. — Cintra, near Lisbon, Portugal.
Geogrnpliical distrihution. — At present known onlj' from the
type locality.
Diagnosis. — Size as in C. russula pulchra (hind foot, 11 '4 to
12 "7, condylobasal length of skull, 18 to 19-2), but colour fully
as dark as in true russula, the back between the mars-brown and
russet of Ptidgway, the hairs with a peculiar, strong, coppery
lustre rarely indicated in the typical race.
Measurements. — External measurements of type : head and
body, 64; tail, 3.3 ; hind foot, 11*4. Average and extremes of
ten specimens from the type locality : head and body, 67 • 6
(64-72); tail, 37-7 (33-42); hind foot, 11 "9 (11 •4-12-7). For
cranial measurements see Table, p. 107.
Specimens examined. — Eleven, all from the type locality.
Remarks. — In its small size the Cintra shrew agrees with the
Spanish race, but the colour is conspicuously darker. Taken as a
whole the series, in winter pelage, is about as dark as in French
and Belgian russula ; but the noticeable coppery lustre is highly
characteristic of the Portuguese form.
6 6, 5 9. Cintra, Estremadura, 0. Thomas (c & p). 98. 2. 2. 10-20.
300-350 m. Portugal. (T7/|;eof subspecies 98. 2. 2. 11.)
CROCIDURA SICULA Miller.
1879. ? Crocidura sicula Giglioli, Wiegmann's Archiv fiir Naturgesch.,
1879, I, p. 96. Nomen nudum : " Crocidura sicula (Giglioli MSS.
sp. nov. ?) Castelbuouo, Sicilien."
1901. Crocidura sicula Miller, Proc. Biol. Soc, Washington, xiv, p. 41,
April 25, 1901 (Palermo, Sicily). Type in U.S. National Museum.
1910. Crocidura sicula Trouessart, Paune Mamm. d'Europe, p. 47.
Type locality. — Palermo, Sicily.
Geographical distribution. — Sicily.
Diagnosis. — Size and colour essentially as in Crocidura russula
pulchra (hind foot, 12 to 13 mm., condylobasal length of skull,
17 'G to 19 mm.); brain-case nearly as much flattened as in C.
leucodon, its depth usually a little less than half greatest width ;
crown ai'ea of molars slightly reduced.
Colour. — Upper parts a light bluish drab tinged with sepia ;
underparts faintly constrasted pale smoke-grey, with or without
a bufly cast. Feet dull whitish grey. Tail obscurely bicolor,
brownish above, whitish grey below.
Skull and teeth. — Except for its distinctly flattened brain-case
CROCIDCKA 109
the skull resembles that of the small races of Crocidura russula.
Teeth as in C. russula, but upper molars with crown area
somewhat reduced, a character readily appreciable on comparison,
and large upper premolar with autero-external cusp slightly
enlarged, its form and relative size approaching the conditions
found in C. leucodon.
Measurements. — External measurements of type (male) and
a second specimen from the type locality : head and body, 68
and 75; tail, 32 and 35; hind foot, 12 and 12. Average and
extremes of six specimens from San Giuglielmo, Castelbuono,
Sicily : head and body, 76-3 (72-80) ; tail, 35-3 (32-41) ; hind
foot, 12'8 (12-13). For cranial measurements see Table, p. 113.
Specimens examined. — Fourteen, all from Sicily. Exact localities :
Palermo, 5 (B.M. and U.S.N.M.) ; Marsala, 2; San Giuglielmo, Castel-
buono, 6 ; Ficuzza, 1.
2 ?. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 2-2*.
1. Marsala. {A. Robert.) O. Thomas (p). 6. 8. 4. 26.
3 cJ, 1 ?. San Giuglielmo, Castel- 0. Thomas (p). 8. 9. 1. 6-9.
buono. (^4. Bobcrt.)
CROCIDURA CANE.ffi: Miller.
1909. Crocidura canese Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,
p. 418, May, 1909. Type in British Museum.
1910. Crocidura canex Trouessart, Fauna Mamm. d'Europe, p. 48.
Type locality. — Crete.
Geographical distribution. — Island of Crete.
Diagnosis. — Size and general appearance as in the smaller
forms of Crocidura russula, and skull with similarly deep cranium ;
but second upper premolar as large as third, and entire anterior
portion of upj^er tooth-row unusually long relatively to cheek-
teeth.
Colour. — The colour does not difter appreciably from that of
dark individuals of C. 7-ttssula.
Skull. — The skull is essentially .similar to that of the smaller
forms of Crocidura russula. Brain-case slightly more than half as
high as wide. Anterior portion of palate between unicuspids
and anterior incisors more nearly parallel-sided, and more
elongate than in any of the related species.
Tectli. — The teeth differ from those of all the other known
European members of the genus in the approximately equal size
of the two small upper unicuspids and in the longer, relatively
narrower crown of the first imicuspid. In the related species the
first unicuspid is so wide posteriorly that it makes an abrupt and
noticeable break in the outline of outer side of tooth-i'ow. In
C. caneie this tooth, though larger than usual, is not sufficiently
wide to project beyond the general line of the outer margins of
the unicuspid teeth. Second unicuspid fully as large as third,
which is of normal size. The unicuspid row is thus distinctly
110 INSECTIVOKA
increased in length, so that the distance from front of large
premolar to front of incisor equals that from front of large
premolar to mesostyle of second molar, while in the related
species^it equals that from front of large premolar to metastyle
of first molar. Large premolars and molars, particularly those
of mandible, more robust than usual though not peculiar in form.
Measurements. — External measurements of type (male) : head
and body, 65 ; tail, 42 ; hind foot, 11 '8. External measurements
of adult male from Canea : head and body, 71; tail, 47;
hind foot, 12" 6 ; ear, 9*5. For cranial measurements see Table,
p. 113.
Specimens examined. — Two, both from Crete.
6 al. Crete. Purchased (LinnEea, 84. 3. 14. 2.
Frankfort).
[Type of species.)
6. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 1.
(C. H. B. Grant.)
CROCIDURA CAUDATA Miller.
1901. Civcidura caxidata Miller, Proc. Biol. See. Washington, xiv, p. 42,
April 25, 1901. Type in U.S. National Museum.
1910. Crocidura caudata Trouessart, Faune Mamm. d'Europe, p. 49.
Ti/pe locality. — Palermo, Sicily.
Geographical flistrihution. — Sicily.
Diagnosis. — Size about as in large specimens of C. 7u.9si(ln
(hind foot, 14 mm.). Tail very long, its ratio todaead and bodj-
about 80, and so thickened that its diameter at middle is 3 mm.
(in other European sjjecies the diameter of tail scarely if at all
exceeds 2 mm.).
External characters. — Except for the unusual length of the
tail Crocidura caudata does not difter in external characters from
C. russula. The tail is so long that when laid forward over back
it extends to between ears. It is distinctly 4-sided, broader
below than above, its greatest diameter at middle 3 mm.*
Colour. — After six months' immersion in alcohol the colour of
the type specimen was essentially as in Crocidura sictda. After
eight years more in the same fluid the back appears to have
assumed a somewhat more brownish cast.
Sliull and teeth. — The only known skull is so injured that the
details of its form cannot be seen. -The rostral portion does not
difter appreciably from that of C. russtda. Teeth essentially as
in C. russula, but lirst upper unicuspid larger, third unicuspid
more crowded against large premolar, and cutting edge of large
pi-emolar higher, its antero-external cusp, however, of the same
form as in C. russula.
* In the type the tail is flattened laterally for about 13 mm. from tip,
evidently as the result of an accident.
CROCIDDRA 111
Measurements. — External measurements of type : head and
body, 63; tail, 52; hind foot, 14. For cranial measurements
see Table, y>. 113.
Specimen examined. — -The type.
Hemarhs. — The tail is actually as well as relatively longer in
this species than in any other European member of the genus.
CROCIDURA CYRNENSIS Miller.
1907. Crocidura cyrnensis Miller, Ann. and Mag. Nat. Hist., 7th ser., sx,
p. 390, November, 1907. Type m British Museum.
1910. Crocidura cyrnensis Trouessart, Fauna Mamm. d'Europe, p. 49.
Type locality. — Bastia, Corsica.
Geographical distribution. — Corsica.
Diagnosis. — Smaller than Crocidura caudata (hind foot, 12 to
12- 4 mm.) but with tail relatively almost as long, its ratio to
head and body about 70.
External characters. — Similar to C. caudata except for the
smaller size ; tail apparently less thickened than in the Sicilian
animal, its diameter at middle only about 2 mm.
Colour. — Back and sides drab washed with a brown inter-
mediate between wood-brown and raw-umber, this especially
noticeable on posterior half of back, but scarcely extending to
sides, which are a nearly clear drab ; underparts a light buffy
drab-grey, inconspicuously contrasted with sides ; tail dull dark
drab, essentially unicolor ; feet (both fore and hind) like tail on
outer half, rather sharply contrasted pale buffy grey on inner
half.
Skull and teeth. — AVhile its general size and form are essentially
as in Crocidura russula, the skull of the Corsican shrew is
distinguishable by its broader, more deepened rostrum. In the
type the mandible is peculiar in the unusual depth of ramus,
though in a second specimen this character is less marked.
Teeth essentially as in C. russula.
Measurements. — External measurements of type (adult male) :
head and body, 67 ; tail, 48 ; hind foot, 12-4. Very old female
from the type locality : head and body, 62 ; tail, 46 ; hind foot,
12*4 ; ear, 8 '2. External measurements of well made skin from
La Foce de Vizzavona : head and body, 72 ; tail, 51 ; hind foot,
12. For cranial measurements see Table, p. 113.
Specimens examined. — Three, all from Corsica.
Bemarlcs. — Crocidura cyrnensis is nearly related to C. caudata,
though readily distinguishable by its smaller size and less
thickened tail. So far as known it is the only shrew inhabiting
Corsica. Whether an animal of this type occurs in Sardinia
is a matter of doubt. Five Sardinian specimens (three from
112 INSECTIVORA
Ovile Seardu and two from Zinnigas) in the Genoa museum
appear to be strictly of the russula type, though without com-
parison of the skulls it is impossible to say whether they are most
nearly related to true russula or to sicula. Their average and
extreme measurements (from spirit specimens) are as follows :
head and body, 65-2 (60-69); tail, 36-8 (33-6-39): hind
foot, 12 (11 -8-12 -2).
1 La Foce de Vizzavona, Col. J. W. Yerbury (c & p). 93. 9. 15. 3.
Corsica.
6 al. Bastia, Corsica. Mrs. Southwell (c & p). 6. 3. 14. 1.
{Type of species.)
9 al. Bastia, Corsica. Mrs. Southwell (c & p). 9. 6. 14. 1.
CROCIDURA BALEARICA Miller.
1901. Crocidura russula Thomas, Proc. Zool. Soc. London, p. 39.
1907. Crocidura halearica Miller, Ann. and Mag. Nat. Hist., 7th ser., xs,
p. 391, November, 1907. Type in British Museum.
1910. Crocidura halearica Trouessart, Faune Mamm. d'Europe, p. 49.
Type locality. — San Cristobal, Minorca, Balearic Islands.
Geographical distribution. — Balearic Islands, Spain.
Diagnosis. — In general similar to Crocidura cyrnensis, but
tooth-row distinctly shorter than in the Corsican form.
Colour. — The three skins are more noticeably brownish above
than in the one skin of ci/rnensis at present known, though the
actual elements of the colour are the same. Feet with the same
colour pattern.
Shdl and teeth. — In cranial and dental characters the Balearic
.shrew agrees with Crocidura cyrnensis, except that the mandible
is less robust (essentially as in the small races of C. russula) and
the tooth-row is distinctly shorter. Brain-case somewhat more
flattened than in the small races of C. russula.
Measurements. — External measurements of type (female) :
head and body, 62 ; tail, 45 ; hind foot, 12-5. External measure-
ments of two other specimens from the type locality (male and
female) : head and body, 71 and 72 ; tail, — and 45 ; hind foot,
12 "5 and 12. For cranial measurements see Table, p. 113.
Sjxcimcns examined. — Three, all from the type locality.
c?, 2 9 San Cristobal, Minorca; 0. Thomas and R.I. Pocock 0. 7. 1. 40-42.
Balearic Islands. (c & p).
(0. 7. 1. 42. Tij2)e of species.)
CROCIDURA
113
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114 insectivora
Family ERINACEID^.
1821. Erinaceidx Gray, London ^Med. Eepos., xv, p. 300, April 1, 1821.
Geogrcqjhical disiribntion. — Tropical and temperate Africa,
Europe and continental Asia ; in Europe west to Ireland, north
to central Sweden and south-eastern Norway.
Characters. — Skull deep and heavy, not specially tapering
anteriorly, most of the sutures persistent ; zygomatic arch com-
plete and heavy ; tioor of brain-case completely ossified ; tympanic
bone annular, not attached to skull ; auditory process of basi-
sphenoid large, sometimes forming an evident half-bulla ; glenoid
surface directed downward (normal) ; a large external pterj'goid
plate ; teeth anterior to molars neither well differentiated by
form into incisors, canines and premolars nor strictly " unicuspid,"
the anterior upper incisor higher than the others but not specially
modified in form ; anterior lower incisor short, oblique ; crowns
of upper molars rather high, sub-quadrate in outline (except the
reduced third), the cusjas sub-equal, subterete, near margin of
crown, the styles and commissures reduced or absent, never
forming an important functional part of the tooth ; form short
and heavy ; eyes and ears well developed ; snout pointed, some-
what produced ; back normally covered with short, stifl" spines.
Remarlcs. — At present this family is usually regarded as
containing the single genus Erinaceus. It is very probable,
however, that several genera are represented among the members
of the group.
Genus ERINACEUS Linmmis.
1758. Erinaceus Linnreus, Syst. Nat., i, lOth ed., p. 52 {E. curopxus).
1857. Erinaceus Blasius, Saugethiere Deutschland.s, p. 152.
1868. Herinaceus Mina-Palumbo, Ann. Agric. Sicil., 2nd ser., sii, p. 37.
Ti/pe species. — Erinaceus europseus Linnaeus.
Geographical distribution. — Essentially coincident with that
of the family.
Characters. — Skull rather short and broad, the zygomatic
breadth distinctly more than half greatest length ; posterior
palatal region conspicuously fenestrate ; auditory process of basi-
sphenoid well developed, concave, sometimes forming a half-bulla ;
external pterygoid plate rather larger than internal pterygoid
plate, formed aliout equally of ectopterygoid and a broad horizontal
outgrowth from palatine; dental formula: i ^'j^, <-'}—, P«lr^,
m -lo = 36 ; canines not diflPerentiated by form from the contiguous
teeth ; third upper molar consisting of a large j'l'otocone and
minute j)aracone, all trace of crushing sui'face absent ; body
short and heavy, the back covered with stifT, sliarply pointed
bristles of uniform length : tail shorter than hind foot.
ERINACEUS 115
BemarJcs. — About twenty-tive species are currently referred
to this genus, four of them occurring in Europe.
KEY TO THE EUROPEAN F0R:MS OF ERINACEUS.
Third upper iucisor definitely 2-rooted ; elevated
portion of posterior lower premolar 2-cusped ; a
bare area among spines at middle of forehead
^ (Spain, Balearic Islands and southern France)... E. algirics, p. 130.
Underparts clouded with brown, at least in inter-
ramial and intercrural regions (Spain and
southern France) E. a. alg!n(s, p. 131.
Underparts entirely whitish (Balearic Islands).... A', a. vagaiis, p. 1.33.
Third upper incisor never definitely 2-rooted ;
elevated portion of posterior lower premolar
3-cusped ; no bare area among spines at middle
of forehead.
Greatest upper length of maxillary greater than
or at least equal to depth of rostrum at middle ;
anterior upper premolar sub-equal to canine,
its posterior border with small though evident
cusp (Eastern).
Hind foot 40 to 43 mm. ; condjiobasal length
of skull in adult male about 58 mm. (Eastern
Germany through Bohemia and Roumania
to Greece) E. roumanicus, p. 127.
Hind foot 35 to 38 mm. ; condylobasal length
of skuU in adult male about 55 mm. (Crete) E. nesiotes, p. 129.
Greatest upper length of maxillary less than
depth of rostrum at middle ; anterior upper
premolar decidedly smaller than canine, its
posterior border with cusp obsolete or absent
(Western) E. europseus, p. 115.
Size larger, the skull in old males exceeding
59 mm. in condylobasal length.
Average colour darker, the face never clear,
pale, huffy grey, but usually with notice-
able blackish markings (Central-western
Europe) E. c. europseus. p. 120.
Average colour lighter, the face usually clear,
pale, buffy grey without noticeable black-
ish markings (Iberian Peninsula) E. c. hupanlcus, p. 122.
Size smaller, the skull in old males not ex-
ceeding 59 mm. in condylobasal length.
Colour of head and shoulders lighter than
that of contiguous spiny area (Italy) E. e. italicus, p. 123.
Colour of head and shoulders darker than
that of contiguous spiny area (Sicily) E. e. coiisolei, p. 12G.
ERINACEUS EUROPSEUS Linnteus.
(Synonymy under subspecies.)
GeoijrnplikaJ distribution. — Europe from the Mediterranean
coast to Scotland and southern Scandinavia ; west to Ireland ;
eastern limits of range not known.
DiagnoniK. — Size large (head and body about 225 to 27-5 ; hind
foot 40 mm. or more ; condylobasal length of skull more than
116 INSECTIVORA
55 mm.) ; spines coarse and heavy, extending in an unbroken
line across forehead and decidedly overtopping ears ; skull with
heavy, deep rostrum, the distance from posterior extremity of
premaxillary to jjosterior extremity of maxillary less than rostral
depth at middle ; auditory process of basisphenoid short, wide-
funnel shaped, not forming a half-bulla ; third upper incisor with
one root, this sometimes partly divided longitudinally ; elevated
portion of posterior lower premolar with three cusps (tig. 25 a).
External characters. — General form short, thick and clumsy,
the legs short, the feet large, plantigrade. Legs, tail, underparts
and head, except ci'own, clothed with coarse fur, the finer more
woolly hairs of which are about 1 5 mm. long, the longer, straight
hairs about 40 mm. in length. The rest of the body is covered
b}^ a densely-set mass of sharply pointed bristles about 25 mm. in
length and 1 mm. in diameter.* The skin on wliich these bristles
are set is loosely attached to body, and jjrovided with a special
system of muscles by which the edges of the spiny area can be
drawn together ventrally over the animal's retracted head and legs,
forming a comjjlete protection for the entire body. Feet robust
with short digits and well developed claws, those on hind feet
longest. Fore foot broad and rounded with very short, thick
fingers, the third and fourth sub-equal and longest, second slightly
shorter, fifth extending to base of fourth, first well developed
but not reaching base of second ; balls of all five large and pad-
like ; palm entirely naked ; three large, semi-coniluent pads
at bases of median digits, a small tubercle (about 2 mm. in
diameter) at base of thumb, and two large pads at back of wrist,
the outer the larger ; skin between pads wrinkled ; hind foot
much like fore foot but longer ; second, third and fourth digits
sub-equal and longest, but their claws noticeably graduated from
second to fourth, fifth digit extending to base of fourth, first not
reaching base of second ; sole naked, the tubercles as on palm
but more crowded, the two posterior sub-equal. Tail short and
thick, its length much less than that of hiud foot. Ear simple,
rounded, shorter than bristles on crown, the meatus without
valves. Muzzle moderately produced, somewhat pointed, the
muzzle-pad well developed, naked, its surface marked with minute
furrows, its lateral edgesfinely scalloped, its lower border continued
as a pair of parallel ridges extending inward to palate. Eye
well develojaed but rather small. Mammie : p 1-1, ff. 2-2,
i 2-2 = 10.
Colour. — Furred portions of body varying from dull brown to
dirty whitish, the under fur usually a dusky hair-brown, the
longer hairs lighter and more buffy. Belly often irregularly
blotched and variegated with whitish and darker or lighter
brown. Feet usually darker than sides. Cheeks and eye-ring
often darker than rest of head. Sjjines buflty at base, then with
* Rarely the bristles are replaced by coarse hair like that on under-
parts. See Natural Science, xiii, p. 156, pi. ii, September, 1898.
EEINACEUS
117
I
a slaty area of variable width, followed by a narrow but sharply
defined buffy annulation and an obscurely darker tip. The
general effect is a coarse grizzle, tlie exact tone of which varies
considerably in the different geographical races as well as in
individuals of the same race.
SIcuU. — General form of skull rather short, heavy and deep,
tlie zygomatic breadth about | upper length, the brain-case not
Fig. 22.
Erinafens eitropaeus.
conspicuously wider than interorbital region, the rostrum short
and deep (ilistance from anteorbital foramen to front of pre-
maxillary less than depth through anterior root of zygoma).
Occiput and interorbital region marked by noticeable" ridges.
118 INSECTIVOEA
Ventral profile straight, the dorsal profile essentially parallel to
it from lambda nearly to front of interorbital region, then sloping
forward at a slight angle (about 15°) ; occiput squarely or some-
what obliquely truncate. General outline of occijiital region as
viewed from behind truncate-triangular, slightly more than half as
high as wide, the base of the triangle formed by line joining tips
of widely projecting mastoid processes, the apex by the narrowly
rounded or bluntly j^ointed lambdal region. Paroccipital
processes nearly as large as mastoid processes and resembling
them in form, though more slender and directed more backward.
Basisphenoid with deep median pit between bases of half-funnel
formed auditory processes, the pit continuous anteriorly with
mesopterygoid fossa. Tympanic ring open jaostero-externally, its
greatest breadth (antero-internal) about 3 mm. Inner and outer
2)terygoid plates broadly triangular, approximately alike in size
and form, each containing more of the pterygoid than palatine
element. Hamular short, strongly curved. Mesopterygoid sjiace
slightly longer than broad. Palate terminating posteriorly in a
high transverse ridge and strongly projecting median spine, the
ridge nearly straight, its median portion well developed. In
front of I'idge the palatine bones are conspicuously and irregularly
fenestrate. Lambdoid crest high. Sagittal crest low but
evident, extending forward to back of interorbital region. Here
it divides into two low, diverging ridges which pass forward
toward lachrymal region. In some specimens they can be traced
as far as the high, well defined ridge which occupies edge of orbit
for a distance of about 7 mm. above lachrymal foramen. Ante-
orbital foramen small, separated from lachrymal foramen by a
space much greater than its own diameter, its anteiior border
over anterior root of large pi'emolar. Upper portion of maxillary
rather short, its length behind posterior point of premaxillary
less than depth of rostrum at middle. Posterior termination of
premaxillary variable in form : nearh' squai'e, broadly or narrowly
cuneate, rounded, or rounded with supplemental inner spicule.
ManiliV)le short and heavy, the greatest depth of I'amus about one-
third length of alveolar line. Coronoid process high, narrow,
sharply hooked backward at tip. Angular process about as wide
as coronoid process, but not so long, its apex slightly bent
inward.
Teeth. — General aspect of teeth as compared with that in
other European members of the order, short, heaA^y and blunt,
distinctly omnivorous rather than strictly insectivorous in type.
Anterior upper incisor about twice as high as the succeeding
small teeth, its shaft subterete, flattened posteriorly, directed
slightly forward and inward, the teeth separated at base by
space about ecjual to height of shaft, at tip by about half this
distance. The four succeeding teeth (two incisors, canine and
anterior premolar) are essentially alike in form, the crown
slightly longer than wide, its height slightly greater than length,
ERINACEUS
119
the blunt point of conical cusp somewhat in front of middle of
crown. Of these four unicuspid teeth the first is smallest, the
second and thii'd sul>equal and larger, the fourth intermediate.
On posterior side of crown of each unicuspid there is a faintly
developed ridge extending to apex of cusp, this ridge tending to
rise posteriorly, especially in fourth unicuspid, to form a verj'
rudimentary secondary cusplet. First and second incisors invari-
ably single-rooted. Third incisorsingle-rooted, but root occasionally
showing trace of longitudinal furrow on outer side. Canine
usually single-rooted, the root with or without longitudinal
furrow ; but in a small series of specimens every stage may be
observed from this condition to a completely two-rooted tooth,
each root with a distinct alveolus. First premolar single-rooted,
the root of ten showing traces of longitudinal division, and perhaps
rarely double. Anterior lower incisor essen-
tially like the corresponding upj^er tooth
but not so high. Its shaft is directed
obliquely forward in line with symphysis,
the teeth of opj^osite sides parallel, separated
throughout by a narrow space.
Three succeeding teeth unicuspid,
the crowns similar in outline to
those of upiaer unicusjiids, but an-
terior cusps obsolete and posterior
cusplets relatively better developed.
Second upper premolar scarcel)"
broader than the unicuspids, but
three-rooted and with a distinct
protocone, metacone and postero-
external commissure. Large upper
premolar with well developed pro-
tocone, hypocone, metacone and pos-
tero-external commissure, the cusps
much as in the molars except that
hypocone is relatively smaller and
metacone and its commissure larger
and more trenchant. Large lower premolar with a high anterior
three-cusped portion similar to first triangle of lower molars,
except that the metaconid is reduced to a slight thickening at
inner base of commissure of protoconid ; second triangle repre-
sented by a mere narrow ledge or thickened cingulum. First
upper molar sub-quadrate in outline, the crown slightly wider
posteriorly than anteriorly. Protocone with somewhat broader
base than the other cusps and with low ill-defined anterior
and posterior commissures. Paracone, metacone and hypocone
sub-equal, the metacone slightly larger than the others. All
three are subterete with faintl}- indicated commissures, that
extending outward and backward from metacone to rudimen-
tary metastyle the most distinct. Parastyle and mesostyle
Tig. 23.
Eriiiacetts europieus. Teeth.
120 INSECTIVORA
absent.* A small but evident metaconule. Second molar like
first but smaller, its crown area about equal to that of large
premolar, its greatest diameter anterior instead of posterior.
Cusps essentially as in first molar, except that paracone is larger
than metacone and hypocone, and metaconule is barely indicated.
Third molar reduced to a protocone nearly as large as in the
other teeth, and a rudimentary paracone, the two connected by
a cutting edge sloping obliquely outward, forward and upward.
The tooth is single-rooted. Lower molars with the usual cusps
and commissures, the cusps more terete and commissures less
trenchant than in other European insectivores. Third molar
consisting of the anterior triangle only, this somewhat smaller
than in the other two teeth.
Erinaceus EUROPyEUS EUROP^us Linujeus.
1758. [Erinaceus'} europxiis Linnteus, Systema Naturae, i, lOfch ed., p. 52
(Sweden).
1779. [Hystrix] erinaceus Blumenbach, Handbuch d. Naturgesch., p. 72
(Germany).
1803. Erinaceus suillus Geoffroy, Catal. ^lammif. du Mus. Nat. d'Hist.
Nat., p. 67 (France).
1303. Erinaceus caninus Geofiroy, Catal. ISIammif. du 'Sins. Nat. d'Hist.
Nat., p. 68 (France).
1857. Erinaceus europxus Blasius, Saugethiere Deutscblands, p. 153 (part).
1858. E\_rinaccus'] canicejjs Hamilton Smitb, Jard. Nat. Libr., 2nd ed., xv
(Mammalia I), p. 148 (near Brussels, Belgium).
1897. [Erinaceus] echinus Scbulze, Abb. u. Vortr. Gesammtb. Naturw. iv,
No. 10, p. 19 (Substitute for europunis).
1900. Erinaceus europibus Barrett-Hamilton, Ann. and ilag. Nat. Hist.,
7tb ser., v, p. 362, April, 1900.
1900. Erinaceus europxus occidentalis Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7tb ser., v, p. 362, April, 1900 (Haddingtonshire,
Scotland). Type in British Museum.
1910. Erinaceus europxus and E. europx-us occicloitaUs Trouessart, Faune
Mamm. d'Europe, p. 38.
Tjiije locality. — Upsala, Sweden.
Oeoijraphical distribution. — Western central Europe from
Scotland, southern Norway, and central Sweden to the Pyrenees
and Alps ; west to Ireland : eastern limits of range not known.
Diagnosis. — Size largest of the European hedgehogs (condylo-
basal length of fully adult skulls usually 61 to 63 aim., seldom
less than 60 mm.) ; colour dark, the sides and underparts seldom
if ever a light buffy grey : face with blackish area extending
from eye to muzzle.
Measurements. — Adult male and female from Upsala, Sweden :
head and body, 265 and 263 : tail, 34 and 37 : hind foot, 44
and 43. Adult male from Innerwick, Haddingtonshire, Scotland :
head and body, 218 ; tail, 17 : hind foot, 42. Adult male and
* The cingulum usually forms a small projection resembling a rudi-
mentary parastyle.
EKINACEUS 121
female from Oundle, Northampton, England : head and body,
■2id and 257 ; tail, 24 and 31 ; hind foot, 40 and 40. Adult
male and female from Maredsous, Namur, Belgium : head and
body, 270 and 251 ; tail, 39 and 33 ; hind foot, 44 and 44.
Adult male from Bouconne, Gers, France : head and body, 220 ;
tail, 20 ; hind foot, 41. Adult male atid female from St. Gallen,
Switzerland : head and body, 279 and 297 ; tail, 41 and 43 ; hind
foot, 44 and 47. For cranial measurements see Table, p. 124.
Specimens examined. — Seventy-three, from the following localities : —
Scotland: Dunphail, Elgin, 2; Gordonstown, Elgin, 2; Innerwick,
Haddingtonshire, 1 ; Glendoc, Inverness, 1 (Wilson) ; Lanarkshire, 1.
England: Kelnsea, Spurn, Yorkshire,!; Leeds, Yorkshire,! (U.S.N.M.);
Somersetshire, 4 ; Shrewsbury, Shropshire, 3 ; Oundle, Northampton, 2 ;
Graftonbury, Herefordshire, ! ; Saifron Walden, Essex, 2 ; Banstead,
Surrey, 1 ; Elstead, near Godalmiug, Surrey, ! : Ockley, Surrey, ! ;
V/andsworth Common, Surrey, ! ; Hampshire, 1 (U.S.N.M.).
Wales : Cardiff, 2.
Ireland: Castle Hamilton, !; Nenagh, Tipperary, 2; Ennis, Co.
Clare, 2 ; Glenmore, Co. Donegal, ! (U.S.N.M.) ; KUmanock, Wexford, 2.
Norway : Asker, near Christiania, !.
Sweden : Upsala, 5 (U.S.N.M.) ; Upland, !.
Denmark : Copenhagen, 5 (Andersen).
Holland : No exact locality, !.
Belgium : Maredsous, Namur, 2.
France : Foret de Bouconne, Gers, ! ; Cranves-Sales, Haute-Savoie, !.
CtERmany : Brunswick, 1 (U.S.N.M.) ; Heidelberg, 1 (U.S.N.M.) ; Strass,
near Burgheim, Bavaria, 3; Ingelheim, Rheinliessen, 1.
Switzerland : GJeneva, 2 (Mottaz) ; St. Gallen, 7 (B.M. and U.S.N.M.) ;
Uzwil, St. Gallen, 1 (U.S.N.M.) ; Heresau, St. Gallen, ! (U.S.N.M.) ; Wolf-
halden, Appenzell, ! (U.S.N.M.) ; Thurgau, 2.
Bemarhs. — The typical race of Erinaceus europseus is dis-
tinguishable from the forms occurring in the Mediterranean
region by its combination of large size with dark colour. It is
more readily confused with the dark E. roumanicus, whose range
adjoins it on the east, and from which it cannot be distinguished
with certainty except by comparison of the skull and teeth. The
cranial character supposed to distinguish British specimens from
the Continental form appears to be too inconstant to warrant the
recognition of an insular race.*
6, 9. Crordonstown, Elginshire. W. E. Ogilvie-Grant II. !. 3. 379-
Scotland. (c & p). 380.
6. Innerwick, Haddington- W. Eagle Clarke (c & 0. 3. 13. !.
shire. p).
{Type of E. e. occidcntalis B.-Ham.)
9. Stockbriggs, Lanarkshire. PJ. R. Alston (c & p). 79. 9. 25. 76.
6, 9. Oundle, Northampton, Hon. N. '0. Roths- !1. 1. 3. 381-
England. child (c & p). 382.
S. Graftonbury, Hereford- W. E. de Winton (c !1. !. 3. 883.
shire. & p).
* For discussion of this character see Lonuberg, Ann. and Mag. Nat.
Hist., 7th sor., v, pp. 542-544, June, 1900, and Barrett-Hamilton, I.e.,
pp. 245-246, August, 1900. The question cannot yet be regarded as
decided, since no adequate series of skulls has yet been brought together.
122
INSECTIVORA
2 6. Saffron Walden, Essex.
{Wright.)
6. Banstead, Surrey.
6. Godalming, Surrey.
6. Ockley, Surrey.
3juv.al. Shrewsbury, Shropshire.
6, 9. Somerset.
i, 9 al. Somerset.
9. Cardiff, Glamorganshire,
Wales.
6. Nenagh, Tipperary, Ire-
land. (W. SmitJuvick.)
S al. c?. Ennis, Clare.
2 6. Kilmanock, Wexford.
Asker, Christiania, Nor-
1.
way. (E).
6 juv. Upland, Sweden. {G. Lord Lilford (p'
KoltJwff.)
1 al. Holland." {Scha Coll.)
2 6. Maredsous, Namur, Bel-
gium.
6. Eoret de Bouconne, Gers,
250 m. France. (-4.
Rohnrt.)
9. Cranves - Sales, Haute-
Savoie. (.4. Robert.)
6, 9. Burgheim, Bavaria, Ger-
many. {Woltcrstorff.)
6. Ingelheim, Rheinhesseu.
2 ^, 9 juv. St. Gallen, 500 m.
Switzerland. (E. H.
ZolUkofer.)
6, 9. Thurgau, 400 m. (E. H.
ZolUkofer.)
G. Barrett-Hamilton 11. 1. 2. 91-92.
C. H.B. errant (c&p). 11. 1. 3. 386.
W.T.Blanford(c&p). 11. 1. 3. 384.
Hon. Ella Scarlett 11. 1. 3. 385.
(c & P).
H. E. Forrest (c & p). 0. 9. 23. 1-3.
Dr. J. Anderson (p). 93. 7. 81. 1-2.
Dr. J. Anderson (p). 93. 7. 31. 3-4.
R. Drane (c ir p). 11. 1. 3. 387.
G. Barrett-Hamilton 11. 1. 2. 95.
(P).
J. W. Scott (c & p). 93. 10. 30. 1-2.
Ct. Barrett-Hamilton 11. 1. 2. 93-94.
(c & p).
Christiania Tiluseum 93. 3. 1. 7.
11. 1. 1. 151.
Lidth de Jeude Coll. 67. 4. 12. 555.
Rev. G. Fouruier (c 1. 6. 2. 1-2.
A- p).
O. Thomas (p). 6. 4. 1. 10.
O. Thomas (p).
Lord Lilford (p).
C. Hilgert (c).
0. Thomas (p).
0. Thomas (p).
6. 4. 2. 1.
11. 1.1.92,150.
8. 11. 2. 5.
4. 4. 5. 27-29.
4. 4. 5. 30-31.
Erinaceus europ.ecs nisp.wicus Barrett-Hamilton.
19C0. Erinaceus curopieus liisjianicus Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., v, p. 363, April, 1900. Type in British Museum.
1910. Erinaceus curojJeeus hispanicus Trouessart, Faune Mamm. d'Europe,
p. 39.
Type loculUi/. — Seville, Spain.
Geograpltical (listribution. — Iberian Peninsula.
Diagnosis. — Size large, essentially as in E. europseus europseiis
(condylobasal length of skull in old individuals about 60 mm.) ;
general colour paler than in the typical race, the fur often a
uniform light buffy grey, seldom with any evident darker area
between eye and muzzle.
Colour. — There is much variation in colour, .some individuals
essentially agreeing with the palest examples of true europseus.
In its extreme phase, however, the colour is a whitish buff",
decidedly paler than the cream-buff of Ridgway, the muzzle and
region about eyes washed with ecru-drab ; feet tinged with drab ;
S2)ines the same whitish buff, about half of them with scarcely any
ERINACEUS 123
dark shading, the rest with a drab sub-terminal band, the general
effect of spiny area scarcel)- siDeckled, and nearly as pale as fur.
In the type specimen the fur is a dull cream-buff, and the feet
are washed with broccoli-brown ; spines drab brown with light
tips, essentially as in true europseus.
Measurements. — Type : hind foot (dry), 40. Adult male and
female from Burgos, Spain : head and body, 270 and 250 ; tail,
30 and 28 ; hind foot, 43 and 41. Adult male and female from
Pajares, Leon : head and body, 252 and 249 ; tail, 22 and 21 ;
hind foot, 44 and 42. For cranial measurements see Table,
p. 125.
Specimens examined.— Twenty, from the followiug localities in Spain :
Arrechavaleta, Vitoria, 1 ; Pajares, Leon, 7 ; Burgos, 5 ; Palacios de la
Sierra, Burgos, 1; Bejar, Salamanca, 2; Seville, 4.
Bemarhs. — The Spanish hedgehog is a moderately well
differentiated form. Extreme specimens are easily distinguish-
able from typical europseus ; but in general the difference between
the two races must be regarded as an average one.
i. Arrechavaleta, Vitoria, 0. Thomas (p). 8. 2. 9. 9.
Spain. {N. Gonzalez.)
(J, 2 ?, 9 juv. Pajares, Leon. (N. Gem- 0. Thomas (p). 8. 2. 9. 2-6.
zalez.)
6, 9, 9 juv. Burgos. G. S. MiUer (c). 8. 8. 4. 17-19.
2 6. Burgos. (N. Gonzalez.) 0. Thomas (p). 8. 2. 9. 10-11.
2 6 juv. Bejar, Salamanca. {N. 0. Thomas (p). 8. 2. 9. 7-8.
Gonzalez.)
i, 2 9, 1. SeviUe. {Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 1-4.
[Type of subspecies 95. 3. 3. 2.)
Erinaceus eueop.eus italicus Barrett-Hamilton.
1857. Erinaceus europxus Blasius, Saugethiere Deutschlands, p. 153 (part).
1900. Erinaceiis europxus italicus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., v, p. 3G4, April, 1900. Type in British Museum.
1910. Erinaceus europasus italicus Trouessart, Faune Mamm. d'Europe,
p. 39.
Type localiti/. — Siena, Italy.
(• eographical distribution. — Italian Switzerland, Ital}' and
Sardinia ; Corsica ?
DuKjnosis. — Colour as in E. europseus europseus or slightly
paler ; size less than in the typical race, the largest skulls
probably not exceeding 59 mm.
Colour. — The colour is about as in the paler iudi^■iduals of
typical europseus, though the speckling of the spin}' area seems in
general to be finer, and the underparts usually lack all heavy
dark clouding.
Jleasureuients. — Adult female irom Curoggio, Ticino, Switzer-
land : head and body, 250; tail, 29 : liind foot, 41. Two adult
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126 INSECTIVORA
males from Siena, Italy : liead and body, 210 and 220 ; * tail, 30
and 28 ; ''' hind foot, 40 and 42.* Two adult females from the
same locality : head and body, 200 and 208 ; tail, — and 32 ;
hind foot, 43 and 38. For cranial measurements see Table,
p. 125.
Specimens examined. — Seventeen, from the following localities : —
Switzerland : Bigorio, Ticino, 1 (U.S.N.M.) ; Curoggio, Ticino, 1
(U.S.N.M.); Gentilino, Ticino, 1 (U.S.N.M.).
Italy : Empoli, Florence, 1 ; Siena, 4 ; Ostia, Rome, 1 ; Rome, 4.
Sardinia : Su Cramu, 1 ; Bare, 1 ; Marusei, 1 ; Trecorgia, 1.
Remarks. — Though not so pale as the Spanish race the Italian
hedgehog seems worthy of recognition as a form distinct from
true eurupseus. Its status is at present unsatisfactory, owing to
the lack of sufficient material ; but specimens from south of the
Alps seem never to attain the large size of Central European
adults. The Sardinian si^ecimens that I have seen are in general
paler than those from the mainland ; but here again the material
is insufficient. A hedgehog is known to occur in Cor.sica, but
no specimens have yet been compared with the Italian race.
9. Empoli, Florence, Italy. A. H. Savage Laudor 97.3.7.1.
(c & p).
2 ^, 2 9. Siena. (S. Brorji.) Dr. E. Hamilton (p). 98. 10. 2. 5-8.
(Type of subspecies 98. 10. 2. 5.)
1. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 7.
3 6. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 3-4. 96.
6. Su Cramu, Sardinia. (IT. O. Thomas (p). 0. 12. 3. 5.
WoUerstorff.)
6. Bare. (17. Woltersforff.) 0. Thomas (p). 0. 12. 3. 3.
9. Marusei. (W. Wolterstorff.) 0. Thomas (P). 0.12.8.6.
9 juv. Trecorgia. (ll'.iroZ^_Tsto>^.) 0. Thomas (p). 0.12.3.4.
Erinaceus europ.eus consolei Barrett-Hamilton.
1900. Erinaceus europxus consolei Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., v, p. 866, April, 1900. Type in British iluseum.
1910. Erinaceus europxus consolei Trouessart, Faune INIamm. d'Europe,
p. 40.
Type locality. — Palermo, Sicily.
Geographical distrihiUion. — Sicily.
Diagnosis. — Size apparently as in E. enrnp'ceus italicus (only
known specimen imperfect) ; colour differing from that of
italiciis in the uniform dusky brown head and shoulders ; quills
unusually robust, their dark and liglit markings strikingly con-
trasted.
Colour. — Whole head and sides of neck and shouldei'S a
uniform dark brown between hair-brow^i and sepia, sprinkled
with bufty grey hairs. The dark brown continues back along
* Type.
ERIXACEDS 127
edge of sj^iny area to tail, but throughout this region it is overlaid
by the uniform light cream-buff of underparts. Spines very dark
drab with light cream-buff tips, the light area shorter on spines
of mitldle of back than on those of sides, thus producing a slight
thougli evident darker median dorsal area. Feet so injured that
colour cannot be determined.
SJciiJI and teeth. — The imperfect skull shows no peculiarities.
Teeth as in specimens from the mainland.
Medsiirementit. — Type (sex not known) : head and body, 252 ;
tail, 50 ; hind foot, 40. For cranial measurements see Table,
p. 125.
Specimen examined. — The type.
Bemai-ks. — If not an abnormal specimen of E. europaeus italicns
the type of console! represents a very distinct local race.
1. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. G. 1.
{Tijpe of subspecies.)
ERINACEUS ROUMANICUS Barrett-Hamilton.
1900. Erinaceus europxus roiimanicus Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., v, p. 365, April, 1900. (Gageni, Roumania.)
Type in British Museum.
1901. Erinaceus danuhicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.
Freunde, Berlin, p. 229, December, 1901. (Prundu, Roumania.)
1910. Erinaceus europxus roiimanicus and E. curopxHs danuhicus Troues-
sart, Fauue Mamm. d'Europe, pp. 40-41.
Type locality. — Gageni, Pi-ahova, Roumania.
Geographical distribution. — From eastern Germany (Konigs-
berg) and northern Bohemia south through Hungarv, into
Greece.
Diagnosis. — In general like Erinacen.s europsens, but colour of
furred parts usually much darker, and chest often with a lai'ge,
conspicuously contrasted whitish area ; skull with upper length of
maxillary greater than depth of rostrum at middle ; teeth
essentially as in E. curopeeus, but first upper premolar with
postero-basal cusp usually more distinct.
Colour. — Furred area dark hair-brown interspersed with bufiy
and whitish hairs, the former most numerous on sides, the latter
forming a clear whitish pectoral area which sometimes spreads
over entire underparts. Feet and tail sepia. Face usually with
a seal-brown suffusion. Claws blackish horn-colour. Quills
with colours usually less contrasted than in E. europeeus, the
general effect of the spiny area darker and less speckled.
Shull and teeth. — The skull differs from that of E. eurrqjieus
in the relati^■ely longer, less deepened rostrum, a peculiarity
which seems chiefly to involve the maxillary bone. Upper length
of maxillary greater than depth of rostrum at middle. Posterior
1 28 INSECTIVORA
transverse palatal ridge usually lower and less developed,
particularly at middle, where there is often a slight angle.
Ridge at margin of orbit in lachrymal region not so long as in
E. curojjseus, and less distinctly marked off from general contour
Fig. 2-J.
Erinaceus rounuinicus. Nat. size.
of skull, its degree of development somewhat as in E. ahjirus.
Teeth not certainly distinguishable from those of Erinaceus
ruropseus, though anterior upper premolar is usually larger
relatively to canine, and its posterior cusplet tends to be better
developed.
Measurements. — External measurements of adult male and
female from vicinity of Konigsberg, Germany : head and body,
285 and 280; tail, 21 and 22; hind foot, 42 and 41. Type
(adult female) : head and body, 206 ; tail, 24 ; hind foot, 40-6.
Adult female from Corfu, Greece : head and body, 263 ; tail,
37 ; hind foot, 43 ; ear, 29. Adult male from Cephalonia,
Greece: head and body, 260; tail, 35; hind foot, 42; ear, 31.
For cranial measurements see Table, p. 132.
Specimens examined. — Eleven, from the following localities : —
Germany: Near Konigsberg, 3 (U.S.N.IM.).
KrsiNACEUS 129
Austria-Hunc4ary: Haida, Arva, Bohemia, 1; Vasoiir Eiscnbur^
Hungary, 1 (U.S.N.M.). °'
EoDMANiA : Gageni, Prahova, 2.
Greece: Corfu, 2; Cephalonia, 1 ; Tatoi, near Atheus, 1.
BemarJcs.— Though at first sight very similar to Erinacetis
europseus, this species is easily recognizable by its cranial characters.
In most specimens there is a strong contrast between the dark
posterior portion of underparts and whitish chest, throat and
shoulders, a pattei-n which appears to be rarely if ever well
developed in the related species.
9. Haida, Bohemia. Lord Lilford (p). 97. 8 14 1
i, 9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 15-16.
, „ ( "'• Dodson.) [Type of species 4. 4. 6. 16.)
.^. 1 otamos, Corfu, Greece. .J. I. S. Whitaker (p). 8 10 1 G
{C. Mottaz.) ■ ■ ■
■?. Argostoli, Cephalonia. J. I. S. Whitaker (p). S 10 1 7
(C. Motfaz.) ....
i. Tatoi, Athens. C. Mottaz (c). 8. 11. ,3. S.
ERINACEUS NESIOTES Bate.
1906. Erinaceim ciimjxns ncsioti's Bate, Proc. Zool. Soc. Loudon 1905 ii
p. 316, April 5, 1906. Type in British Museum. ' ' '
1910. Erinaceus europxvs ncsiotes Trouessart, Faune Mamm d'Eurone
p. 40. ^ '
Type locality. — Near Gonia, western Crete.
Gt'ograpMcal distribution.— Island of Crete.
-Dm»7«o.s;s.— Similar to Erinaccui^ roumanicm but smaller (hind
fo_(>t, 35 to 3S ; condylobasal length of skull in adult male, about
55 mm.) ; spines not so coarse as in the related species ; first
upper premolar distinctly larger than canine and of essentially
the same height.
(Jolour.—The colour is like that of E. roimnnicux. Under-
parts, sides and face dull whitish grey with faint dark clouding
between eye and muzzle, and in one of the three skins with a
dai-k wash on posterior half of under])arts.
Skull.— As in E. roumamnis, but not attaining as large size.
Mra.vu-ements.~Adu\t male"- and female from the type
locality: head and body, 208 and 204; tail, 29 and 19 • hind
foot, 40 and 38 (dry, 38 and 35). For cranial measurements see
lable, p. 132.
SjK'cimens f.vcnnined.—lhreL', all from Crete.
Bemarks.— The Cretan hedgehog is nearly related to Erinaccu>t
nmmuniruH, though well differentiated by its smaller size and by
the relatively large fourth unicuspid tooth.
'-, V. Gonia, Crete. Miss D. Bate (c). 5 10 o ii_i^
-,,,,. (5. 12. 2. 11 Type of speciesO ^
V. Mcsoghia. Miss D. Bate (c). 5. 12. 2. 13.
* Type.
130
IXSECTIVORA
ERINACEUS ALGIRUS Duvernoy and Lereboullet.
(Synon3-my under subspecies.)
Gcogrupliical distrihution. — Northern Africa ; also in southern
Spain, southern France, and the Balearic Islands.
Diagnosis. — Smaller than Erinaceiis europsens (hind foot less
than 40 mm.) : sjiines not so coarse ; middle of forehead with
bare area, among the spines ; skull with sagittal crest extending
forward to middle of frontal : a wide flattened area on each side
of bony palate behind transverse ridge ; third upper incisor with
two perfectly distinct roots ; elevated poi'tion of posteiior lower
premolar with only two cusps (fig. 2b h).
External characters. — Externally Erinaceus alijlnis is dis-
tinguishable from E. eurofseus by its smaller size, shorter, more
slender and apparently more densely-set bristles, and by the bare
area at middle of forehead. This bare area is about 7 mm. wide
and extends back about 10 to 15 mm. from front line of spines.
In dried skins it is sometimes partly hidden by shrinking. Fur
more dense and less coarse than in the larger animal. Claws on
front feet seldom attaining a length of 8 mm.
Colour. — The colour resembles in general that of the paler
races of E. euro^setis.
Slull. — The form of the skull is essentially as in E. i nropseus
except that rostrum is less elevated posteriorly, so that the dorsal
profile tends to become slightly concave. Sagittal crest when
fully developed extencUng forward to middle of frontal, while in
E. eiiropseiis it is usually confined to parietals, rarely encroaching
on posterior edge of frontal. Ridge at margin of orbit veiy short,
scarcely more than a process above lachrymal foramen. Bony
palate extending behind transverse ridge as a M-ell-defined Hat area
divided along median suture by a longitudinal ridge representing
the median spine of E. europaeus. Basisjahenoid pit narrower,
relatively deeper, and with more overhanging edges than in
E. enropsetis.
Teeth. — In general the teeth show no departure from those
of E. curopstus. The posterior lowei' pie-
molar, however, lacks all trace of the meta-
conid, so that the resemblance of the elevated
portion of the tooth to the first triangle .of
iH^ and 7/(.j is completely destroyed. In the
upper jaw the third incisor, canine, and
a h first premolar are two-iooted, apparenth'
i'lG. 25. without exception.
Large lower premolar of Measurements. — Head and body about
a.uiE.^aigiru!fh).'^i^l 200 to 250 ; tail, 25 to 40 : hind foot, 32 to
37 ; condylobasal length of skull, 54 to 59 mm.
Ilemarlcs. — This species is readily distinguishable from £v/«ace;f*"
europ»U8 by the bare spot among spines of forehead, the perfectly
two-cusped large lower premolar', and the t^\"0-rooted third upper
liRINACEUS 131
incisor. Although occurring wild in southern Spain, on the
Balearic Islands, and in south-eastern France, it seems not
improbable that the animal owes its presence in Europe to the
agency of man.
Ekixackus algirus AL(iiKus Duvernoy and LerebouUet.
1840. Erinaceus algirus Duvernoy and LerebouJlet, M6m Soc Mus
Q Hist. Nat. Strasbourg, iii, fasc. 2, p. 4.
1898. Erinaceus algirus de Winton, Proc. Zool. Soc. London, 1897, p. 955.
Type locality. — Oran, Algeria.
Geographical distribution.— 'i^orthern Africa ; also in .southern
Spain and south-eastern France.
J>ia(/Mos/s.— Condylobasal length of skull in individuals with
distinctly worn teeth 57 to 59 mm. ; underparts dusky through-
out or with at least an evident dark wash in interramial and
intercrural regions.
CoZor«»-.— Furred area buffy white to base of hairs, except on
muzzle, cheeks, interramial region, a narrow band aloncr sides
bordering quills and spreading posteriorly to cover tail hind
legs and intercrural region, all of which are a dark brown very
nearly the bister of Ridgway. Feet a lighter shade of the' same
brown. Occasionally the brown suffusion extends over most of
ventral surface. Quills dull horn-colour, each with a whitish
sub-terminal area about 7 mm. in length, the extreme tip usually
(ark. Throughout the spiny area the whitish strongly pre-
dominates, especially when animal is viewed from in front
Claws light yellowish horn-colour.
Measurement.s.~Adult male from Schaf-el-Kab, Morocco
(teeth much worn) : head and body, 206 ; tail, 2(i ; hind foot 32
For cranial measurements see Table, p. 1.32. '
Sp('ci77iens examined.— Nnmeious specimens from Northern Africa • also-
an adult from " Andalucia," Spain; a young, less than half grown, from
h.lche, Alicante, Spain ; and a still younger .specimen from Lecques, Var
Bcmarks.— Owing to the unsatisfactory nature of the Hpani.sh
and French material the status of the Continental European
hedgehogs of the Erinaceus algirus group is at present doubtful
Should they prove to be identical with tlie North African form
It would seem probable that they have been introduced within
historic times. The specimen from Elche, though undoubtediv a
wild-bred animal, is too young to be positively determined as' to
geographical race. The same is even raoi-e true of that from Var
The adult from " Andalucia " lacks detailed history. Externally
it resembles the African form as compared with E. ah/irns vaqans
but the skull is small,* es.sentially as in the Balearic" race. "
* Coudylolmsal length, 53-8; zygomatic breadth, 32-0; least inter-
orbital^breadth, 14-8 ; mandible, 41 -2 ; upper tooth-row, 27-0; lower tooth-
132
INSECTIVORA
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KRINACEUS 133
1. Andalucia, Spain. Lord Lilford (p). 94. 6 11 5
_i juv. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 1(3.
juv. al. Lecques, Var, France. Dr. P. Siepi (p). 98. 8. 25. 1.
Erinaceus ALGiRu.s VAGANs Thoma.s.
1901. Erinacens alr/irus imgans Thomas, Proc. Zool. Soc. London, p. 38.
Type in British Museum.
1910. Evinaccus algirus vagans Trouessart, Faune Mamm. d'Europe, p. 39.
Tyi^e locality. — Qsrn Cristobal, Minorca, Balearic Islands.
Geographical distribution. — Balearic Islands.
Diatjnosis. — Condylobasal length of skull in individuals with
worn teeth about 54 mm. ; furred area of body whitish through-
out, with no evident dark wash on face, in interramial or
intercrural regions, or on feet.
Measurements. — Type (adult male) : head and body, 250 ;
tail, 40 ; hind foot, 37. Adult male from Inca, Majorca : head
and body, 218 ; tail, 31 ; hind foot, 35. For cranial measure-
ments see Table, p. 132.
Specimens examine cl.~0\x6 from ^Majorca and five from Minorca
Balearic Islands. '
Bemarlcs. — The Balearic form of Erinaceus algirus is dis-
tinguishable from the African race by its paler colour and smaller
size. Its relationship to the animal occurring on the mainland
of southern Spain is not at present clear.
i. Inca, Majorca ; 300 m. O. Thomas & R. I. Pocock 0. 7. 1. 6.
Balearic Islands. (c & p).
4 6, 9. San Cristobal, ]\rinorca. 0. Thomas & R. I. Pocock 0. 7 1 35-39
(c & p).
(0. 7. 1. 36. Type of subspecies.)
134
CHIKOPTERA.
Order CHIROPTERA.
1779. Gldmptcra Blumeiibacli, Handbucb der Naturgescliichte, -p. 74.
Geofirajilrieal disirihutiou. — Practically cosmopolitan ; only
absent from the treeless arctic and antarctic I'egions, and from
the most remote islands of the Pacific and South Atlantic
oceans.
Characters. — Terrestial placental mammals with the anterior
limbs modified for true flight, the fingers greatly elongated (third
usually at least as long as head and body) and joined together
by a membrane which extends to sides of body and legs ;
shoulder girdle much more developed than pelvis, the sternum
usually keeled ; knee directed backwaixl.
Beniarhs. — ^The order Chiroptera, containing the only living
vertebrates, except birds, capable of true flight, is the most
sharply circumscribed of the main groups of mammals. Not
only are its living members invariably distinct from all other
recent forms, but the fossils also are, so far as known, equallj"
well differentiated. Therefore no intermediate stage has yet
been found connecting the bats with any other order. That
they are, however, not distantly related to the Insectirora, is
shown by numerous peculiarities of structure, among others the
relative!}' simple character of the brain. Two sub-orders are
recognized among the recent members of the order, the
MeijacMroptcra, not represented in Europe, with less highly
modified skeleton of fore limb, and more modified teeth, and the
Microchiropiera with more highly developed wing and in most
instances more primitive teeth.
Sub-Order MICROCHIROPIERA.
18'21. Inscdivora Gray, London Medical Repository, xv, x^. 299, April 1,
1821.
1872. Animalivora C-i-ill, Arrangement of the Families of Jlammals, p. IG,
November, 1872.
1875. Microcliiroptcra Dobson, Ann. and Mag. Nat. Hist., 4tli ser., xvi,
p. 346, November, 1872.
1878. Microcliiroptera Dobson, Catal. Ghiropt. Brit. ]Mus., p. 2.
1907. Microcliiroptcra Miller, Families and Genera of Bats, p. 78, June 29,
1907.
GeographicaJ di.vtribtit ion. -Thf: saiue as that of the order.
In Europe north to the limits of tree growth, west to Ireland
and the Azores.
MICROCHIROPTERA
135
Characters. — Anterior limb ^ery highly modified, the second
linger scarcely if at all independent of third, its ungual phalanx
never present, the humerus with trochiter and trochin large, the
former usually articulating with scapula ; mandible with angular
process well developed, long and narrow ; teeth usually not
modified for frugivorism (never in European species), the cheek-
teeth of upper and lower jaws very difierent from each other
(except when excessively reduced, as in the South American
Dcsmodontidse) ; margin of ear not forming a ring ; tragus usually
present. Mammaj in all European genera, jj 1— 1 = 2.
BemarJcs. — The sub-order Microchiroptera is essentially cos-
mopolitan in distribution. Though more highly modified than
the Megacliiroptcra in wing structure, the members of this group
foi' the most part I'etain the primitive tuberculo-sectorial type of
molar tooth, though certain South American frugivorous forms
show the stages through which the Megachiropterine molars
have probably passed. At jd resent 17 families and nearly 150
genera are recognized ; the species ai-e too imperfectly known to
permit any approximate estimate of their number. Three
families and ten genera are found in Europe.
KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES
OF MICROCHIROPTERA.
Tragus absent ; muzzle with leaf-like outgrowths ;
premaxillaries represented by palatal branch
only, not fused with surromiding parts (often
lost in prepared specimens) PJdnolophidx, p. 136.
Tragus present ; muzzle without leaf-like out-
growths ; premaxillaries represented princi-
pally (entirely in European genera) by nasal
branch, very early and completely fused with
surrounding parts.
Fibula robust, its diameter about half that of
tibia ; tail projecting conspicuously beyond
hinder edge of narrow interfemoral mem-
brane Molo&sidx, p. 276.
Fibula very slender, its diameter much less than
half that of tibia ; tail scarcely or not pro-
jecting beyond hinder edge of broad inter-
femoral membrane Vespcrtilionidie, p. 165.
Presternum with median lobe much smaller
than body of bone ; coracoid curved out-
ward ; second phalanx of third linger not
specially elongated Vespei-tilioninie, p. 165.
Presternum with median lobe larger than
body of bone ; coracoid straight, directed
inward ; second phalanx of third finger
nearly three times as long as first Miniopteriiue. p. 26S.
i;56
CHIROPTEEA
KEY TO THE GENERA OF EUROPEAN BATS.
(A wholly artificial key based primarily on external characters.)
^luzzle with leaf-like outgrowths Rhinolophus, p. 137.
Muzzle without leaf -like outgrowths.
Tail projecting conspicuously beyond membrane ... Xi/ctinomus, p. 276.
Tail not projecting conspicuoush' beyond membrane.
Ears joined.
Ear longer than head Plccotus, p. 25G.
Ear shorter than head Barbastella, -p. 268.
Ears separate.
Second phalanx of tliird finger nearly throe
times as long as first Miii'wptenis, p. 268.
Second phalanx of third finger less than twice
as long as first.
Fifth finger alsout as long as metacarpal of
fourth or third Nydalus, p. 242.
Fifth finger much longer than metacarpal of
fourth or third.
Ear wider than high, its lower margin
forming a small pocket near angle of
mouth Vcispciiilio, p. 2.38.
Ear higher than wide, its lower margin
not forming pocket near angle of
mouth.
Upper cheek-teeth 6-6 Mijotis, p. 166.
Upper cheek-teeth less than 6-6.
Upper cheek-teeth .5-5 Pipistrcllus, j). 202.
Upper cheek-teeth 4-4 Ei^tcsic/is, p. 224.
Family RHINOLOPHID^.
1827. Bhmulophina Lesson, Man. de jMammalogic, p. 81 (part).
1857. Phyllostomata Blasius, Saugethiere Deutschlands, p. 26.
1866. Rhinolophidai Gray, Proe. Zool. Soc. London, p. 81 (part).
1878. Bhiiiolophidie, Dobsou, Catal. Chiropt. Brit. Mus., p. 100 (x)art).
1907. RJiinolopliidx Jliller, Families and Genera of Bats. p. 106, .June 29,
1907.
Geographical clistrlbution. — Tropical and temperate portions of
the Old World from Ireland east to the Philippine Islands,
Solomon Islands and north-eastern Australia. In Europe north
to northern England and the Baltic coast of Germany.
Characters. — Ear without tragus ; muzzle with conspicuous
leaf-like cutaneous outgrowths {i\g. 26) consisting of a horizontal
anterior horseshoe, a perpendicular median sella, and a posterior
erect lancet ; skull with premaxillaries rejaresented by palatal
branches only, the two bones partly cartilaginous and not fused
with surrounding parts (often lost in prepared specimens) :
.shoulder girdle highly abnormal, the seventh cervical and first
dorsal vertebrae, first and second ribs, and presternum fused into a
continuous ring ; secondary articulation of humerus with scapula
small l)ut distinct ; fibula thread-like ; foot normal, the hallux
with two phalanges, the other toes with three.
liciaarJi-s. — The I\lnnolophid;r are the most widely distributed
HHINOLOPHUS
137
of the Old World leaf-nused bats, and the only t'ainil}' known
to occur in Europe. Notwithstanding its extensive distribution
and its large number of species the group is represented by a
single genus.
Genus RHINOLOPHUS Lacepede.
1799. Hliinolophiis Lacepede, Tabl. des div. sousdiv. oidrcs et genres des
Maminif^res, p. 15 {femtm-cqtiinum).
1836. Rhinocrepis Gervais, Diet. Pittoresque d'Hist. Nat. iv, pt. 2, jj. 617
(attributed to Geoffroy and Cuvier, Mag. Encyclop6dique, 1795,
but the name does not occur in the paper alluded to).
1847. Aquias Gray, Proc. Zool. Soc. London, p. 15 (lucius and irifoliatus).
1857. lihinoloplius Blasius, Siiugethiere Deutschlands, p. 26.
1866. Phyllotis Gray, Proc. Zool. Soc. London, p. 81 {pliilijiprnsis) not
Phijllotis Waterhouse, 1837.
1866. CcelophyUuH Gray, Proc. Zool. Soc. Loudon, p. 427 {coslophyUus).
1878. Rhinolophus Dobson, Catal. Chiropt. Brit. Mus., p. 100.
1901. Euryalus Matschie, Sitz.-Ber. GeseUsch. naturforsch. Freuude,
Berlin, p. 225 [mchelyi).
1904. Euryalus Matschie and Andersen, Sitz.-Ber. Gesellsch. naturforsch.
Freunde, Berlin, p. 71 (euryalc group).
1907. Jlhinolophus Miller, Families and Genera of Bats, p. 108, -Tune 29,
1907.
Type species. — Vespertilio ferrum-equinum Schreber.
Geographical distribution. — Same as that of family (p. 136).
Cliaractcrs. — Dental formula: « .7-^^ , c i^ , j)m |~| , )h |^ = 32.
Upper incisor very small, but usually well formed and with
distinct rounded crown with slight cusp on inner side. Lower
incisors tritid, the outer larger than inner, the four teeth
forming a continuous row between canines. Upper canine
heavy, but without secondary cusps or conspicuous cingulum.
Lower canine rather weak. Anterior upper premolar (jnn^) and
middle lower premolar {pm.^) small, functionless, usually crowded
Huite out of tooth-row. Other teeth showing no special peculiari-
ties ; m^ and m- ^\'ithout hypocone, m^ with live cusps and
three commissures (in many species a rudimentary- fourth com-
missure), the crown area much more than half that of m^ or
m'-. Skull with large brain-case and much shortened, globularly
inflated rostrum, beyond which the maxillaries, bearing the large
canines, conspicuously project ; palate so deeply emarginated
both anteriorly and posteriorly that its median length is less
than least distance between tooth-rows. Tail well developed,
e.xtending to edge of wide interfemoral membrane. Calcar
slender. Ears large, .separate, without tragus. Muzzle with
conspicuous leaf-like cutaneous outgrowths, consisting of a hori-
zontal anterior horseshoe, a perpendicular median sella and an
erect posterior lancet (fig. 26).
Remarks. — Among European bats the members of the genus
lihinoloplms are at once recognizable by the presence of the nose-
138
CHIP.OPTERA
leaf and absence of tragus. The skull dift'ers from that of all
other members of the fauna" in the short, globularly inflated
rostrum and long, projecting maxillaries, between which lie the
Xoseleaf of RhiiKilopIn'ifen-mn-equinum (a), li. hrppaxiilerus (b), 11. /■iii-yole (o),
and R. blasii{A). Nat. size.
horizontal free premaxillaries (often lost in prei)ared specimens).
About 100 forms have been described,* eight of which occur in
Eui'ope.
KEY TO THE EUROPEAN FORMS OF RHINOLOPHUS.
Noseleaf with connecting process broadly
rounded above ; skull with nasal swellings
long, rising gradually above line of fore-
head.
Forearm over 50 mm. ; condylobasal length ^
of skull over 20 mm. ; sella pandurate ;
large upper premolar in contact with
canine (Greater Horseshoe) B. ferrum-eqitiniim, -p. 139.
Wing relatively long ; forearm 54-58 mm.,
longest finger 84 to 92 mm. (Southern
and central Continental Europe) i:. f. ferruvi-eijuiinnn, p. 142.
Wing relatively shorter ; forearm 52 to
55 mm., longest finger 83 to 88 mm.
(England) Ji. f. inaulanus, -p. 147.
* See Andersen, Ann. and Ma,g. Nat. Hist.. 7th ser., xvi, pp. G48-662.
RHINOLOPHUS 139
Forearm under 43 mm. ; condylobasal length
of skull under IG mm. ; sella cuneate ;
large ujJiper premolar not in contact
with canine (Lesser Horseshoe) 7?. hipiiomdews, p. 147.
Greatest length of skull 14 • -5 to 15-5 mm.
(Jlediterranean region) it. h. minimiu^, ]}. 151.
Greatest length of skull more than
15* 5 mm.
Forearm 80 -3 to 39 mm. (England and
Ireland) n. /,. m!inau>f, p. 154.
Forearm 89 to 41 -7 mm. (Central Fjurope) j;. h. hiiiimxidemit, p. 149.
Noseleaf with connecting process acutely
pointed above ; skull with nasal swellings
short, rising abruptly above level of fore-
head.
First phalanx of fourth finger more than
half as long as second ; sella bluntly
cuneate ; no marked contrast between
crown areas of anterior and posterior
lower premolars (Eastern Mediterranean
region) li. blasii, p. 162.
First phalanx of fourth finger less than half
as long as second; sella parallel-sided,
broadly rounded above ; a marked con-
trast between crown areas of anterior
and posterior lower premolars.
Size smaller, forearm 44-6 to 49 mm.,
upper tooth-row 6*2 to 6*6 mm.;
gradation between phalanges of fourth
. finger abrupt (ratio of first to second
about 38) ; ])oint of lancet gradually
narrowed, never linear R. curtjalc, p. 155.
Size larger, forearm 48 '6 to 51 '4, upper
tooth-row about 7 mm. ; gradation
between phalanges of fourth finger
less abrupt (ratio of first to second
about 44) ; point of lancet linear 7?. mclicliji, p. 159.
RHINOLOPHUS FERRUM-EQUINUM Schreber.
(S^^lonymy under subspecies.)
(reograpliicdl (Jistribntion. — From southern Japan and China,
through the Hiiiiahxyas, the Mediterranean sub-region (exclusive
of Egypt), and central Europe to southern England (Andersen).
Diar/noftis. — Hize largest of the European species (forearm
more than 50 mni., condylobasal length of skull about 21 mm.,
mandible, 15 to IG mm.); noseleaf with horseshoe less than
10 mm. wide, the sella pandurate, the connecting process low,
abruptly rt)unded ; fourth finger with first phalanx considerably
mor(! than half as long as second ; large upper prenujlar broadly
in contact with canine, the small ])remolar minute (sometimes
absent), completely external to tooth-row.
Extern<d characters. — Size large and form leather heavy
(among the European members of the genus). General outhne
of noseleaf a rather elongate ovate-pyriform, the width of horse-
shoe slightly greater than distance from flat area at base of sella
140
CHIROPTERA
to ti]i of lancet. Sella broadly rounded at tip, noticeably con-
stricted somewhat above middle, the resulting outline pandurate.
Connecting process rising slightlj" but evidently above level of
sella, its upper border straight anteriorly, its tip rather abruptly
rounded off. Eai' large, extending when laid forward slighth'
beyond extremity of muzzle, abruptly narrowed to a rather acute
recurved tip ; antitragal lobe less than half as high as conch,
its width about eclual to its height, its upper border nearly
horizontal. Wings broad, the membrane attached to ankle.
First phalanx of foui-th linger noticeably more than half as long
as second. Foot slender, nearly half as long as tibia.
Colour.- — General effect a light greyish or drabby brown
pi'oduced by varying combinations of the pale ecru-drab under
colour and the darker hair tips, the region between ears, across
shoulders and at sides of lumbar region usually jialer than back.
The colour of the tips is usually either a clear, rather light .sepia,
or a buffy brown resembling the wood-brown of Ridgway. While
intermediate shades occvir the extremes are more often met with,
evidently representing two dichromatic phases. Underparts
usually rather lighter than back but never approaching whitish.
Slciill. — General outline of skull long and narrow, the breadth
of brain-case much less than twice that across canines, and con-
tained about 2 A times in greatest length ; zygomata projecting
slightl}' beyond general outline ; interorbital region greatly con-
stricted and conspicuously hour-glass shaped ; occipital portion of
brain-case noticeably overhanging foramen
magnum and marked oti' from main j^ortion
Iw a slight transverse depression corre-
sponding to suture between parietals and
unusually large interparietal ; sagittal
crest well developed, extending forward to
nari'owest portion of interorbital region :
lambdoid crest low but evident ; auditory
bulla' small, covering less than half surface
of large cochleae, between which the floor
of brain-case is reduced to a longitudinal
bridge less than 1 mm. in diameter :
mesopterygoid fossa slightly widei-
anteriorly than posteriorly, about li
times as long as wide, its rounded
anterior margin at level of postero-
internal angle of ;/(- ; palate with an
evident emargination on each side ex-
tending between m^ and mesopterygoid space ; anterior palatal
emargination extending back to line joining protocones of
anterior molars ; premaxillary ligulate, somewhat wider pos-
teriorly than anteriorly, the innei' border entire, the outer border
with a deep almost circular emargination posteriorly, the two
bones closely applied to each other along inner margin and to
y\i^£^
Jihinolopliusferrum-eqiiimi
Nat. size.
liHINOLCIPHUS
Ul
bottom of palatal emai'gination posteriorly, but otherwise free:
rostral inflation evident but low, its posterior border running
gradually into that of interorbital region, its anterior margin
over middle of anterior molar ; maxillary triangular in outline
when viewed from the side, the heavy canine projecting forward
and downward from its anterior apex ; anteorbital foramen small,
over middle of second molar and directly beneath minute
lachrymal foramen, the plate forming outer wall of canal
thread-like, occasionally absent.
Teeth. — Except for the minute ]iremolars and upper incisors the
teeth are robust and heavy relatively to size of skull. Upper
incisor minute, low, the crown subterete, wider than root, lower
externally than internally, each tooth placed near middle of
oblique anterioi' border of premaxillary, the space between the
two nearly double diameter of crown. Lower incisors strongl}-
imbricated, forming a short, very convex row between canines,
their ci-owns longer than high, compressed (the outer tooth less
than the inner), deeply and et|ually trilid. Upper canine very
large, and noticeably the highest tooth in the maxillary series,
the root oblique, the shaft abruptly bent downward at level of
well developed cingulum, its length along cinguluin about three-
fourths height ; cross section of shaft triangular, the inner surface
flattened, slightly concave neai' cingulum, the posterior edge
trenchant, the antei-ior edge narrowly sub-ti'enchant, the outer
surface with well developed median longitudinal ridge and
noticeable posterior concavity. Lower canine not so large as
upper, the flattened surface of its shaft directed posteriorly to
o})pose front of upper canine in mastication. Anterior uppei'
premolar minute, resembling upper incisor in both size and form,
entirely external to tooth-row, occasionally absent. Posterior
uj)per premolar large, closely crowded against canine, its crown
area about equal to that of second molar, without secondary
cusps, its main cusp intermediate in height between canine and
metacone of first molar ; posterior border of crown slightly but
evidently emarginate. Anterior lower premolar less than half as
high as posterior premolar and with barely half its crown area,
the two teeth crowded closely together between canine and first
molar ; shaft of anterior tooth with slightly developed concave
area on inner side, that of posterior tooth with better defined
posterior concavity, its inner side convex ; middle lower premolar
resembling anterior upper premolar, its position equally external
to tooth-roM'. Upper molar with large though not unusually high
protocone, behind which in m^ and iii- there is a low-lying
lieel (best developed in »(*), but no indication of a true hypocone ;
paracone and metacone well developed, the latter slightly the
higher in m^ and m-, this reversed in m^ ; styles and commis-
sures in m^ and m- well developed, forming a normal VV-pJittern :
in »K^ the metastyle and fourth commissure are absent, and the
third commi.ssure is much reduced in length : crown area of in^
142 CHIROPTEEA
about two-tliirds that of //('- ; lower molars with protoconid
noticeably higher than the other cusps, the posterior triangle
slightly the wider of the two in m,^ and lu.^, a little the narrower
in m^ ; behind entoconid the cingvdum forms a minute though
evident accessory cusp.
Bemarks. — Among the leaf-nosed bats of Europe this species
is easily recognizable by its large size taken in connection with
the low, rounded upper margin of the connecting process of sella.
In B. meliehji, which approaches it in size, the connecting process
is sharply pointed above. Two geographical races ai'e known, one
occupying the Continental range of the species, tlie other confined
to Great Britain.
Rhinolophus ferrum-equinum februm-equinum Schreber.
1774. Vcspertilio fcrntm-cqiiimtm Sclireber, Siiugthiere, i, pi. lxii, upper
figures ; description, i, p. 174 under name : Die Hufeisennase
(part). Prance ; based primarily on Daubeuton.
177G. Vcspertilio equinus P. L. S. Miiller. Natursyst. Suppl. u, Kegist.-
Band, p. 20 (part), France.
1779. [T'esperiiZJo] perspiciUatus Blumenbacb, Handb. d. Naturgescb., p. 75
(part : included the leaf-nosed bats of Europe and Soutli America).
1785. [Vespertilio] ungula Boddaert, Elenchus Auimaliuni, i, p. 71
(Burguud}-).
1792. Vcsp\crfilio] fer[fnni\-equ[inuvi] viajor Kerr, Anim. Kingd., p. 99
(not 1'. molossus major Kerr, I.e., p. 97), Prance.
1798. Vespertilio hippocrcpis Sehrank, Pauna Boica, i, p. G4 (Renaming
of fcrrum-cquinum Schreber).
1803. Bhinolophiis major Geoffroy, Catal. Mamm. ;\Ius. Nat. d'Hist. Nat.,
Paris, p. 50 (Burgundy).
1S13. Bhinolophiis unihastatus (4cofii'oy, Ann. INIns. d'Hist. Nat., Paris,
xs, p. 257 (France).
1829. '! Ehinolophns nnifer Kaup, Entw.-Gcscb. u. Natiirl. Syst. Europ.
Tbierwelt, i, p. 104 (nomen nudum).
1857. Bhinolophus fcrrnm-cquinuni. Blasius, Siiugetliiere Deutschland.^.
p. 31.
18G3. \_Bhinolvphus frrrum-cqiiiniiiiij a var. qcniiaiuciis Kocli, Jalirb. des
Vereins fiir Naturlcunde im Herzogtlium Nassau, xviii, p. 522
(Wiesbaden, Hesson-Nassau, Germany).
18G3. [Blmiolophns fcrriaii-cqiiiniim] B var. italiciis Koch, Jalirl). des
Vereins fiir Naturkunde im Herzogthum Nassau, xviir, p. 523
(Italy).
1878. Bldnolopihiis fvrriiin-cqiiinum Dobson, Catal. Cliiropt. Brit. Mus.,
p. 119 (part).
18S5. Bhinolophus ■unihastatus^ var. hoinorodahnasicnsis Daday, Orvos-
Termeszettudomanyi Erteset(i, Kolozsvar, x, p. 274 (Homorod-
ALmas cave, Hungary).
1886. Bhinolophus nniliastatus var. Jiomorodalmasicnsis Daday, Verhandl.
u. Mittheilungen des Siebenbiirgiseben Vereins fiir Naturwissensch.
in Hermannstadt, xxxvi, p. 79.
1887. Bhinolophus fcrriim-cquinum var. homorodcnsis Daday, EIrtekezesek a
Term^szettudomanyok Kor^bi'd, Budapest, xvi, pt. 7, p. IH
(Renaming of homorodalniasiensis).
RHINOLOPHUS
143
1904. Bliinolophus ferrtiin-equinuvi obscurus Cabrera, ilem. Soc. Espafi.
Hist. Nat. Madrid, ii, p. 257 (Valencia, Spain).
1905. Bhinolophus ferrum-cquinum hjpicus Andersen, Proc. Zool. Soc.
London, 1905, ii, p. 113, October 17, 1905.
1905. Bhinolophus femtfn-equinum obscurus Andersen, Proc. Zool. Soc.
London, 1905, ii, p. 116, October 17, 1905.
1910. BMvolophus feirmn-equinum, B. fcmmi-equinum obscurus and B.
ferrum-equinum homowdensis Trouessart, Faune IManim. d'Europe,
pp. 4-5.
Type loralitji. — Burgundy, France.
Geographical distribution. — Central and southern Continental
Europe.
Diagnosis. — Wing relativel}^ long ; forearm, 54 to 58 mm. ;
third finger, 84 to 92 mm.
Measurements. — Average and extremes of six males from
8ilos, Burgos, Spain: head and body, 62 •! (58-64 '4); tail,
39-9 (37-6-43); tibia, 23-2 (21-8-24); foot, 11-7 (11-13);
forearm, 55-1 (54-8-55-4); third finger, 89-3 (88-92); fifth
finger, 72-5 (72-73); ear from meatus, 24-9 (24—26). Average
and extremes of five females from Silos, Burgos, Spain : head
and body, 61-8 (60-66); tail, 38-8 (37-42); tibia, 23-1
(22-6-24); foot, 11-7 (11-13) ; forearm, 56-2 (56-57) ; third
finger, 89-6 (88-91); fifth finger, 73-4 (71-76); ear from
meatus, 25-1 (24-8-26). Two males from Granada, Spain:
forearm, 54 and 54. Two females from the same locality :
forearm, 56 and 58. Two males from Elche, Alicante, Spain :
forearm, 54 and 55. Average and extremes of six adults
(3 males and 3 females) from St. Genies, Gard, France : tibia,
24-1 (23-4-24-6); foot, 12-1 (11-8-12-8); forearm, 55-4
(54-65) ; third finger, 87-3 (84-91) ; fifth finger, 71-5 (69-74).
Female from Marseilles, France : forearm, 57. Male and female
from near Genoa, Italy : forearm, 57 and 57. Two females from
Kimini, Ital}^ : forearm, 53 and 57. Female from Siena, Italy :
forearm, 54. Male from Rome, Italy : forearm, 56. Male from
the Parnassus region, Greece : forearm, 54. Adult female from
Tubingen, Wiirtemberg, Germany : head and body, 63 ; tail, 37 :
tibia, 24-4; foot, 11; forearm, 56-6; third finger, 88; fifth
finger, 72; ear from meatus, 24; width of ear, 16. Adult
female from Ofener Mts., Hungary : head and body, 61 ; tail, 40 ;
tibia, 24; foot, 11-6; forearm, 56; third finger, 87; fifth
finger, 73; ear from meatus, 23-4; width of ear, 16. Two
males from Herkulesbad, Hungary : forearm, 56 - 6 and 58. Four
females from Herkulesbad, Hungary: forearm, 56, 56-4, 56*6
and 57-4. For cranial measurements see Table, p. 144.
Specimens c.vamincd. — One hundred and forty-one, from the following
localities : —
Portugal: Cintra, 1.
Spain : Silos, Burgos, 22 ; Granada, 4 ; Elche, Alicante, 3 ; San
Cristobal, IMinorca, Balearic Islands, 4.
144
CHIROPTEKA
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CO CM 00 CI C' CO -ti
.H rH T-H 0-1 CM tH CM
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ft w
RHINOLOPHUS
145
S .SO
.■M
o
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<M
o
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<M
cq
<M
o
<M
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<M
o
o
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cq
CS
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03
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OS
05
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cq
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'SI
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oq
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14G
CHIKOPTERA
Fkaxce : Troubate, Hautes-Pyrenees, 6 ; St. Genios, Gard, 4:3 (ISIottaz) ;
Marseilles, 1 (U.S.N.M.) ; Meounes, Var, 1.
Italy: Turin, 1; near Genoa, 8 (B.M., U.S.N.M. and Genoa); Grotta
de Isoverde, Liguria, 7 (B.M. and Genoa) ; Rimini, 2 (U.S.N.M.) ; Siena, 1
(U.S.N.M.); near Rome, 5 (U.S.N.]\I. and Genoa); no exact locality,!;
Sicily, 2.
Corsica : Commune di Barbaggio, 1.
Sardinia : Sassari, 1.
Greece : Parnassus region, 1 (U.S.N.M.) ; Island of Syra, Cyclades,
1 (U.S.N.M.) ; Mt. Goria Monastery, Crete, 3.
Germany : Tiibingen, Wiirtemberg, 1.
Switzerland : Geneva, G (U.S.N.M. and J\Iottaz) ; Boiidry, Neuchatel,
2 (Mottaz) ; Tremona, Ticino, 1 ; Mendrisio, Ticino, 2 ; Lugano, Ticino, 3.
ACSTRIA-HUNGARY : Herkulesbad, Hungary, G ; Ofener Mts., Hungary, 1.
BeiaarJcs. — Spanish specimens of BhinolopJiuti ferruui-equinum
have been regarded by both Cabrera and Andersen at represent-
ing a pecuhar race, R. f. ohscuius, distinguished by small size.
The material which I have examined, however, indicates that
the Iberian animal cannot be treated as distinct.
6.
Cintra, 500 mm. Portug
;al.
0. Thomas (c A- p).
98. 2. 2. 1.
G al.
Silos, Burgos, Spain.
N. iV- S. Gonzalez
8. 7. 7. 38-4.3.
2 0, 2 V.
Silos, Burgos.
G. S. MiUer (c).
8. 8. 4. 1-4.
6, 9.
Granada, 2250 ft.
G. S. Miller (c).
8. 8. 4. 8-9.
2 i, ?.
Elche, Alicante, 20 m.
G. S. Miller (c).
S. 8. 4. 5-7.
6, -2 V.
San Cristobal, Minorca,
learic Islands.
Ba-
0. Thomas and
R. I. Pocock
(c \- P).
0. 7. 1. 24-26.
6 al.
San Cristobal, ;\Iiuorca.
0. Thomas and
R. I. Pocock
(c & P).
0. 7. L G8.
G c^..
Troubate, Hautes-Pyrenees,
0. Thomas (p).
G. 4. 1. 1-6.
France. [A. Robert.)
1 al.
Meounes, Var.
Dr. K. .Jordan
(c & p).
Prof. Bonelli (p).
8. 3. 15. 1.
V al.
Turin, Italy.
9 al.
Isoverde, Genoa.
0. Thomas (CvV p).
88. 12. 7. 1.
Skeleton.
Italy. (I'riuce Bonaparte.)
Tomes Collection.
7. 1. 1. 727.
2 6 al.
Sicily.
rfal.
Barbaggio, Corsica.
Dr. C. I. Forsyth
Major (c & p).
6. 4. 14. 1.
6a\.
Sassari, Sardinia.
Marquis G. Doria
6. 12. 1. 11.
6, 2 9.
Mt. Goria Monastery, Crete.
Miss D. Bate (c).
5. 12. 2. 1-3.
9al.
Tiibingen, Wiirtemberg, Ger-
Dr. A. Giinther
66. 2. 1. 1.
many.
(P)-
9.
Tremona, Ticino, Switzerland.
0. Thomas (p).
2. 8. 4. 1.
[E. H. Zollikofer.)
2 9.
Mendrisio, Ticino. {E.
Zollikofer.)
H.
0. Thomas (p).
2. 8. 4. 2-3.
3.
Lugano, Ticino. {E. U. Z>
kofer.)
olll-
0. Thomas (p).
4. 4. 5. 1-3.
2 i, 4 9.
Herkulesbad, Hungary.
Hon. N.C.Roths-
child (p).
7. 9. 16. 1-6.
9al.
Ofener :Mts., Budapest.
Budapest iluseum
(E).
94. 7. 18. 5.
RHINOLOPHUS
147
JviiixoLOPHUS PERKUM-EQUINUM iNsuLANUs Barrett-Hamiltoii.
I'JJO. J titiiiolophns ferriivi-equimivi insulamts Barrett-Hamilton, Ann. and
Mag. Nat. Hist., 8tli ser., v, p. 292, March, 1910. Type in British
?tluseum.
lyiO. Rlnnoloplius ferriim-cqitiiium ijtsulaiiits Trouessarfc, Faune Mamm.
d'Europe, p. 273.
Ti/jtc locdlifij. -Clieddar, Somersetshire, Englaiul.
(T/'oiiraphiral (lii^frihuiion. — Central and southern England.
DlitfjDOsix. — Wing relatively short ; forearm, 52 to 55 ; third
finger, S.3 to 88.
Mfamireinents. — Type (adult male) : head and body, 67 ; tail,
•■i7 : tibia, 23-4 ; foot, 11 ; forearm, 54 ; third finger, 80 ; fifth
finger, 70; ear from meatus, 23 '6. Average and extremes of
seven males from the type locality : head and body, 65 (63-67) :
tail, 36-9 (35-40): tibia, 22-8 (22-24); foot, 11-2 (11-12):
forearm, 53 '5 (52-54) ; third finger, 85-5 (83-87) : fifth finger,
Gii'9 (69-71) ; ear from meatus, 24*2 (23-25). Tliree females
from the type locality : head and body, 64, G6 and 64 : tail, 35,
34 and 36; tibia, 23-4, 23 and 23; foot, 10-6, 12 and 12:
forearm, 55, 54*6 and 54: third finger, 87, 86 and 88: fifth
finger, 72, 72 and 72 ; ear from meatus, 25, 24 and 24. For
cranial measurements see Table, ]>. 144.
Specimens examined. — Twenty-four, from tlie following localities in
England : — Cheddar, Somersetshire, 10 ; Wells, Somersetshire, 10 ; Dorset-
shire, 3 ; Bonclmrch, Isle of Wight, 1.
5, 9. Cheddar, Somerset, J. A. Coward (c & p). 7. 1. 10. 1-2.
England.
(J al. Cheddar. • -T. A. Coward (c & p). 7. 1. 10. 3-10.
<, 2 9 al. Wells. Hon. N.C. Rothschild 5.1.23.1-3.
3 i. 6 juv., 9 St. Wells. Hon.N. C.Rothschild /I. 9. 3. 1-4.-
(p). \2. 9. 6. 1.
.^,9. Wells. Hon. N.C.Rothschild 11.1.3.3-4.
(P)-
2 9. Dorset. W. M. Hardy (c & p). 11. 1. 3. 1-2.
1. Bonchurch, Isle of Rev.C. A.Burv (c A" p). 11. 1. 3. 388.
Wight.
RHINOLOPHUS HIPPOSIDEROS Beclistein.
(Synouymx under subspecies.)
(icoijraphiral (Uxfrihittioii. — From Uilgit through the Mediter-
ranean sub-region and central Europe to Ireland ; north in
continental Europe to the Baltic, and in Great Britain to about
the southern border of Scotland.
DifKjnosix. — Size small, the forearm less than 43 mm. in
length, condylobasal length of skull about 14 to 15 mm., mandible
about 10 mm. : no.seleaf (fig. 26 b) with bluntly cuneate sella
and low , broadly rounded connecting process ; fourth finger with
L 2
148 CHIROPTERA
first phalanx slightly more than half as long as second ; large
upper premolar separated from canine by a noticeable space
occupied by the well developed small premolar, which lies per-
fectly in the tooth-row.
External characters. — A much smaller, more delicately formed
animal than Uliinoloplms ferni/in-equimim. General outline of
noseleaf nariower than in H. fcrriim-equinum, the width of horse-
shoe less than distance from flat area at base of sella to tip of
lancet ; sella narrowly rounded at tip, the sides straight, slightly
convergent above, the resulting outline bluntly cuneate ; connect-
ing process essentially as in the larger sjjecies but relatively
wider and lower, its upper extremity about on level with that of
sella ; lancet slender, scarcely or not contracted at middle, the
tip cuneate. Ear when laid forward extending about o mm.
beyond extremity of muzzle, the narrow tip abruptly curved
backward ; antitragal lobe more than half as high as conch, its
width less than its height, its upper margin noticeably obli(iue.
Wings and feet essentially as in B. ferruvi-equinavi.
Colour. — The colour does not difl'er appreciably from that of
Rliinoloplivs ferrum-equinum.
Skull. — The skull differs from that of Blii)wlophu.s ferruui-
equinum chiefly in its conspicuously smaller size (greatest length
about 16 mm. instead of about 24 mm.). There are also some
slight peculiarities in form, the principal of which is the greater
contrast between width of brain-case and anterior maxillary
region, the breadth of former being about twice that of latter.
As the breadth of brain-case is contained distinctly less than
2.1 times in greatest length it is probable that the diffei-ence
between the two animals is due rather to enlarged brain-case in
the smaller than to widened palate in the larger. Sagittal crest
slightly develojied ; lambdoid crest obsolete. Nasal region
relatively more inflated than in J?, ferrum-equinum, particularly
at side of nares, but dorsal outline not rising abruptly above
level of interorbital region. Mesopterygoid fossa so conspicu-
ously widened anteriorly that it occupies almost entire extent of
palate between posterior molars, leaving no space for lateral
palatal emarginations, a peculiarity by which the skull may be
distinguished from that of all other European members of the
genus.
Teeth. — Though in general resembling those of Mhiiiohq/hu.s
ferrum-equinum, ajmrt from their much smaller size, the teeth of
B. hipposideros show several notable peculiarities. Upper canine
relatively small, its apex in line with large cusps of molars, anrl
slightly below that of posterior premolar ; lower canine corre-
spondingly .short ; anterior upper jiremolar a well develojjed
functional tooth lying perfectly in tooth-row, with crown area
equal to nearly half that of canine, its shaft subterete though
flattened posteriorly, and only a little less than half as high as
main cusp of large jiremolar : anterior and posterior lower pre-
RHINOLOPIIUS
149
molars less contrasted in size than in the larger animal, and
separated from each other by a slight space in which lies
the minute middle premolar, less
crowded outward than in B. ferruvi-
eqwiniim. Large upper premolai
and upper molars essentially as in
R. ferrum-eqmnum except that in
m^ there is a distinct meta-
style and fourth commissure,
and area of tooth is nearly
equal to that of m"^. Lower
molars as in Hhinolophus
ferrKrn-t'quinum, except that
in M«3, almost exactly resem-
bles the other teeth, the second
triangle having undergone
practically no reduction.
Measurements. — Head and
body about 40 mm., tail about
.'jO mm., forearm, 34-5 to
4rl • 7 mm.,condylobasal length
of skull, 13 "8 to 16 mm.
(greatest length 1 4 • 5 to 1 6 • 2).
Details under sub.species.
Bemarks. — Three imperfectly differentiated forms of Bhino-
lophus hipposideros occur in Europe : a larger central race, a
smaller Mediterranean race, and an intermediate form peculiar
to Great Britain and Ireland. Without examination of much
more complete material than that now available it is impossible
to reach any wholly satisfactory conclusion as to the status and
interrelationships of these forms. The characters here given are
those published by Andersen in 1905.*
Fig. 2S.
ItliiiiolnpliHS Jiipposidems. Teetli X 10
Rhinolophus hipposideros hipposidkros Bechstein
1789. Vespertilio ferrum-equinum B., Die kleine Hufeiseunase, Bechstein.
Gemeinn. Naturgesch. Deutschlands, i, 1st ed., p. 186.
1792. Vcsi}[crtilio] fcrlnimyequlinum^ minor Kerr, Anim. Kingd., p. 99
(not 1". molossun minor Kerr, I.e., p. 97) (France).
1800. Vespertilio hijjposidcros Bechstein, Thomas Pennant's .\llgemeine
tJebersicht der vierfiissigen Thiere, ii, p. 6'29.
1803. lilmiolophus minor Geoffrey, Catal. ]\Iamm. !Mus. Nat. d'Hist. Nat.,
Paris, p. 57 (Neighbourhood of Paris).
181:5. Rhinolophus bihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xx,
p. 259 (Neighbourhood of Paris).
1816. ? I'hijllorhina mimtia Leach, Syst. Catal. Spec. Indig. Mamm. and
Birds Brit. Mus., p. 5 (uomcn nudum : " Small Leaf nose ").
* For further discussion of the subject see Mottaz, M6ni. Soc. Zool. de
France, Paris, xx, pp. 21-22, September, 1907 ; Andersen, Ann. and ^lag.
Nat. Hist., 7th ser., xx, pp. 384:^89, November, 1907.
150 CHIROPTERA
1S29. ? B]dnolo2)hubi bifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ.
Thierwelfc, i, p. 104 (nomeu nudum).
1S57. ? Rhinolophus hipposideros Blasius, Siiugetbiere Deutschlauds, p. 29.
1863. illhinolophus hipposideros] a var. typus Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 5.30 (Wiesbaden).
1863. [Rhinolophus hijiposidcros] var. alpinus Koch, Jahrb. des Vereius
fiir Naturkunde im Herzogthum Nassau, xviii, p. 530 (Alps).
1870. niiinoloj^ltus cggcnlioffncr Fitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Clasae, lxi, Abth. i, p. 151
(MS. synonym of bihastatas).
1878. Rhinolophus hipposideros Dobson, Catal. Chiropt. Brit. Mus., p. 117.
1885. Rhinolophus bihastatus var. kisnyiresicnsis Daday, Orvos-Terme'sz-
ettudoniAnyi Ertesitti, Kolozsvar, x, p. 274 (Kis-Nyires, Szolnok-
Dobaka, Hungary).
1886. Rhinolophus bihastcdus var. kisnyiresiensis Daday, Verhandl. u.
Mittheilungen des Siebenbiirgischen Vereins fiir Naturwisseusch.
in Hermannstadt, xxxvi, p. 80.
1887. Rhinolophus hipposideros var. troglophilus Daday, Ertekezesek a
Term6szettudomdnyok Korebol, Budapest, xvi, pt. 7, p. 8 (Re-
naming of kisnyircsiensis) .
1904. Rhuwlophus euryalc helvetica Bretscher, Vierteljahrsschrift der
Naturforsch. Gesellsch. in Ziirich, xlix, p. 256. See Mottaz,
Bull. Soc. Zool., Geneve, i., p. 172, 1908 (Baar, Zug, Switzerland).
1905. Rhinolophus liipposidenis ti/picns Andersen, Proc. Zool. Soc. London,
1905, II, p. 141, October 17, 1905.
1910. Rliinolophiis Idpposidcrns Trouessart, Paune ]Mamm. d'Europe, p. 9.
Tyije localitij. — France.
Oev'jrajjhical distribution. — Central Europe, north of the Alps,
ea.st through Ai'uienia and north-west Persia to the Himalayas.
Diagnosis. — Forearm* usually 39 to 41 '7 mm.; greatest
length of skuUf about 16 mm.
Mcasnrevients. — Two males from 8trass near Burgheim,
Bavaria, Germany: tibia, 17 and 18*4; foot, 7 '8 and 7*4:
forearm, 38-8 and 39-2 ; third finger, 61 and 60 ; fifth finger,
54 and 53. Female from the same locality: tibia, 17'4 ; foot,
7-4; forearm, 38-8; third finger, 60: fifth finger, 53. The
six following extremes of forearms of Eurojuean specimens are
given by Andersen (I.e. p. 142) : N. Bulgaria (1), 39 ; Roumania
(13), 39 to 41-2 ; Transsjdvania (2), 40 to 41 ; S. Carj^athians
(1), 39-3; Schlangenbad, Nassau, Germany (2), 40 to 40*1 :
Strassburg, Germany (3), 39 to 40*1. Forty-nine males from
the vicinity of Geneva, Switzerland (Mottaz collection), J ."7 -3
to 39. Thirty females from the same locality (Mottaz collection), J
38-5 to 40-6. Four males from the same locality (U.S.N". M.),
37 to 38-4 Six females from the same locality (U.S.N.M.),
37*4 to 40. Adult female from Dions, Gard, France (Mottaz
collection), 40. For cranial measurements see T\'ible, p. 152.
* Bases of metacarpals included.
t From back of occiput to front of canine.
% Measured by Chas. Mottaz (base of metacarpals included) and verified
by Gerrit S. Miller.
RHINOLOPHUS
1.11
Specimens examined. — One huudred and seventeen, from the following
localities : —
France: Dions, Gard, 1 (Mottaz) ; St. Cergues, Haute-Savoie, 1.
Germany : Strass, near Burgheim, Bavaria, 3 ; Mainz, 1 (Strassliurg) ;
Bitsch, Alsace, 1 (Strassburg).
Austria-Hungary : Hatszeg, Hiinyad, Hungary, 1.
Switzerland: Near Geneva, 9-i (B.M., U.S.N.M. and Mottaz); St.
Jloritz, 1 ; Thayngen, Schaffhausen, 2 (U.S.N.M.) ; Roggwil, Thurgau,
7 (B.:M. and U.S.N.M.) ; Canton Thurgau, 5 (B.M. and U.S.N.M.).
6. St. Cergues, Haute-Savoie, A.Robert (c & p). 5.4.9.1.
France.
■2 ?. Burgheim, Bavaria, 375 m. Lord Lilford (p). 11. 1. 1. 117-
Germanv. 118.
c5. Hatszeg, Hunyad, Tran- C.G.Danford (c). 8.11.8.1.
sylvania.
3 6. Grand Pr^, Geneva, Swit- C. jMottaz (c & r). 6. 2. C. 1-3.
1. St. Moritz, Grisous. Leon O. Galliard 75. 9. 20. 3.
(c X- P).
3 i, 2 ?, iram. Roggwil, Thurgau. (E. H. O. Thomas (p). 2. 8. 4. 4-9.
Zollikofer.)
9al. Ihurg&u. {E.H. Zollikofer.) 0. Thomas (p). 2.8.4.55.
RiiixoLOFiius HiPPOSiDERCs MINIMUS Heuglin.
1861. Rhinolophus minimus Heuglin, Nov. Act. Acad. Caes. Leop. -Carol.,
XXIX, Abliandl. viii, p. 6 (articles separately paged) (Ker^n,
Abyssinia).
18G'B.' [RhinoloiJhus hipiM.ndcro:^] y y&r. piallid-us Koch, Jahrb. des Vereins
fiir Naturkunde im Herzogthum Nassau, xviii, p. 531 (Mediter-
ranean region).
1904. Bhinolophus phasma Cabrera, Mem. Soc. Espan. Hist. Nat., Madrid,
II, p. 252 (Madrid, Spain).
1904. Tili[inolophus] h[ippiosidcnis] minimus .\ndersen, Ann. and ^lag. Nat.
Hist., 7th ser., xiv, p. 456, December, 1904.
1905. Rhinolophus hij^posidcrits minimus .\ndersen, Proc. Zool. Soc.
London, 1905, ii, p. 140, October 17, 1905.
1910. RJiinolophus hippiosidci'us minimus Trouessart, Faune Maram.
d'Europe, p. 10.
Ti/jje lornliiij. — Keren, Abyssinia.
Geofjraphical distribution. — Mediterranean region.
Diagnosis. — Forearm usually 34 ■ 7 to .'58 mm. ; greatest
length of skull, 14-5 to 15 '5 mm.
Measurements. — The following measurements of forearms are
given by Andersen (I.e. p. 141) : Keren (type), 86' 3 ; Cvprus (6),
34-7 to 37-7 ; Malta (8), 36 to 37 ; Ostia, Italy (2)," 35-7 t«
36-8; Corsica (1), 37-7; Balearic Islands (7), 36-2 to 37-6;
Seville, Spain (1), 37*7 ; Cintra, Portugal (1), 36 -2. Specimens
in U.S. National Museum : Ficuzza, Sicily ( ? ), 35 • 2 ; Rome ( ^ ),
36-4; Siena (3 $), 36-8 to 38; Marseilles, France ( ? ), 38;
Dions, Gard, France (cJ), 34 '8, (9), 36-6 Two females from
Ax-les-Thermes, Ariege, France, 36-6 and 38*4. Two males
from Silos, Burgos, Spain, 36 and 37. Female from the same
locality, 38. Five males from Ticino, Switzerland, 37 '0
(36-2-38). Mottaz collection: St. (denies, Gard, France (15,
152
CHIROPTERA
-5 '^
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154
CHIROPTERA
Loth sexes), 36 "4 to .SS-2) ; Sardinia (2 9 ), 37-4 to 37 -S. For
cranial measurements see Table, p. 153.
Specimens cxaminvd. — Eighty-one, from the following localities : —
Spain: Inca, Majorca, Balearic Islands. 1; San Cristobal, Minorca,
Balearic Islands, 3; Elche, Alicante, "1; Silos, Burgos, 5; Seville, 1.
Portugal: Ciutra, 1.
Fbance: As-les-Thermes, Ariegc, 6; St. Geuies, Gard, 17 (U.S.N.M.
and Mottaz); Marseilles, 1 (U.S.N.M.).
Switzerland : Locarno, Ticino, 4 (U.S.N.M.) ; Gordola, Ticino, 2
(U.S.N.M.) ; Minusio, Ticino, 6 (U.S.N.M.).
Italy : Liguria, :3 (U.S.N.M.) ; Siena, 3 (U.S.N.M.) ; Faenza, 1
(U.S.N.M.); Rome, 3; Osiia, Rome, 2; Ficuzza, Sicily, 6.
Sardinia : No exact locality, 2 (!Mottaz) ; Zimmigas, Siliqua, 2.
Corsica : Dintorni di Patrinionio, 3; no exact locality, 1.
^Ialta : 7.
?. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 1-2.
Islands. Pocock (c & p).
i, 9. San Cristobal. Minorca. O. Thomas and R. I. 0. 7. 1. 27-28.
Pocock (c iV' p).
1 al. San Cristobal, i\Iinorca. 0. Thomas and R. I. 0. 7. 1. G9.
Pocock (c & p).
?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 11.
c!. Elche, Alicante, Spain. G. S. Miller (c). S. 8. 4. 12.
5 al. Cintra, 500 m. Portugal. O. Thomas (c & p). 98. 2. 2. 57.
S,9. .4x-les-Thermes, Ariege, CV. S. Miller (c). 8.8.4.124-
2400 ft. France. 126.
6. Rome, 37 m. {C. Colt.) G.Barrett-Hamilton 11.1.2.42.
(P)-
2. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 6-6.
6 9. Ficuzza, Sicily. (A. J!ol>crt.) O. Thomas (p). |g- g" f l^'^^^'
6, ? al. Zimmigas, Siliqua, Sardinia. Marquis G. Doria (p). 6. 12. 1. 20-21.
(7?. Meloni.)
3 6, 2 ?. Malta. {J. Micalleff.) Lord LiKord (p). 11. 1. 1. 119-
123.
2. :\[aUa. Lord Lilford (p). 95. 3. 2. 1-2.
Rhinolopiius hipposidekos MixuTus Montagu.
1808. Vespcrtilio niiniitus Montagu, Trans. Linn. Soc. London, ix, p. 163.
1905. Rhinolophiis liipposidcrus minutus Andersen, Proc. Zool. Soc.
London, 1905, ii, p. 142, October 17, 1905.
1910. lilnnokphits liipposidcruf! ■mhnitus Trouessart, Fauna Mamm.
d'Europe, p. 10.
Type locality. — Wiltshire, England.
Geographical iJitftrihution. — England and Ireland.
Diagnosis. — Forearm, 36 '3 to 39 mm.; greatest length of
.skull about 16 mm.
Measuronents. — Forearm in 30 Engli.sh and Irish specimens
measured by Andersen {i.e. p. 142), 37 • 6 (36-3 to 39). For
cranial measurements see Table, p. 152.
Specimens examined. — Sixteen, from the following localities in England :
— Great Grimsby, Lincolnshire, 1 ; Bowdou, Cheshire, 1 ; St. Asaph,
Denbighshire, 3 ; Conway, Carnarvonshire, 1 ; Hope End, Herefordshire, 1 ;
Wells, Somersetshire, 1 ; Devizes, Wiltshire, 1 ; Zeals, Wiltshire, 2 ; Devon-
shire, 2 ; Ragley House, Warwickshire, 3.
RHINOLOPHUS
1 55
3 6. St. Asaph, Denbighsliire, Charles 0]dham (c .^- i'). 11.1.3.5-7.
Wales.
1. Conway, Carnarvonshire. Sir W. Jardine (c Ar p). GO. 9. 17. 1.
9. Great Grimsby, Lincoln- G. Barrett-Hamilton (p). U. 1. 2. 'J8.
shire, England. (Caton
Uaigh.)
6. Bowdon, Cheshire. {T.A. G. Barrett-Hamilton (i-). 11.1.2.99.
Coioard.)
6 al. Hope End, Herefordshire. N. C. Hewitt (c .t p).
6 al. Wells, Somerset. S. Lewis (c .fc p). 5. 1. 24. 1.
1. Devizes, Wiltshire. J. E. Hartins (c & p). 87.2.21.1.
i, 9 al. Zeals, Wiltshire. F. Norgate (c & p). 4. 11. 6. 1-2.
2 9. Devonshire. Oxley Grabham (c >t p). 11. 1. 3. 8-9.
3. Ragley House, Warwick- Tomes Collection. 7. 1. 1. 289-
shire. 291.
RHINOLOPHUS EURYALE Basius.
1853. PMnoloplms euryale Blasius, Wiegmann's Archiv fiir Naturgesch.,
1853, I, p. 49 (Milan, Italy).
1857. Bhinolophus euryale Blasius, Siiugethiere Deutschlands, p. 35.
1878. Bhinolophus euryale Dobson, Catal. Chiropt. Brit. IMus., p. 116.
1904. E{uryalus] toscanus Andersen and Matschie, Sitz.-Ber. Gesollsch.
Naturforsch. Preunde, Berlin, p. 77 (Caverna di Parigiiano, lit.
Pisani, Italy).
1904. E[uryalus] atlanticus Andersen and Matschie, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 77 (St. Paterne, ludre-et-Loire,
France) .
1904. E[nryalus] cabrene Andersen and Matschie, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 78 (Alcah'i de Henares, Madrid,
Spain).
1910. Rhinoloph-us euryale, R. euryale atlanticus, R. euryale toscanus,
R. euryale cabrerai Trouessart, Faune I\Iamm. d'Europe, pp. 5-7.
Type Jocalitij. — Milan, Italy.
Geixjraphkal distribution. — Southern Europe from Portugal
to Greece, north to Hungary and central France.
Diagnosis. — Size medium, forearm, 44 "6 to 49, condylobasal
length of skull, 16 '4 to 18, mandible, 12 to 1-3, upper tooth-row
6 • 2 to 6 • 6 ; noseleaf with parallel-sided, bluntly rounded sella
and high, sharply pointed connecting process, the lancet gradually
narrowing to a bluntly cuneate tip ; fourth finger with first
lihalanx slightly more than one-third as long as second (ratio
about 38) ; large upper premolar separated from canine by a
narrow space occupied by the much reduced small premolar.
External cliaractera. — Size intermediate between that of the
Greater and Lesser Horseshoes. General outline of no.seleaf
(fig. 26 c) about as in lihinolophus ferrum-equintoii ; sella parallel
sided, rounded ofi" above, connecting pi'ocess .sharply linear-
pointc^d, rising conspicuously above .sella ; lancet with slight con-
striction above middle, beyond wliich the tip narrows gradually to
a bluntl\- cuneate point. Ear when laid forward extending about
5 mm. beyond extremity of nuizzle, its tip less attenuate and le.ss
noticeably curved backward than in lihinolophii.'^ //■rrum-eqiiinuui
156
C'HIROPTEi;A
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,, slightly worn.
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RHINOLOPHUS
157
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CHIROrXERA
and I\. Jiipposidenis ; antitragal lobe about lialf as High as conch,
its width about equal to its height. Wing pecuUar in the
shortening of the tirst jshalanx of fourth finger to a little more
than one-third that of second (average of 1 specimens from Gard,
Fi'ance : tirst phalanx, 6'G mm.; second phalanx, 17 '4 mm.;
ratio of first to second, 38 -f )• -Foot averaging slightly more
than half as long as tibia.
Colour. — While essentially as in Illiinoloijlins ferrmii-eqmnuni
and M. hipposideros, the colour usually differs slightly in the more
evident contrast of the light area between ears and the more
drabby general effect of underparts. Median region below
occasionally rather paler than usual, sometimes nearly as in
R. mehehj).
Skull. — In both size (greatest length about 19 mm.) and form
the skull is somewhat intermediate between that of mdywlophun
ferrum-equinum and B. hipposideros. It resembles or surpasses
the latter in the breadth of brain-case relatively to naTro^\"
maxillar}' region, but more nearly agrees with the former in the
nearly parallel-sided mesopterygoid space, the anterior boi'der of
which is separated from posterior molars by well defined palatal
eraarginations. The mesopterygoid .space is, however, shorter in
proportion to its width than in B. ferrum-equinum. Floor of
brain-case between cochlese less narrowed than in the preceding
species. Nasal region slightly less inflated than in R. hipposideros,
and inflated area relatively shorter, its posterior border rising
abruptly above interorbital level.
Tectli. — In all respects the teeth closely resemble those of
Blii iiolophu8 ferrum-equinum (apart from their smaller size), excejjt
that the upper canine is relatively less robust, the anterior upper
premolar is less reduced (its crown area about double that of
uj:)per incisor), and anterior lower premolar is less crowded
between canine and posterior premolar.
Measurements. — For cranial and external measurements see
Tables, pp. 156 and 160.
Specimens e.vamined. — About 130, from the following localities : —
Portugal : Cintra, 6.
Spain: Villalba, Lugo, 1; ^Madrid, 2; Silos, Burgos, 1.
Feance : St. Paterue, Indre-et-Loire, 3 (B.M . and U.S.N.M.) ; St. Genies,
Gard, about 50 (Mottaz) ; Gapeau Kiver, Var, 12.
Italy: Near Genoa, 33 (B.M., U.S.N.M., Genoa, and Mottaz); Monte
Pisanino, 2; Siena, 3 (U.S.N.M.) ; Rome, 2 ; Velletri, Home, 5 (U.S.N.M.) ;
Nicotera, Calabria, 1 ; Marsala, Sicily, 2.
Sardiijia: Mount Gennargentu, 3 (U.S.N.]M.).
Austria-Hungary: Ofener Mountains, 2 ; Orsova, 1.
Dalmatia : Zara, 1.
Greece: Missolungi, Acarnania, 2 (U.S.X.M.).
JteiiiarJcs. — Mhinolopihus euryale is so readily distinguishable
from all the other European membsrs of the genus, except
R. mehehji, as to renuire no special comparisons. From R. meheli/i
it is most easilv distinguished bv tlie form of the lancet and
RHINOLOPHUS
159
aiititragus, together witli its rather smaller size and iisnall\-
darker colour. With the material at hand I am unable to
recognize the local forms of this species described by Andersen
and Matschie, as the alleged differences appear tf) be within the
range of normal indi^'idual variation.
2 9. Cintra, 500m. Portugal. (). Thomas (c & p). US. 2. 2. 2-8.
2 i, 2 9 al. Cintra, 500 m. 0. Thomas (c & r). >JS. 2. 2. 53-5G.
9 al. Villalba, Lugo, N.W. Dr. V. L. Seoane (p). 'J4. 1. 1. 1.
Spain.
5, 9 al. Madrid. A. Cabrera (p). 5. 2. 3. 1-2.
1 al. St. Paterne, Indre-et- Roj^al Army Medical "J. 1. 4. 9.
Loire, France. College (p).
9 al. St. Paterne, Indre-et- (t. E. Dobson (p). 80. 12. 14. 3.
Loire.
'J al. Cxapeau River, Var. Dr. K. Jordan (C & p). 8. 3. 15. 2-10;
3 i, 9 al. Finalborgo, L i g u r i a , Marquis G. Doria (p). 6. 12. 1. 14-17.
Italy. (A. Gagi'iu.)
2 al. Monte Pisanino,Lignria. Lord Lilford (p). 73. 1. 8. (i.
•2 6. Rome. {C. CoIi.) G. Barrett-Hamilton 11. 1. 2. 40-41.
W-
c$ al. Nicotera, Calabria. Florence Museum (b). 85.7.6.1.
2 d. :\Iarsala, Sicily. {A. O. Thomas (p). G. 8. 4. 10-11.
Robert.)
6, 9 al. Ofeuer Mts., Budapest. Budapest Museum (e). 94. 7. IS. 2-3.
9. Orsova, Hungary. Hon. W. Rothschild 7. 9. 16. 7.
9. Zara, Dalmatia, 50 ni. Lord Lilford (p). 11. 1. 1. 128.
{Kolombafocic.)
1 al. S. Europe. Purchased (Parreys). 47. 5. 27. 44.
RHINOLOPHUS MEHELYI Matschie.
1901. RhinolopliHS mehehji ^Matschie, Sitz.-Ber. Gesellsch. Naturforscli.
Freunde, Berlin, p. 225 (Bucharest, Roumauiat.
1904. Rhinolophufi carpetcuiua Cabrera, ]\Iem. Soc. Espafi. Hist. Nat., ii,
p. 254 (Madrid, Spain).
1910. Rhinolopliua euryalc vwlieliji and R. cavpctanus Trouessart, Fauue
5Iamm. d'Europc, pp. 7-8.
Type locaHty. — Bucharest, Uoumauia.
(Teix/rapJucal flistribidioii. — Roumania, southern France
(Uard), Sardinia, central Spain. Details of distribution not
known.
Diagnosis. — Like Hhiiiolophns luryaJr but larger (forearm,
48 'G to 51 "4; upper tooth-row about 7 mm.); noseleaf with
lancet abruptly narrowed to a linear tip ; ear with antitragal
lobe relativel)^ broad and low ; fourth linger with iirst phalan.\
decidedly more than one-third as long as second (ratio about 44) :
colour usually paler than in tlie related animal.
J E.rirrtial characters. —i>\ight\y larger and more robu.st than
Rhinoloplnis ciiryalc, a difference especially noticeable in freshlv
killed individuals. Noseleaf as in I{. ciiryale, except tluit the
lancet is very abruptly narrowed above middle to a distinctlv
linear tip. Ear as in B. enryulr but bioader. the antitragal lobe
160
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162 CHIROPTERA
scarcely half as high as conch, its width sHghtly greater than
height. Wing difl'ering from that of the related animal in the
less degree of shortening of the first phalanx of fourth linger as
compared with second (average of ten specimens from Gard,
France, first phalanx, 8'1 ; second phalanx, 18'0; ratio of first
to second, 44 +). Foot as in B. euryale.
Colour. — Though not invariably distinguishable the colour is
usually paler than that of Bhinolophus euryale, a difference
especially noticeable in the region between ears, on sides of face
and neck, on chin and throat, and along median portion of chest
and belly, all of which are frequently a very pale almost whitish
drab-gre}'.*
Skull and teetJi. — Except for its slightly greater average size
(greatest length about 20 mm.) the skull agrees with that of
Bhinolophus euryale. Teeth more robust than those of the
related animal, but not peculiar in form.
Measurements. — For external and cranial measurements see
Tables, pp. 161, 163.
Specimens examined. — About fifty-five, from the following localities : —
Spain: Near Madrid, 1 (paratype of carpetcums).
France ; Near St. Genies, Gard, about 50, skins and in flesh (!Mottaz).
Sardinia : Sassari, 3 (B.M. and U.S.N.M.).
RouMANiA ; Bucharest, 1 (U.S.N.M.) ; Dobrudscha, 1 (Mottaz).
Remarks. — At first sight this species apjjears very similar to
Bhinolophus euryale, together with which it occurs ; but its
characters when once understood are readily appreciable. The
ranges of the two animals will probably be found to be essentially
coincident, though Bhinoloplius mehelyl may prove to be more
strictly confined to the Mediterranean region than the smaller
form.
9. Madrid. A. Cabrera (p.) 6. 2. 3. 1.
(Paratype of R. carpetayius Cabrera.)
2 i al. Sassari, Sardinia. Marquis G. Doria (p). 6. 12. 1. 18-19.
RHINOLOPHUS BLASII Peters.
1857. Bhiiiolopihus cliuosiis Blasius, Saugethiere Deutschlands, p. 33. Not
of RiippeU, 1824 (Italy, Sicily, Istria and Dalmatia).
1866. Bhinoloplius blasii Peters, Monatsber. k. Akad. Wissensch. Berlin,
p. 17 (Renaming of clivosus Blasius).
1878. Rhinolophus blasii Dobson, Catal. Chiropt. Brit. Mus., p. 117.
1910. Bhiiwlophus blasiusi Trouessart, Faune Manim. d'Europe, p. 9.
Type locality. — South-eastern Europe.
Geographical distrihution. — Eastern portion of the Mediter-
ranean region : Cyprus, Greece, Italy?
Diagnosis. — Size essentially as in Bliinolojjlnis euryale; nose-
* When seen by candle-light flying in caverns these bats are said to
appear entirely white.
RHINOLOPHUS
163
Observations.
Teeth not worn.
„ slightly worn.
,, moderately worn.
,, not worn.
„ slightly worn.
,, not worn.
,, slightly worn.
„ not worn.
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• 6. 12. 1. 18
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86536
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„ 415
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37508
Locality.
R. mehelyi.
lloumauia: Dobrudscha .
Bucharest
Sardinia : Sassari .
France : near Nimes, Gard
Spain : near ^ladrid
R. blasii.
Cyprus ....
Greece: Nauplia
M 2
1G+ CHIROPTERA
leaf with cuneate sella and very bigli, sharply poijited connecting
process ; fourth fingei' with iirst phalanx more than half as long
as second ; no marked contrast between crown areas of anterior
and posterior lower* premolars, a character unique among the
European members of the genus.
Colour. — The only skin of this species which I have examined
is in bad condition. It indicates that the colour is not essentially
different from that of Bhinolophus euryale.
Skull. — In general the skull resembles that of RhlnolophiiH
euryale, with which it agrees in size and in the form of the nasal
swellings as well as in that of mesopterygoid fossa and posterior
portion of palate. Constriction at front of interparietal more
pronounced than in any of the other European species, noticeably
marking off the occipital region from rest of brain-case.
Teeth. — Incisors, canines and molars as in Mhinoloplms euryale.
Small upper premolar slightly less reduced, perfectly in the
tooth-row, but showing no tendency to develoj) a cusp. Large
upper premolar with anterior and posterior margins of crown
essentially parallel, the posterior border nearly straight. Lower
premolars differing from those of all the other European membei's
of the genus in the approximately equal crown ai'eas of the two
larger teeth, the anterior subterete, the posterior with trapezi-
form section ; shaft of posterior tooth with diameter in axis of
tootli-i'ow much less than transverse diametei', the cusp when
viewed from the side appearing to rise from middle of crown with
noticeable flat area before and behind it.
Meanurements. — Average and extremes of four females from
Cyprus: head and body, 48-4 (44-51); tail, 24-7 (24-25);
tibia, 19-3 (19-20) : foot, 9-7 (9-4-10) ; forearm, 45 -^ (44-6-
47) ; thumb, 7-5 (7-8) ; third tinger, 70 3 (69-72) ; tifth finger,
57 "7 (56—60) ; ear from meatus, 19 "7 (19-20) : ear fi-om crown,
15-5 (15-16-4); width of ear, 14-7 (14-15). Adult from
Nauplia, Greece : tibia, 18 - N ; foot, 9 • 6 : forearm, 44 • 6 ; thumb,
8 ; third finger, 09 : fifth finger, 57. For cranial measurements
see Table, p. 163.
Specimens exaviincd. — Five, from the following localities : —
Cypbus : No exact locality, 4.
Geeece : Nauplia, 1 (U.S.N.:^[.).
Meuiurls. — This .species is so readily distinguished from the
other European members of the genus by the peculiarities of its
noseleaf and lower premolar as to reejuire no special comparisons.
Its range appears to be strictly confined to the eastern portion
of the Meditei-ranean i-egion, not extending west of Italy.
vespertilionin.e 165
Family VITSPERTILIONID^.
1821. Vespertilionichx Oray, London IMecl. Repos., xv, p. 299, April 1, 1821
(part).
1857. Vespertiiiones Blasius, Siiugethiere Deutschlands, p. .37.
1878. Vcspertilionidx Dobson, Catal. Chiropt. Brit. ]\Ius., p. 1G7 (except
the genera Natalus and Thyroptera).
1907. Vespertilionidx Miller, Families and Genera of Bats, p. 195, -Tune 29,
1907.
Geographical distribution. — Eastern and western hemispheres
to the limits of tree growth ; in the Atlantic to the Azores, and
in the Pacitic to the Galapagos and Hawaiian Islands from
America, and to Australia, New Zealand and Samoa from Asia.
Characters. — Ear with tragus ; muzzle without distinct leaf-
like outgrowths ; skull with premaxillaries represented b}' nasal
branches only, the two bones very earl\' fused with surrounding
parts ; median length of palate greater than least distance between
tooth-rows ; auditory bulla not emarginated on inner side ; shoulder
girdle normal, without fusion of its elements : secondary
articulation of humerus with scapula better developed than in
the Rhiiiolvphidse ; fibula very slender, not adding appreciably to
strength of leg ; foot normal, the toes slender ; tail not project-
ing conspicuously beyond membrane.
Memarks. — This family is the most widely distributed group
of bats as well as one of the richest in genera and species. Forty-
one genera are at present known, eight of which occur in
Europe.*
Sub-Family VESPERTILIONIN^.
1878. Vcspertilioius Dobson, Catal. Chiropt. Brit. IMus., p. 168 (except
genera Kcrivoula and Harpiocephalus) .
1907. Vespertilioninx Miller, Families and Genera of Bats, p. 197, June 29,
1907.
Oeof/rapliicdl distrihniion. — Same as that of the family
Vespertilionidse.
Characters. — Sternum slender, its entire length considerably
more than twice greatest width of presternum ; median lobe
very much smaller than body of pre.sternum : six ribs connected
with sternum ; seventh cervical vertebra not fused with first
* The American Nycteris cinerea has been recorded (under the name
Vespertilio pruinosiis) from South Ronaldshay, Orkney Islands, but the
occurrence seems open to question (see Wolley, The Zoologist, vii, p. 2343,
1849; Yiii, pp. 2G95-96, 2813-14, 1850; Barrett-Hamilton, Hist. Brit.
Mamm., i, pp. 222-224, March, 1911).
A specimen of another North American member of this genus
(N. borealis), bearing the label: " Villeveque (Maine & Loire), 8. 1. 89," is
hgured by Trouessart in Bull. Soc. Zool. de France, xxx, p. 152, 1905.
This is copied, with change of locality from Maryland to France, from a
figure published in Bull. U.S. Nat. Mus., No. 39, pt. N, 1899 and 1901.
Though intended merely as a guide in preparing specimens, Dr. Trouessart's
publication might be misinterpreted as a French record of the species.
166 CHIROPTERA
dorsal ; scapula with coracoid curved outward ; nostrils simple ;
lower incisors in all known genera, 3-3.
BemarJcs. — The sub-family Vespertilioninse contains all but
eight of the known genera of VespertUionidse, and all but one,
Miniopferus, of the eight found in Europe. It is the central,
least specialized portion of the family.
KEY TO THE EUROPEAN GENERA OP VESPEBTILIONID^.
Cheek teeth ~. Mijotis, p. 166.
Cheek teeth less than Jp^.
Upper premolars 1-1,
Rostrum noticeably concave on each side of middle
line ; nares extending about halfway to inter-
orbital constriction ; palatal emargination
broader than deep Vcspcrtilio, p. 238.
Rostrum evenly convex laterally ; nares not ex-
tending halfway to interorbital constriction;
palatal emargination deeper than broad Eptesicus, p. 224.
Upper premolars 2-2.
Lower premolars 3-3.
Auditory bulla large, its greatest diameter
more than twice width of basioccipital ;
ear much longer than head ; second
phalanx of third finger shorter than first Plecotus, p. 256.
Auditory bulla small, its greatest diameter
about equal to width of basioccipital ; ear
shorter than head ; second phalanx of
third finger nearly three times as long as
first Miniopterus, p. 268.
Lower premolars 2-2.
Fifth finger shortened, its length only a little
more than that of metacarpal of fourth
or third Nyctalus, p. 242.
Fifth finger normal, its length greater than
that of metacarpal and first phalanx of
fourth or third.
Upper surface of rostrum convex ; ears
separate Pipistrcllns, p. 202.
Upper surface of rostrum concave ; ears
joined Barbastclla, p. 263.
Genus MYOTIS Kaup.
1829. Myotis Kaup, Entw.-Gesch. u. Natixrl. Syst. Europ. Thierwelt, i,
p. 106 (ynyotis).
1829. Nystactes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 108 (bcchsteinii).
1830. Lcuconoc Boie, Isis, p. 256 {datthcntonii) .
1839. Vespertilio Keyserling and Blasius, Wiegmaun's Archiv fiir Natur-
gesch., 1839, i, p. 306 (Not Vespertilio Linnfeus, 1758).
1841. Sclijsius Bonaparte, Iconogr. Faun. Ital. i., Introd. alia Classe
Mamm., p. 3 {m7jslaciniis).
1841. Capaccinius Bonaparte, Iconogr., Fauna Ital., i, Indice Distrib.,
p. 1 {capaccinii).
1842. Trilatitus Gray, Ann. and Mag. Nat. Hist., x, p. 258, December,
1842 (hasseltii, macellus = adversus and blcpotis).
MYOTIS 167
1849. Tralatitus Gervais, Diet. Univ. d'Hist. Nat., xiii, p. 213 (Modifica-
tion of Trilatitus).
1856. Brachijotus Koleuati, Allgem. deutsch. Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 {nujstacimis, daubentonii, and dasycnemc).
Not Brachijotus Gould, 1837.
1856. Isotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, neue
Folge, II, p. 131 {nattciTri and cmargiiiatiis).
1857. Vespertilio Blasius, Siiugethiere Deutschlands, p. 78. Not Vespertilio
Linnaeus, 1758.
1866. Tralatitius Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 90,
February 1866 (Modification of Trilatitus).
1867. PtcrnoiAerus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin,
p. 706 (sub-genus of Vespertilio = Mijotis, type lobipes = muricola).
1870. Exochurus Fitziuger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 75 (macrodactyius, horsfieldii =
adversus &nd macrotarsus).
1870. Aeoresies Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 427 (villosissimus, albescens, and
nigricans).
1870. Comastes Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.-
Naturwiss. Classe, LXII, p. 565 (capaccinii, incgapodius, dasycneme,
and limnophilus).
1878. Vespertilio Dobson, Catal. Chiropt. Brit. Mus., p. 284. Not Vespertilio
Linnaeus, 1758.
1897. Myotis Miller, Ann. aud Mag. Nat. Hist., 6th ser., xx, p. 382,
October, 1897.
1899. Euvespertilio Acloque, Faune de France, Mammif^res, p. 38 {emargin-
atus, mystacinus, nmrinus = myotis, nattereri, and bechsteinii) .
1907. Myotis Miller, Families aud Genera of Bats, p. 201, June 29, 1907.
Type species. — Vespertilio myotis Borkhausen.
Oeoiirapliical distribution. — Entire mainland of Eastern and
Western hemispheres to limits of tree growth ; also the Malay
Archipelago, New Guinea, Australia and Samoa, and in America
the Lesser Antilles.
Characters. — Dental formula : i ||, c J^, pm ^, m ;J-^ = 38.
General form slender and delicate, even in such large species as
M. myotis, the skull slender and lightly built ; muzzle narrow ;
ear narrow and rather long, without special peculiarities of form,
the tragus at least half as high as conch, straight or slightly
curved, tapering gradually to a narrow or acute point.
Bemarks. — The genus 3[!iotis is the most widely distributed
of the genera of bats. It is also probably the richest in species,
though these are at present so imperfectly known that no estimate
of their number can be made. Nine occur in Europe. These
present considerable differences in size, ranging from nearly the
smallest to nearly the largest members of the group ; they also
differ considerably among themselves in certain details of
structure, notably in the relative size of the hind foot ; but all
are recognizable, apart from their dental formula, by a certain
slenderness and delicacy of form, especially noticeable in the
muzzle, ear, tragus and skull.
168 CHIROPTERA
IvEY TO THE EUROPEAN SPECIES OF MYOTIS.
Size large (forearm 53 to 64 mm., condylobasal
length of skull 18-6 to 23-6 mm., upper tooth-
row 8'2 to 10"6 mm.) ; middle upper premolar
normally crowded inward from axis of tooth-
row ; third lower molar with second triangle
much smaller than first and noticeably different
from it in form.
Condylobasal length of skull 22 to 23 '6 mm.;
mandible 17 '8 to 19 mm. ; maxillary tooth-
row 9-8 to 10-6 mm. (Central and southern
Europe) M. mijotis, p. 192.
Condylobasal length of skull 18 '6 to 21-4: mm. ;
mandible 15 '2 to 17 "2 mm. ; maxillary tooth -
row 8-2 to 9'4 mm. (Mediterranean region) M. oxijgiuitlius, p. 199.
Size small or medium (forearm 34 to 47 mm., condy-
lobasal length of skull 12-4 to 16 '8 mm., upper
tooth-row 5 to 7 mm.) ; middle upper premolar
not crowded inward from axis of tooth-row ;
third lower molar with second triangle nearly
as large as first and essentially like it in
form.
Foot relatively large, obviously more than half
as long as tibia; calcar about twice as long
as free border of interfemoral membrane ;
skull broad, the width of brain-case more
than half greatest length ; crown area of
molars relatively small ; upper molars with
evident protoconule.
Forearm about 47 mm. ; condylobasal length
of skull about 16 mm. (Central and
southern Europe) M. dasijcnonc, ]^. 189.
Forearm less than 45 mm. ; condylobasal length
of skull never more than 15 mm.
Tibia and adjacent membrane densely furred ;
forearm about 42 mm. ; condylobasal
length of skull 14-0 to 14-8 mm. (Medi-
terranean region) 3/. ccqxiccinii, p. 187.
Tibia and adjacent membrane not furred ;
forearm about 35 mm. ; condvlobasal
length of skull 13-2 to 13 -"S mm.
(Distribution general) J\I. daubcntonii, p. 184.
Foot relatively small, about half as long as tibia ;
calcar about as long as free border of inter-
femoral membrane ; skull narrow, the width
of brain-case less than half greatest length ;
crown area of molars relatively large ; upper
molars without protoconule.
Ear not specially elongated, extending slightly
beyond nostril when laid forward.
Forearm about 40 mm. ; condylobasal length
of skull about 15 mm.; posterior border
of ear conch with deep, almost angular
emargination slightly above middle M. cmarginatus, p. 177.
. Forearm about 34 mm.; condylobasal length
of skull 12-6 to 13-2 mm.; posterior
border of ear conch with shallow, in-
conspicuous emargination M. mystacinus, p. 169.
Ear elongated, extending conspicuously beyond
nostril when laid forward.
MYOTIS 109
Combined length of tibia and foot less than
25 mm. ; condylobasal length of skull
14 • to 14 • 6 mm. ; ear narrow, its width
about 10 mm. ; tragus conspicuously
more than half as high as conch M. nattcrcri, p. 174.
Combined length of tibia and foot about
30 mm. ; condylobasal length of skull
16 to 17 mm. ; ear broad, its width about
14 mm. ; tragus scarcely half as high as
conch M. bechsteinii, ]^. 179.
MYOTIS MYSTACINUS Kuhl.
1819. Vcs^xrtiiio mystacinus Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 202.
1821. Vespertilio collaris Schinz, Das Thierreich von Cuvier, i, p. 177
(Mt. Blanc, Haute-Savoie, France).
1834. Tcs2iertilio humcmlis Baillon, M^m. Soc. Royale d'Emulatiou
d' Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1837. Vespertilio schinzii Brehm, Ornis, Heft iii, p. 27 (Renthendorf,
Thiiringen, Germany).
1843. V[espcrtilio] schrankii Wagner, Wiegmann's Archiv fiir Naturgesch.,
IX, Bd. II, p. 25 (Munich, Germany? See Fitzinger, Sitzungsber.
kais, Akad. Wissensoh. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 219, 1871).
1857. Vespertilio mystacinus Blasius, Saugethiere Deutschlands, p. 90.
1863. [Brachyottis mystacinus'] var. nigricans Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden,
Nassau, Germany).
1863. [Brachyotiis mystacinus'] var. rufofuscus Koch, Jahrb. dcs Vereins
fiir Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden,
Nassau, Germany).
18G3. [Bruchyotus mystacinus] var. aureus Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 445 (Breisgau,
Germany).
1869. [Vespertilio mystacinus] var. nigricans Fatio, Faune Vert. Suisse,
I, p. 92 (Switzerland). Not of Koch, 1863.
1869. IVespertilio] lugubris Fatio, Faune Vert. Suisse, i, p. 93 (Alternative
for nigricans Fatio).
1871. Vesjwrtilio mystacinus, nigro-fuscus Fitzinger, Sitzungsber. kais.
Akad. Wisseusch. Wien, jNIath.-Naturwiss. Classe, LXii, pt. 1,
p. 217 (Renaming of V. schinzii Brehm).
1878. Vespertilio mystacinus Dobsou, Catal. Chiropt. Brit. Mas., p. 314.
1900. Myotis mystacinus M6hely, Monogr. Chiropt. Hungarise, p. 200.
1910. Myotis mystacinus Trouessart, Faune Mamm. d'Europe, p. 33.
Geoffra^hical distribution. — Entire Continent of Europe north
to about the hmit.s of tree g!-owth ; west to Irehind ; ea.st into
Asia.
Diagnosis. — Smalle.st species of European Myotis (forearm
about 34 mm., longest finger about 60 mm., condylobasal length
of skull 12'() to 13 "2 mm.) : ear moderately long, extending 1 to
2 mm. beyond tip of muzzle when laid forward, its posterior
border with shallow incons})icuous emargination ; foot about half
as long as tibia ; wing membrane extending to ba.se of outer
toe ; last caudal vertebra free.
170
CHIROPTERA
External characters. — General form slender and delicate, the
legs and tail rather long, the membranes thin and semi-trans-
parent. Muzzle with rather noticeable glandular swellings.
Ear extending slightly beyond nostril when laid forward, its
general form rather slender, the tip narrowly rounded off, the
posterior border with shallow inconspicuous concavity extending
from just below tip to near middle of conch ; inner surface of
conch without well defined transverse striations. Antitragus
small (length about 2 mm.) but well defined Tragus a little
more than half as high as conch, its width slightly above level
of antei'ior base contained about 2^ times in length of anterior
border, the anterior border straight, the posterior border convex
below ; from widest region it narrows rather rapidly upward to
a rather blunt point ; posterior basal lobe well defined. Wing
rather narrow, with no special peculiarity of form, the third,
fourth and fifth metacarpals sub-equal, their distal extremities
falling short of elbow by about 3 mm.; membrane inserted at base
of outer toe. Foot about half as long as tibia ; calcar slender,
with barely indicated rudiment of keel and terminal lobe, its
length slightly greater than that of free border of interfemoral
membrane. Tail about as long as head and body, and twice as
long as tibia, its terminal vertebi'a free from membrane.
Fnr and colour. — The fur is soft and loose, the hairs on middle
of back about 10 mm. in length, tho.se of underparts a little more
than half as long ; it is closely confined to body, extending on
wings to extreme base of membrane only, slightly farther below
than above, and on interfemoral membrane over basal fifth or
fourth of both surfaces ; free border of uropatagium not fringed.
Colour of upper j^arts a clear light brown resembling the wood-
brown of Ridgway, but usually more yellow and always with a
distinct metallic gloss, the basal portion of the hairs blacki.sh-
slate, this colour sometimes appearing at the surface and
producing a general darkening effect ; underparts paler and more
buffy, usually not forming any decided contrast with back, but
occasionally almost whiti.sh, especially on
chest ; muzzle and cheeks dusky ; ears and
membrane blackish.
Skull. — The skull is slender and lightly
built, the breadth of brain-case conspicuously
greater than that of rostrum and slightly
but appreciably less than half greatest length.
Dorsal profile rising abruptly above low ros-
trum in interorbital region and forming a
strong convexity o%er anterior portion of
Fig. 29. brain-case ; occipital region distinctly pro-
MtiothmyHacinvs. ^^^^^^^ backward and rising slightly but evi-
dently above level of anterior portion of
brain-case, from which it is marked off by a shallow but notice-
al)le transverse constiiction foUowinjj anterior margin of inter-
171
parietal. Ventral profile rising slightly but e^sidently in region
of tloor of brain-case. Greatest depth of brain-case about three-
tjuarters mastoid width ; sagittal and lambdoid crests slightly
indicated in fully adult individuals. Interorbital region obscurely
short hour-glass shaped. Anteorbital foramen small, its posterior
border over posterior root of large premolar. Posterior palatal
region rather narrow, its width immediately behind molars about
equal to its greatest length, the median spine broad and short ;
mesopterygoid fossa slightly wider than long, the hamulars turned
inward. Auditory bulla moderate, its greatest diameter nearly
equal to distance between inner margins of bullje.
Teeth. — Upper incisors sub-equal, about half as high as canine,
each pair in contact or nearly so at base but diverging at tips, the
cingulum of inner tooth horizontal, that of outer tooth oblique ;
crown of outer incisor squarish in cross section, that of inner
somewhat elongated in axis of tooth-row ; near point of contact
each shaft bears a secondarj^ cusp, this usually though not always
better developed in outer than in inner tooth ; distance between
canine and outer incisor about equal to diameter of incisor, that
between pairs a little more than
twice as great. Lower incisors
slightly but evidently imbricated,
forming a continuous, broadly
V-shaped row between
canines ; crown of i^ sub-
terete, its area more than
half that of canine and
about equal to that oi pm^,
its blunt main cusp, the
highest in the incisor series,
situated at outer side, the
three smaller cusps (third
obsolete) on inner mai'gin ;
^1 and /., sub-equal, slightl}-
lower than i'g, their crowns
compressed, longer than
high, tritid, that of second
wider posteriorly than an-
teriorly, and usually with
a minute postero-internal
cusp. Upper canine rela-
tively large, slightly higher than main cusp of large premolar,
its shaft somewhat triangular in cross section, with broad
postero-internal concavity and anterior and postero-external
groove ; posterior cutting edge well developed ; cingulum com-
plete but not forming evident secondary cusps. Lower canine
lower and less acutely pointed than upper, its tip about on level
with highest cusps of molars ; cingulum usually forming a slight
secondary cusp anteriorly. First and second upper premolars
Flo. »l.
My Otis mystaciiius. Teeth X 10.
172 CHIROPTERA
alike in foim, crowded between canine and large premolar but
perfectly in the tooth-row, the first about the same size as the
ujjper incisors, the second with slightly more than half the
height and crown area of first, both with fully developed
cingulum and subterete, conical cusp. Large upper premolar
with crown area only a little less than that of first molar, the
main cusp large and with strongly trenchant posterior cutting-
edge, its height about eijual to that of largest molar cusps :
anterior and ^^osterior borders of crown slightly concave, inner
border narrowly rounded, usually with a distinct though small
cusp anteriorly, sometimes with another barely indicated pos-
teriorly ; a shght though evident concave crushing surface
between cingulum and inner base of main cusp. First and
second lower premolars essentially similar to the corresponding
upper teeth but slightly less reduced in size and with somewhat
higher cusps ; third lower premolar similar to first and second
Init with rectangular crown nearly twice as large as that of second
tooth, and main cusp as high as protoconid of first molar ; a
small cingulum cusp usually present at antero-internal angle.
Upper molars rather large relatively to size of skull, the crown
area of second tooth slightly greater than that of first ; anterior
and posterior borders slightly concave, inner border narrowly
rounded, especially in m'^, the antero-internal and postero-internal
outlines often flattened or even a little concave ; protocone large,
its base occupying entire inner border of tooth, its cusp a little
in front of middle, its anterior commissure simple, extending
uninterrujitedl}' outward to parastyle, its posterior commissure
terminating in a thickened rib-like rudiment of a hypocone,
between the outer base of which and inner base of metacone lies
a deep furrow : metacone larger than paracone ; styles and outer
commissures well developed ; third upper molar with crown area
equal to about three-quarters that of iu^, the protocone as in the
other teeth liut smaller, paracone larger than metacone, the first
outer commissure longer than in vi'^ and m'^, the second and third
about as in the other teeth but set at a different angle, the fourth,
together with metastyle absent. First and second lower molars
with second y slightly larger than first in cross section, this
condition reversed in m^ ; protoconid decidedly higher than
hypoconid in all three teeth ; inner cusps of about equal height
throughout (slighth? more than half as high as protoconid) ; a
distinct cingulum cusji behind entoconid.
Measurements. — Adidt female from Skane, Sweden : tibia,
15 ; foot, 7 • 6 ; forearm, 32 ; thumb, 6 • 2 ; third finger, 49 ; fifth
finger, 38 ; ear from meatus, 1 2 ; width of ear, 8. Adult male from
Madrano, Tyrol: head and body, 38; tail, 38; tibia, 16; foot,
8 ; forearm, 35 : thumb, 6-2 ; third finger, 56 ; fifth finger, 47 ;
ear from meatus, 13 : width of ear, 9. Adult female from the
Carpathian Mountains : head and body, 44 ; tail, 40 ; tibia, 15 • 4 ;
foot, 8 ; forearm, 34 ; thumb, 7 ; third finger, 54 ; fifth finger.
MYOTIS 173
45 ; ear from meatus, 14 ■ 2 ; width of ear, 9. Forearm, in other
specimens : Waremme, Liege, Belgium, 33 and 33 ; tStrass, near
Burgheim, Ba^■aria, 31 '6
Switzerland, 33 and 34 '4.
p. 182.
Dresden, 33 "2 ; Grotte de Vallorbe,
For cranial measurements see Table,
Specimens examined. — Thirty-five, from the following localities ; —
England : Newby Bridge, Lake Windermere, Cumberland, 1 ; Aberia,
Merionethshire, 2; Colwyn, Denbighshire, 1; Cheadle, Staffordshire, 1;
Manchester, Lancashire, 1 ; Ragley House, 1 ; Welford, Herefordshire, 2 ;
Macclesfield, Cheshire, 1 ; Pewsey, Wiltshire, 1 ; Dover, Kent, 2 ; Hastings,
Sussex, 1 ; Bath, Somerset, 1.
Sweden: Skullno, 1; Skane, 1 (U.S.N.M.).
Belgium: Waremme, Liege, 2 (U.S.N.M.).
Germany: Moritzburg, Saxony, 1 (U.S.N.M.) ; Dresden, 1; Strass,
near Burgheim, Bavaria, 1 ; Bavaria, no exact locality, 2 (U.S.N.M.).
ArsTRiA-HuNGARY : Haida, Arva, Bohemia, 1 ; Carpathian Mts., 1
(U.S.N.M.) ; Csallokoz-Somorja, Pressburg, 2 ; Tatra Mts., Hungary, 1.
Switzerland : Geneva, 1 (Mottaz) ; Grotte de Vallorbe, Vaud, 2
(Mottaz); Stein, Appenzell, 1 (U.S.N.M.); St. Gallon, 1 (U.S.N.M.); no
exact locality, 1 (Geneva: type of luguhris Fatio).
Remarks. — Mijotis mystacinus, the smallest European member
of the genus, is recognizable by its small size and relatively short
foot in combination with the insertion of wing membrane at base
of outer toie. From the small species of Pipistrellus it is at once
distinguished by the narrow muzzle and slender tragus.
2 9.
Aberia, Merionethshire,
G. H. Caton Haigh
11. 1. 3. 13-14
Wales.
(c & P).
9 St.
Colwyn, Denbighshire.
Hon. N. C. Roths-
child (p).
6. 2. i. 1.
1 al.
Newby Bridge, Cumber-
land, England.
T. Paul (c & p).
94. 9. 3. 1.
9.
Cheadle, Staffordshire.
E. Blagg (c & p).
11. 1. 3. 15.
6.
Manchester, Lancashire.
C. Oldham (p).
11. 1. 3. 10.
6.
Ragley House, Warwick-
shire.
Tomes Collection.
7. 1. 1. 498.
2 9.
Welford, Herefordshire.
Tomes Collection.
7.1.1.496-497
6.
Pewsev, Wiltshire.
(P. S.'Hembly.)
C. H. B. Grant (p).
11. 1. 3. 11.
9.
Dover, Kent. {B. Hesse.)
C. H. B. Grant (p).
11. 1. 3. 12.
1.
Hastings, Sussex.
MissI.Roods(c&p).
49. 1. 16. 1.
6 St.
Bath, Somerset.
G. Dalgleish (c & p).
4. 10. 13. 3.
1.
Skullno, Sweden.
Stockholm ^luseum
46. 1. 2. 22.
9.
Haida, Bohemia. (Ti"u/-
terstorff.)
Lord Lilford (p).
11. 1. 1.6.
1.
Csallokoz - Somorja,
Budapest Museum
94. 3. 1. 20-21
Pressburg, Hungary.
(E).
lal.
Tatra Mountains,
Dr. R. CoUelt (p).
91. 1. 21. 1.
Hungary.
174 CHIEOPTEEA
MYOTIS NATTERERI Kuhl.
1818. Vespertilio nattereri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 33.
1857. Vespertilio nattereri Blasius, Siiugethiere Deutsclilands, p. 88.
1863. [Isotus nattereri'] var. typhis Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 430 (Wiesbaden, Nassau,
Germany).
1863. [Isotus nattereri] var. spelxus Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 430 (Erdbach, Nassau,
Germany) .
1878. Vespertilio nattereri Dobson, Catal. Chiropt. Brit. Mus., p. 88.
1900. Myotis nattereri M6hely, Monogr. Chiropt. Hungarise, p. 179.
1904. Myotis escalerai Cabrera, Mem. Soc. Espau. Hist. Nat., ii, p. 279
(Bellver, Lerida, Spain).
1910. Myotis nattereri and M. escalerai Trouessart, Faune ilamm. d'Europe,
pp. 29-30.
Type locality. — Hanau, Hessen, Germany.
Geographical distribution. — Central and southern Europe,
west to Ireland, north to southern Sweden.
Diaijnosis. — Size medium among the European species (forearm
about 38 mm., longest linger about 70 mm., condylobasal length
of skull, 14 "0 to 14 '6 mm.); ear elongated, extending about
5 mm. beyond tip of muzzle when laid forward, the conch narrow
(about 10 mm.), obscurely emarginate on upper half of posterior
border, the tragus relatively longer than in any other European
species, its height distinctly more than half that of conch ; foot
about half as long as tibia ; wing membrane extending to base of
outer toe ; edge of interfemoral membrane fringed.
External characters. — General form essentially as in Myotis
mystacinus, but differing in the following particulars : ear much
longer and relatively narrower, extending conspicuously beyond
nostril when laid forward, its extremity more broadly rounded
off (owing to more uniform convexity of anterior border), and
concavity on posterior border even less evident ; antitragus
about as large as in 31. mystacinus but less well defined ; tragus
relatively longer and more attenuate than in any other European
bat, its height conspicuously more than half that of conch, its
gi'eatest width contained about 3^ times in length of anterior
Iwrder, its terminal third almost linear, sometimes faintly
recurved, the basal lobe usually small and ill defined ; insertion
of wing membrane as in 31. mystacinus, but membrane at first
very narrow, so that in some specimens, particularly those that
have been hardened in strong alcohol, the point of insertion
appears to be on side of metatarsus ; * tail rather shorter than
head and body, only the cartilaginous extreme tip free ; calcar
l)etter defined than in 31. mystacinus, about as long as the
distinctly fringed free border of interfemoral membrane.
* Such specimens seem to have formed the basis of the Myotis escalerai
of Cabrera.
MYOTIS 175
Fur and colour. — Quality and distribution of fur essentially as
in M. mydacinus, but free border of interfemoral membrane
distinctly though not very densely fringed with hairs about
1 mm. long. Colour of upper parts a lighter and less yellowish
brown than in M. mystacinua, the exact shade intermediate
between the wood-brown and broccoli-brown of Ridgway, the
longer hairs with faintly darker tips visible in certain lights ;
underparts rather sharply contrasted whitish buffy grey ; a well
defined line of demarcation extending from shoulder to base of
ear ; basal portion of hairs clove-brown ; muzzle and cheeks dusky,
but not so dark as in M. mi/stncinus ■ ears and membranes dark
brown.
Shill. — In all its dimensions the skull is appreciably larger
than that of Myotis mysiacinus. In general form it is slightly
less slender. Forehead rising moi'e abruptly but occipital region
not higher than main portion of brain-case, so that dorsal profile
shows a stronger concavity in interorbital region than that of
M. mystacinus, while over greater extent of brain-case it is
essentially flat, or slightly falling away posteriorly, instead of
rising by two well defined curves to lambdal region. Posterior
palatal region about as in M. mystacinus, except that mesopterygoid
space extends further forward, and median spine is better
developed. Auditory bulla relatively smaller than in M. mysta-
cinus, its greatest diameter decidedly less than distance between
bullae.
Teeth. — In general the teeth resemble those of Myotis
mystacinus apart from their greater size. They differ, however,
in certain details of form : crown area of outer upper incisor
appreciably greater than that of inner tooth ; angle at front of
lower incisor row wider, though evident ; upper canine relatively
weaker, its posterior cutting edge less developed, the cross section
of its shaft half-terete owing to the obsolescence of postero-
external and anterior longitudinal furrow ; lower canine with
length of base much greater in proportion to height of shaft, and
cingulum more oblique ; second upper premolar nearly equal to
first in cross section ; crown area of large premolar not so great
relatively to that of first molar, the cusps on inner border
obsolete or absent ; lower premolars with less slender crowns,
the cingulum of the third forming a very low, sometimes obsolete
antero-internal cusp ; molars, especially m^, with narrower
crowns.
Measitrenwnts. — External measurements of adult male from
Colpin, Brandenburg, Germany, and adult male from Magdeburg,
Germany: head and body, 50, 44; tail, 41, 40; tibia, 17, 16 ;
foot, 8-2, 9-0; forearm,"' 39-2, 40-2 ; thumb, 7-0, 6-2; third
finger, 71,71 ; fifth finger, 56, 56 ; ear from meatus, 18, 18 ; tragus,
11*0, 11 • 2. Two adult females from Spain (Xo. 94. 1 . 1.8, Se-\-ille,
and No. 8. 7. 23. 4 from Deliver, Lerida, paratype of Jlf. escalerai) :
head and body, 46, 46 ; tail, 4 1 , 43 ; tibia, 1 5, 1 7 ; foot, 7 • 6, 9*2;
176
CHIROPTERA
forearm, 36 -4, 40" 6 ; thumb, 7 -2, <S'4 ; third finger, 69, 72 ;
fifth finger, 52, n6 ; ear from meatus, 16, 16" 6; tragus, 10*2,
10±. For cranial measurements see Table, pp. 182-183.
Specimens examined. — Sixty-seven, from the following localities: —
Scotland : Inverary, Argyllshire, 1.
England: Harlech, Merionethshire, .3; Arrow Church, G; Alcester,
Warwickshire, 8; Lilford Hall, Northamptonshire, 5; Henley-on-Thames,
Oxfordshire, 3 (B.M. and U.S.N.M.) ; Bradfield, Berkshire, 1 ; Queen
Camel, Somersetshire, 1 ; Devonshire, 1 ; no exact locality, 3.
Ireland: Co. Longford, 1; Woodpark, Co. Galway, 1.
Germany: Oberlausitz, Silesia, 1 ; Colpin, Brandenburg, 1 ; Magdeburg,
Saxony, 2; Moritzburg, Saxony, 3 (U.S.N.M.).
Austria-Hungary: Haida, Arva, Bohemia, 1.
Switzerland: Canton Thurgau, 3 (U.S.N.M.); St. Gallen, 9 (B.-M.
and U.S.N.M.).
Italy: Arezzo, 4 (B.M. and U.S.N.M.) ; Valesia, 1 (U.S.N.M.) ; Siena, 1
(Mottaz) ; no exact locality, 4 (U.S.N.M.).
Spain: Bellver, L^rida, 2 (B.M. and Genoa; paratypes of escalerai
Cabrera) ; Seville, 1.
lii'hKi.rJis. — This species is easily recognized by its rather small
size, large ear, and very long, attenuate tragus. The exact point
of insertion of wing membrane on side of foot is probably in all
specimens the base of outer toe, but owing to a peculiar narrow-
ing of tlie membrane along edge of metatarsal, the point of
insertion sometimes appears to be at middle of side of foot. This
effect is often increased by the action of strong alcohol.
9. Inverary, Argyllshire,
Scotland.
1 al. Longford, Ireland.
9. Woodpark, Galway.
9, 9 juv. Harlech, Merioneth-
shire, Wales.
G. Arrow Church, Warwick-
shire, England.
Alcester, Warwickshire.
Lilford Hall, Northamp-
tonshire.
Bradfield, Berkshire.
Henley, Oxfordshire.
Henley, Oxfordshire.
Queeji Camel, Somerset.
England.
9. O be rlaus it z, Silesia,
340 m. Germany.
(W. Baer.)
6 al. Colpin, Brandenburg,
Prussia.
6 al. 3Iagdeburg, Saxony.
Duke of Argyll (p)
58. 8. IG. 1.
Dr. G. E. Dobson 76. 11. 3. 2.
(c & P).
R. F. Hibbert (p). 11. 1. 3. 20.
J. Backhouse (p). 11. 1. 3. 16-18.
8 al.
3 9 al
9.
9 St.
6 St.
(Jal.
1 al.
Tomes Collection.
Tomes Collection.
Lord Lilford (c & p).
N. H. Joy (c & p).
J. G. Millais (c & p).
Heatley Noble (c & p).
R. H. Read (p).
Dr. J. E. tTray (p).
Lord Lilford (p).
7.1.1.488-493.
7.1.1.736-743.
72. 8. 21. 3-.5.
72. 11. 12. 13.
11. 1. 3. 19.
I. 11. 2. 1.
0. 3. 23. 1.
II. 1. 3. 21.
•51. 1. 29. 12.
99. 1. 9. G.
Dr. H. Gadow (c & p). 82. 7. 31. 2.
Dr. W. Wolterstorff 92. 12. 1. 1.
3 S, 9.
jMagdeburg, Saxony.
{Wolterstorff.)
Haida, Bohemia. (ll'oZ-
terstorff.)
St. C^allen, GOO m. Swit-
zerland. {E. H. Zolli-
kofer.)
Lord Lilford (p).
Lord Lilford (p).
0. Thomas (p).
11. 1. 1. 8-9.
11. 1. 1.
4. 4. 5. 4-8.
MYOTIS 177
2 al. Arezzo, Italy. Florence Museum (e). 85. 7. 6. 4-5.
9 al. Bellver, Lc^Tida, Spain. A. Cabrera (p). 8. 7. 23. 4.
(Paratype of M. escalcrai, Cabr.)
1 al. Seville. Dr. V. L. Seoane 94. 1. 1. 8.
(c & p).
MYOTIS EMARGINATUS Geoffroy.
1806. Vesp{ertilio] emarginatus Geofiroy, Ann. Mus. d'Hist. Nat., Paris,
VIII, p. 198 (Charlemont, Givet, Ardennes, Prance).
1844. Vesp\crtilio'] rufescens Crespon, Paune Meridionale, i, p. 20 (near
Nimes, Gard, Prance). Type in Nimes Museum.
1853. V[espertilio] ciliatus Blasius, Wiegmann's Archiv fiir Naturgesch.,
1853, I, p. 287 (near Cologne, Germany).
1856. V[es2}ertilio] schrankii Kolenati, Allgem. deutsche Naturhist. Zeitung,
Dresden, neue Polge, ii, p. 178. A nomen nudum (ex Koch and
Giebcl) cited as sj'nouym of emarginatus. Not of Wagner, 1843.
1858. Vespertilio ciliatus Blasius, Siiugethiere Deutschlands, p. 91.
1878. Vespertilio emarginatus Dobson, Catal. Chiropt. Brit. Mus., p. 303.
1880. M)/otis ciliata var. buclapcsticnsis Margo, " INIagyar orv. es term^s-
zetvizsg. xx,nagygyul. munk, p. 255" (Budapest, Austria-Hungary).
See M6hely, Monogr. Chiropt. Hungarife, p. 170.
1890. Vespertilio neglectus Patio, Arch. Sci. Phys. et Nat., Geneve, 3rd ser.,
XXIV, p. 512, November 15, 1890 (Valavran, near Geneva, Swit-
zerland). Type in Geneva Museum.
1900. Mijotis emarginatus Mehely, Monogr. Chiropt. Hungarise, p. 170.
1910. Mrjotis e^narginatus Trouessart, Paune Mamm. d'Europe, p. 28.
Tj/jje locality. — Charlemont, Givet, Ardennes, France.
Geographical distribution. — Central and southern Continental
Europe.
Diagnosis. — Size essentially as in 3Ti/otis naitereri (forearm
about 40 mm., condylobasal length of skull about 15 mm.) ; ear
moderately long, extending 2 to 3 mm. beyond tip of muzzle
when laid forward, its posterior border with a deep almost
angular emargination slightly above middle ; foot about half as
long as tibia ; wing membrane extending to base of outer toe : free
margin of interfemoral membrane sometimes fringed ; fur some-
what woolly in texture, the hairs of back tricolored, drab at
base, yellowish at middle and dark brown at extreme tip.
External characters. — AVhole animal larger and more robust
than Mijotis miistacinus, though of essentially the same form and
proportions. Ears and membranes relatively thick and leathery,
not semi-transparent as in the related small species. Ear
moderately long, extending about 2 mm. beyond nostril
when laid forward, its size and general form much as in M.
mystacinus except that emargination of posterior border is deep
and conspicuous, in most specimens forming an evident angle
below. Inner siu'face of conch marked bv seven or eight short
but well developed cross ridges. Antitragus small and ill-defined.
Tragus slightly more than half as high as conch, its greatest
width contained nearly three times in length of' anterior border,
both its margins essentially straight from level of anterior base
N
178 CHIROPTERA
to rather acute tip. Wing as in the related species ; metacarpals
falhng short of elbow by about 3 mm. ; membrane inserted at
base of outer toe. Calcar slender, its termination usually marked
by a distinct lobe, its length about equal to that of free border
of interfemoral membrane. Tail relatively shorter than in 31.
mystacinus, extending to between ears when laid forward, onh'
the minute cartilaginous tip free from membrane.
Fur and colour. — Fur shorter and more dense than in M.
mystacinus and M. naUereri, and of a slightly woolly texture
unique among the European species, the hairs at middle of back
about S mm. in length. Distribution of fur not peculiar ; free
border of uropatagium sometimes with evident fringe. General
colour buff, light and clear on underparts, dulled and irregularly
clouded by darker brownish hair tips throughout upper parts ;
basal half of hairs drab ; muzzle and cheeks dusky ; ears and
membranes an indefinite rather light brown.
SJiulI. — In form the skull does not differ appreciably from
that of M. naUereri, except that the rostral portion and palate are
relatively more elongate and interorliital concavity is less strongly
pronounced ; occipital region similarly low as compared with that
of M. nnjsfacinns. Mandible slightly more robust than in 31.
naUereri, but of similar form.
Teeth. — The teeth are larger than those of 3Iiiot!s mystacinus,
in this respect agreeing with those of M. natteieri. Upper
incisors higher and more slender than in the preceding species,
their crowns sub-equal in cross section ; lower incisors very
slightly imbricated, forming a broadly and e^•enly convex
(U-shaped) row without anterior angle, the crowns of (j and i.,
alike in form and distinctly 4-cusped, that of i^ not thickened
posteriorly and with no trace of postero-internal cusp. Canines
about etiual to those of 31. nattereri in size, but shaft of upper
tooth with evident postero-external longitudinal groove. First
and second upper premolars even more strongly contrasted in
size than those of 31. mystacinus, the cusp of second only a little
exceeding cingulum of first ; the two teeth less crowded between
canine and large premolar than in 31. mystacinus and M. nattereri ;
lower premolars essentially as in 3T. nattereri, but less closely
crowded, and second relatively larger. Upper molars as in
M. nattereri, but crowns less narrowed ; a rudimentary commissure
extending outward from base of hypocone and another extending
inward from base of paracone ; lower molars not peculiar.
3'leasurements. — Two adult females from Florence, Italy :
head and body, 46-6 and 50 ; tail, 40 and 4-2 ; tibia, 19 and 19 ;
foot, 8-4 and 8-6; forearm, 40 and 41 ; thumb, 8 and 7-8;
third finger, 67 and 70 ; fifth finger, 56 and 57 ; ear from meatus,
16-6 and 17 ; width of ear, 11-4 and 12. For cranial measure-
ments see Table, p. 183.
Specimens examined. — Six, from the following localities : —
Holland : Maastricht, 1 (U.S.N.M.).
MYOTIS 179
Prance : Near Nimes, Gard, 1 (Nimes : type of rufcscens Crespon).
AusTBiA-HuNGARY : Herkulesbad, 2.
Switzerland: Valavran, near Geneva, 1 (Geneva: type of nenlectus
Fatio). -^
Italy: Florence, 2 (U.S.N.M.) ; no exact locality, 1.
Remarks. — The peculiar form of the ear, the short, somewhat
woolly fur, and the yellowish colour are highly characteristic of
this well-detined species.
2 9. Herkulesbad, Hungary. Hon. N. C. Roths- 7. 9. IG. 9-10
child (p).
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 73:3.
without parte.)
skull.
MYOTIS BECHSTEINII Kuhl.
1818. Vespertilio bechsteinii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 30 (Hanau, Hessen, Germany).
1857. Vespertilio bechsteinii Blasius, Siiugethiere Deutschlands, p. 85.
1878. Vespertilio bechsteinii Dobson, Catal. Chiropt. Brit. Mus., p. 308.
1900. Myotis beclistcinii M6hely, Monogr. Chiropt. HungariiB, p. 184.
1905. Vesplertiliol bechst{einii\ ghidinii vel Vcs2}[ertilio'] ghidinii Fatio,
Arch. Sci. Phys. et Nat., Geneve, 4th ser., xix, p. 511, May 15,
1905 (Lugano, Ticino, Switzerland). Type in Geneva Museum.
1906. Myotis beclistcinii favonicus Thomas, Ann. and Mag. Nat. Hist.,
7th ser., xviii, p. 220, September, 1906 (La Granja, Segovia,
Spain). Type in British Museum.
1910. Myotis bechsteini and M. bechstcini favonicus Trouessart, Fauue
Mamm. d'Europe, pp. 30-81.
Ti/pe localiti/. — Hanau, Hessen-Nassau, Germany.
Geographical distrihution. — Central and southern Europe,
west to England, north to southern Sweden.
Biagnods. — Size slightly greater than that of M. nattererl
(forearm about 40, condylobasal length of skull, 16 to 17) ; ear
elongated, extending about 8 mm. beyond tip of muzzle when laid
forward, the conch broad (about 15 mm.), its posterior margin
obscurely emarginate above, the tragus scarcely half as high
as conch ; foot about half as long as tibia ; wing membrane
extending to base of outer toe.
External characters. — In general like Myotis nattereri, but with
broader ears and relatively as well as actually larger legs and
feet (combined length of tibia and foot about oO mm. instead of
less than '25). Muzzle with moderately developed glandular
swellings (these less evident than in M. mystacinus). Ear extend-
ing considerablj' beyond nostril when laid forward, its length thus
about as in M. nattereri, but its breadth so much greater (about
17 mm. instead of about 10 ram.) that the ear is relatively larger
than in any other European bat except Plecotus auritns ; form
of ear essentially as in M. mystacinus, the upper half of posterior
border faintly concave ; inner surface of conch with about eight
rather ill-delined transverse ridges near posterior border ; anti-
tragus about '1 ' 5 mm. in length along base, rather abruptly
N 2
180 CHIROPTERA
rounded off above, its inner margin not continuous with posterior
border of conch. Tragus about half as high as conch, its form
essentially as in M. mystacinus though a little more slender
(greatest width contained nearly three times in height of
anterior border) ; basal lobe small but well developed. Wing
essentially as in M. mystacinus ; third, fourth and tifth meta-
carpals sub-equal, falling short of elbow by about 5 mm. ;
membrane inserted at base of outer toe. Calcar and free border
of interfemoral nieml^rane as in M. mystacinus. Tail shorter
than head and body (laid forward it extends to middle of crown),
the terminal vertebra free.
Fm- and colour. — Quality and distributicjn of fur as in Myotis
iity.^tacinas and M. nattereri. Upper jaarts uniform wood-brown
with a slight tinge of umber, the exact shade almost impossible
to describe exactly ; underparts a huffy grey in strong contrast,
irregularly clouded by the slaty brown of underfur ; ears and
membranes an indefinite dark brown.
Skull. — In general aspect the skull resembles that of ilf//o//.s'
myotis, due allowance being made for its much smaller size, since
it is much more slender than in any of the other small species ;
breadth of brain-case equal to about one-half distance from
lambda to posterior border of narial emargination. Dorsal
profile rising gradually above rostrum and forming a strong
convexity over anterior jiortion of brain-case, behind which it is
essentially flat to lambda ; ventral profile nearly flat, slightly
elevated posteriorly ; occipital region slightly projecting, just
sufficiently to conceal condyles when viewed from above. Brain-
case ovate, noticeably wider than rostrum ; sagittal crest low but
evident ; lambdoid crest moderately develo2:>ed at sides, obsolete
at middle ; greatest deptli of brain-case about two-thirds mastoid
breadth ; floor of brain-case with obscurely defined lateral grooves.
Interorbital region not evidently hour-glass shaped, owing to the
slight widening at lachrymal level ; anterior upper border of
orbit slightly ridged. Rostrum slender, widening a little at
anterior extremity, the dorsal surface smoothly rounded off at
sides, though with indication of a slightly developed median
groove ; narial emargination narrow but deep, extending back
about half way to level of anterior rim of orbit ; rostral depth
at front of orbit about equal to distance from orbit to middle of
canine ; palate long and narrow as compared with that of the
other small species, both transverse and lateral concavities evident,
especially just behind middle ; anterior emargination rather large,
sub-circular in general outline, extending back about to level of
space between canine and first premolar ; posterior extension of
palate about as broad as long ; interpterygoid space wider than
long, encroached on by blunt median palatal spine. Mandible
essentially as in Myotis myotis, the coronoid process higher and
with more obli(iue posterior border than in the other small species.
Teeth. — The teeth are rather small relatively to size of skull.
MYOTIS 181
Upper incisors essentially as in 31. mijutaciniis but more slender.
Lower incisors slightly imbricated, the row as a whole (J -shaped
or very broadly V-shaped, the form of the individual teeth not
peculiar. Canines with no special peculiarities. Small upper
premolars completely in tooth-row, not crowded, the crown area of
first eijual to about half that of canine, that of second to about
two-thirds that of first, the height of first a little less than half
that of canine, that of second about two-thirds that of first ;
cingulum complete but not forming secondary cusps. Large
upper premolar with crown area nearly three-(|uarters that of
first molar, its antero-internal cusp well developed. Lower
premolars essentially as in 31. nattereri. Upper molars with no
special peculiarities except that wi^ is more reduced than in the
other small species, its tranverse diameter through metacone
noticeably less than half length of anterior border. Lower
molars normal, but second triangle of m^ a little more reduced
than in the other small species, though much less so than in
3/1. myotis and M. oxygnatlius.
Measurements. — Adult female from Sweden : head and body,
46 ; tail, 38 ; tibia, 19 • 6 ; foot, 9 • 8 ; forearm, 42 ; thumb, 9 • 6 ;
third finger, 67 ; fifth finger, 56 ; ear from meatus, 22 ; ear
from crown, 19; width of ear, 13-6; tragus, 10. Two adults
from the New Forest, Hampshire, England : tibia, 18 '8 and 20 ;
foot, 9 • 8 and 10 • 6 ; forearm, 39 and 42 • 4 ; thumb, 8 • 4 and 10 ;
third finger, 64 and 68 ; fifth finger, 53 and 57. Adult male
from Lugano, Switzerland (Geneva : type of ghidinii Fatio) :
tibia, 19-4; foot, 10; forearm, 41-2; thumb, 10-4; third
finger, 65 ; fifth finger, 53 ; ear from meatus, 21 ± ; ear from
crown, 19; width of ear, 13; tragus, 11±. Adult female from
Ste. Baume, Var, France : head and body, 53 ; tail, 37 '4 ; tibia,
20 ; foot, 10' 4; forearm, 42-6 ; thumb, 9 ; third finger, 70 ;
fifth finger, 57 ; ear from meatus, 25*6 ; ear from crown, 24;
width of ear, 16*2 ; tragus, 11 "8. Adult male from Cadillac,
Gironde, France : head and body, 45-|- ; tail, 42 ; tibia, 20 ;
foot, 9; forearm, 41 ; thumb, 8-8; third finger, 67; fifth
finger, 56; ear from meatus, 26; width of ear, 15 "6. Adult
male from La Granja, Segovia, Spain (type of favoniciis Thomas) :
head and body, 48 ; tail, 38 ; tibia, 20 ; foot, 8 • 4 ; forearm, 40 ;
thumb, 8-2 ; third finger, 66 ; fifth finger, 55 ; ear from meatus,
23; ear from crown, 19; width of ear, 14-6; tragus, 10 '4.
Adult female from Zay-Ugrocz, Hungary : head and body, 46 • 6 ;
tail, 44 ; tibia, 22 ; foot, 9 ; forearm, 44 ; thumb, 9 ; third
finger, 73 ; fifth finger, 59 ; ear from meatus, 25 ; ear from
crown, 22"6; width of ear, 16; tragus, 11 '8. For cranial
measurements see Table, p. 183.
Specinmns examined. — Seventeen, from the following localities : —
England : Henley-on-Thames, Oxfordshire, 1 ; New Forest, Hamp-
shire, 6.
Sweden : Skdne, 1 (U.S.N.M.).
182
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184
CHIROPTERA
France : Etupes, Doubs, 1 (Mottaz) ; Ste. Baume, Var, 1 (Genoa) ;
Cadillac, Gironde, 1 (Lataste).
Switzerland: Lugano, Tioino, 1 (Geneva: type of gJiidinii Fatio).
Spain: La Granja, Segovia, 1 (type oi favonicu^ Thomas).
Austria-Hungary: Zay-Ugrocz, Trencsen, 4 (B.]\L and U.S.N.M.).
Bemarks. — This animal is recognizable among European bats
by its medium size and very large ears. On the basis of the
material examined I am unable to distinguish a Spanish or
western geographical race.
6. Henley-on-Thames, Ox- J. G. Millais (p). 6. 9. 14. 1.
fordshire, England.
1. New Forest, Hampshire. Dr. W. E. Leach (p). 56. a.
2. New Forest, Hampshire. G. W.H.Blagg(c&p). 7.7.16.1-2.
6 al. La Granja, Segovia, M. de la Escalera (c). 6. 11. 4. 1.
Spain. {Type oi M. favoiiiciis Thos.)
6, ? al. Zay-Ugrocz, Trencsen, Budapest IMuseum G. 6. 20. 1-2.
Hungary. (e).
1. Europe. Leydeii Museum. 37. 4. 28. 23.
MYOTIS DAUBENTONII Kuhl.
1819. Vcspcrtilio danbcntonii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 195 (Hanau, Hessen, Germany).
1839. Vespcrtilio xdilis Jenyns, Ann. Nat. Hist., in, p. 73, April, 1839
(Aukland St. Andrew, Durham, England).
1844. V[espertilio] lanatus Crespon, Faune Meridionale, i, p. 15 (South of
Nimes, Gard, France). Type in Nimes Museum.
1857. Vcspcrtilio daubcntonii Blasius, Stiugethiere Deutschlauds, p. 98.
1871. Vcspertilio capucincUus "Koch, Bayr. Fauna," Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wioi, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 206 (Bavaria?).
1871. Vcspcrtilio minutcllus " Koch, Bayr. Fauna," Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 206 (Bavaria?).
1871. Vcspertilio daubcntonii, albus Fitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, p. 210
(Kenaming of V. xdUis Jenyns).
1878. Vcspertilio daubentonii Dobsou, Catal. Chiropt. Brit. Mus., p. 297.
1890. Vcspcrtilio staufferi Fatio, Faune Vert. Suisse, v, 3"'" suppl. aux
JIamm., p. 6 (Lucerne, Switzerland). jNIS. synonym; repudiated.
1900. Myotis daubcntonii M6hely, Monogr. Chiropt. Hungaria", p. 164.
1910. Myotis daubentoni Trouessart, Faune Mamm. d'Europe, p. 27.
Type locality. — Hanau, Hessen-Nassau, Germany.
Geo(jrapliical dhtrihution. — Europe from the Mediterranean
north to central Norway, west to Ireland, east into Asia.
Diagnosis. — 8ize nearly as small as in Myotis viystacinas
(forearm about 35 mm., longest finger about 60 mm., condylo-
basal length of skull, 1.3-4 to 13 '8); ear moderately long,
extending about 2 mm. beyond tip of muzzle when laid forward,
its posterior border with shallow inconspicuous emargination ;
foot decidedly more than half as long as tibia ; wing membrane
extending to side of metatarsus ; last caudal vertebra free.
MYOTIS 185
External characters. — Smallest of the large-footed European
species. General form less slender and delicate than in M.
mystacinun, the tail and legs relatively shorter. Ear rather
short, extending, when laid forward, about to tip of muzzle ;
anterior border faintly and evenly convex from basal lobe to
narrowly rounded-off tip ; posterior border with shallow though
evident concavity above ; antitragus small and ill-defined ; tragus
about half as high as conch, its greatest width contained three
times in length of anterior border, the posterior border
moderately and evenly convex, the anterior border nearly
straight, the tip rather blunt ; posterior basal lobe relatively
large and well defined. Wing slightly broader than in 31. mi/sta-
cinus, the three main metacarpals evidently graduated from
third to fifth, the third very slightly shorter than forearm ;
membrane inserted at middle of metatarsus. Foot large,
appreciably more than half as long as tibia ; calcar slender,
very long, without keel on posterior border and with slight
terminal lobe, its length fully double that of free border of
interfemoral membrane. Tail about as long as body without
head, the terminal vertebra free from membrane except for an
exceedingly narrow strip extending outward along each side.
Fur and colour. — Fur slightly shorter and more dense than
that of 31. mystac'mus, but with no peculiarities of distribution
except that it tends to spread farther outward along dorsal
surface of interfemoral membrane ; free border of urojiatagium
not fringed, but a slight fringe is usually present along basal
half of calcar. Colour above essentially as in ilf. nattereri,
though usually inclining more definitely toward wood-brown ;
underparts buffy grey visually less contrasted than in 31. nattereri,
though sometimes pale enough to produce a distinct line of
demarcation along sides of neck. Muzzle and cheeks dusky.
Membranes and ears an indefinite brown.
Skull. — The skull of Myotis dauhentonii is smaller tlian that
of any other European species except 31. niystacimifi. From this
it is immediately distinguishable by its noticeably greater breadth
both of rostrum, palate and brain-case, by the relatively lower
occipital region, and relatively deeper rostrum. Posterior exten-
sion of palate short and broad, the width just behind molars
greater than length to tip of hamular ; median projection angular,
seldom forming a distinct spine. Greatest breadth of brain-case
slightly though appreciably more than greatest length of skull.
Mandible with coronoid process low, scarcely rising above level
of condyle.
Teeth. — Teeth relatively smaller than those of 3lyotiS mysta-
cinus, a difference particularly noticeable in the crown areas of
the first and second upper molars. Upper incisors as in mystacinus,
but with cingulum less developed. Lower incisors very slightly
imbricated, tlie form of the row vacillating between (J-shaped
and broadly V-shaped, the cusps as in 31. mystacinus but less
186
CHIROPTERA
Fig. 31.
Miiotif: dauheidonii. Teetli X 10.
developed. Canines both above and below weaker and less
trenchant than in M. mijist acinus, their form essentially as in
M. nattereri. Premolars
and lower molars not
obviously diflerent from
those of M. Dii/stacinus.
Upper molars peculiar as
compared with those of
the small-footed European
species in the presence of
an evident protoconule on
anterior commissure of
protocone, the small cusp
provided with a small
but distinct secondary
commissure extending to
base of paracone ; m^ and
fii^ with a small commis-
sure extending from base
of hypocone to base of
metacone and partly fill-
ing depression lying be-
tween these cusps.
Measurements. — Two
adult males from Upsala,
Sweden : head and body, 43 and 44 ; tail, 34 and 36 ; tibia, 17
and 17 '4 ; foot, 11 and 11 ; forearm, 37 and 37 ; thumb, 8 '4
and 8 ; third finger, 62 and 59 ; fifth finger, 49 and 49 ; ear
from meatus, 13 and 13 ; width of ear, 10 and 9 "6. Adult male
and female from Lecco, Italy : head and body, 42 and 45 ; tail,
36 and 39; tibia, 16 and 17; foot, 10'6 and 11; forearm,
37 and 38 ; thumb, 8 and 8 ; third finger, 62 and 62 ; fifth finger,
51 and 51 ; ear from meatus, 13 '6 and 13' 6 ; width of ear, 9*6
and 9 "4. For cranial measurements see Table, p. 190.
Specimens examined. — Seventy-three, from the following localities : —
Scotland : No exact locality, 1.
England : Bowdon, Cheshire, 1 ; Knutsford, Cheshire, 1 ; Stratford-
on-Avou, Warwickshire, 2 ; Hillingdon, Middlesex 1, ; Epping, Essex, 2 ;
Northamptonshire, 1 ; Henley-on-Thauies, Oxfordshire, 2 (B.M. and
U.S.N.M.); Christchurch, Hampshire, 1; Devonshire,!.
Sweden: Upsala, 3 (B.M. and U.S.N. M.); upland, 1; no exact locality, 2.
Switzerland : Geneva, 1 (INIottaz).
Fbance : Near Nimes, Gard, 1 (Nimes : type of lanatus Crespon).
Italy : Lecco, Lombardy, 21 (U.S.N.M.) ; Pavia, 1 (U.S.N.M.);
Florence, 30 (Mottaz).
Heniarks. — This species is immediately recognizable among
the European members of the genus by its small size, large foot,
and naked upper surface of legs.
1. Scotland. Dr. J. !Macgiilivray (p).
Bowdon, Cheshire, Eng- T. A. Coward (c&p). 11. 1. 3. GO.
land.
6.
MYOTIS 187
9. Knutsford, Cheshire. T. A. Coward (c & r). 11. 1. 3. 22.
2. Stratford-on-Avon, War- Tomes Collection. 7.1.1.486-487.
wickshiro.
i al. Hilliugdon, Middlesex. O. Thomas (c & p). 84. 1. 29. 1.
2 al. Epping, Essex. H. Doubleday (c &p). 44. 10. 21. 1-2.
al. Northampton. Mrs. Jenyns (p).
9 St. Henley, Oxfordshire. J. G. Millais (c & p). 1. 11. 2. 2.
1 al. Christchurch, Hamp- Lord Lilford (p). 87. 'J. 1. 1.
shire.
6. Upland, Sweden. Lord Lilford (p). 11. 1. 1. 25.
(G. KoUhoff.)
2. Sweden. Stockholm Museum 46. 6. 2. 15.
(e). 48. 6. 28. 3.
MYOTIS CAPACClNll Bonaparte.
1837. Vespertilio capaccinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx
(Sicily). Type in British Museum.
1839. Vespertilio megapodius Temmiuck, Monogr. de Mamm., ii, p. 189
(Sardinia).
1841. Vesp[crtilio'] dasypiis de Selys-Longchamps, Atti della seconda
Riunione degli Scienziati Italiani, Torino, 1840, p. 249 (Published
as synonym of capaccinii).
1844. Vesplcrtilio] pellucens Crespou, Fauna Meridionale, i, p. 16 (Cave
near Pont-du-Gard, Gard, France).
1857. Vespertilio capaccinii Blasius, Saugethiere Deutschlands, p. 101.
1877. Vespertilio blasii Major, Atti Soc. Tosc. Sci. Nat., Pisa, iir, p. 108
(New name for the capaccinii of Blasius should it prove to be
different from that of Bonaparte).
1878. Vespertilio majori Ninni, Atti Reale Instit. Veneto, 5th ser., iv,
pt. 1, p. 721 (Substitute for blasii Major).
1878. Vespertilio capaccinii Dobson, Catal. Chiropt. Brit. Mus., p. 293.
1901. Myotis capaccinii Thomas, Proc. Zool. Soc. London, p. 37.
1910. Myotis {Leuconoe) capaccinii Trouessart, Faune Mamm. d'Europe, p. 26.
Type locality. — Sicily.
Geographical (ligtrihiition. — Mediterranean region, north to
Italian Switzerland, east into A.sia.
Diagnosis. — Not so small as Myotis dauhentonii (forearm about
42 mm., longest linger about 68 mm., condylobasal length of
skull, 14"0 to 14*8 mm.), but similar in form and proportions;
wing membrane extending to ankle ; last caudal vertebra free ;
tibia and adjacent membrane densely furred.
External cltaracters. — In all essential features the external
form is as in M. dauhentonii. The foot, however, is relatively
larger, and the wing membrane is inserted at the ankle.
Fur and colour. — Fur rather dense and short, the hairs at
middle of back about 6 mm. in length ; distribution peculiar
among the European members of the genus in its tendency to
spread on membranes, forming a tlistinct patch on upper surface
of wing at elbow and extending over entire uropatagium to level
of feet, the furry covering of tibia and immediately adjacent
membrane (both above and below) especially dense. Colour
188
CHIROPTERA
above a light drab tinged with grey or with ecru-drab, the
general effect jjaler and more greyish than in any of the other
European species ; underparts pale butfy grey, rather strongly
contrasted and with moderately well-defined line of demarcation
along sides of neck. Underfur slaty black. Muzzle and
cheeks faintly dusky. Ears and membranes an indefinite rather
Hght brown.
Skull. — In all resi^ects the skull so closely resembles that of
Myotls daubentonii that it is only distinguishable by its larger
size. From the skulls of M. nattereri and M. emarginatus it
differs in its greater breadth, a character perhaps most readily
appreciable in the form of the jjost-palatal region.
Teeth. — Except for their larger size the teeth I'esemble those
of Myotls daubentonii. The crown area of upper molars is
relatively less than in M. nattereri and 31. einanjinatus, though
»r retains the broader outline characteristic of M. daubentonii
and M. mystacinus. Upper molars with protoconule and its
accessory small commissure, and m^ and ?n'- with commissure
between hypocone and metacone as in jW. daubentonii.
Measureynents. — Average and extremes of forearms in four
males and eight females from Lugano, Ticino, Switzerland :
males, 39-9 (38-8-41); females, 39-4 (38-8-40-4). Sicily
(type): tibia, 15* 6 ; foot, 12 ; forearm, 39-2; thumb, 10 ; thii-d
finger, 56-2 ; fifth finger, 50. Adult male from Corleone, Sicily :
head and body, 49 ; tail, 38 ; tibia, 16 ; foot, 10 ; forearm, 41 ;
thumb, 8 • 2 ; third finger, 68 ; fifth finger, 55 : ear from meatus,
15 ; width of ear, 10 '4. Two adult males from Sassari, Sardinia :
head and body, 50 and 51 ; tail, 37 and 37 ; tibia, 16 '4 and 17 ;
foot, 10-6 and 11 ; forearm, 39-4 and 41*6 ; thumb, 8-4 and
8"6 ; third finger, 66 and 69 ; fifth finger, 51 and 56 ; ear from
meatus, — and 14*4; width of ear, 10 and 10. For cranial
measurements see Table, p. 191.
Specimens e.vamincd. — Seventy-two, from the following localities: —
Austria-Hungary: Herkulesbad, 1.
Switzerland : Near Lugano, 49 (B.M., U.S.N.M., and Mottaz).
Italy : Finalborgo, Liguria, 2 (Genoa) ; Pavia, 1 (U.S.M.M.) ; Ostia,
Rome, 2; Marsala, Sicily, 1; Corleone, Sicily, 1 (U.S.N.M.); Sicily, no
exact locality, 1 (type).
Sardinia : Cagliari, 4 (B.M. and U.S.N.M.) ; Sassari, 5 (U.S.N.M.) ;
Grotte de Sardale, 2.
France: Marseilles, 1 (U.S.N.M.).
Spain : Inca, Majorca, Balearic Islands, 1 ; Elche, Alicante, 1.
Itemarlis. — In general appearance this species resembles
Myotic daubentonii ; but it is immediately recognizable by its
even larger foot, and by the densely pubescent upper surface of
leg, the latter character unique among European bats.
4 6, 9. Near Lugano, Ticino, 0. Thomas (p). 2. 8. 4. 10-14.
Switzerland.
(E. H. Zullikofer.)
2. Ostia, Rome. Dr. L. Samhon (c&p). 1.1.2.3-4.
189
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 734.
without parte.) {Type of species.)
skull.
9.
2al.
2al.
Marsala, Sicily. O. Thomas (p). 6. 8. 4. 21.
(.4. Robert.)
Cagliari, Sardinia. Florence Museum (e). 85. 7. 6. 6-7.
Grotte de Sardale, Sar- O. Thomas (p). 0. 12. 3. 1-2.
dinia.
Inca, Majorca, Balearic O. Thomas and R. I. 0. 7. 1. 3.
Islands. Poooek (c & p).
Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 13.
MYOTIS DASYCNEME Boie.
1823. Vcspertilio mijstaciniis Boie, Isis, p. 9G5. Not Y. viystacinus Kuhl
(Jutland, Denmark).
1825. Vespertilio dasycneme Boie, Isis, p. 1200 (Renaming of 7ni/slacinns
Boie).
1839. Vcspciiilio limnopldlus Temmiuck, Monogr. de Mamm., ii, p. 176
(Holland).
1857. Vespertilio dasycneme Blasius, Siiugethiere Deutschlands, p. 103.
1878. Vespertilio dasycnetne Dobson, Catal. Chiropt. Brit. Mus., p. 295.
1904. Myotis dasycneme Trouessart, Catal. Mamm. Tam viv. quam foss.,
suppl., p. 88.
1910. Myotis dasycnetne Trouessart, Faune Mamm. d'Europe, p. 27.
Type locality. — Chalk quarries at Dagbierg, near Wiborg,
Jutland, Denmark.
Geographical distribution. — Central and southern Europe,
west to the Atlantic coast,* north to Sweden, east into Asia.
Diagnosis. — Form and proportions essentialh' as in 3Iyotis
dauhentonii and M. capaccinii, but size much larger (forearm
about 47 mm., longest linger about 75 mm., condylobasal length
of skull about 16 mm.) ; tibia and adjacent membrane naked.
External characters. — Form not essentially different from the
other European members of the large-footed group. Tragus
relatively shorter than in any other European Mj/otis, its height
distinctly less than half that of conch, its anterior border slightly
concave, its posterior border slightly convex below, then more
abruptly convex to bluntly rounded ofi' tip, the two margins
essential!}' parallel through lower half. Free border of uro-
patagium without fringe.
Fur and colour. — Distribution of fur as in Myotis dauhentonii ;
quality not peculiar, the longest hairs on middle of back about
8 mm. in length. Colour of upper parts a light yellowish wood-
brown ; underparts strongly contrasted greyish white with a
tinge of but!', the line of demarcation along sides of neck well
defined. Muzzle and cheeks scarcely contrasted dusk\-. Under-
fur slaty black. Ears and membranes an indefinite dark brown.
* The supposed British record is probably erroneous (see Barrett-
Hamiltou, Hist. Brit. I\Iamm., i, pp. 157-158. December, 1910).
190
CHIROPTERA
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192 CHIROPTEEA
Skull. — The skull is large, sligbtly exceeding that of Myotis
hecltsteinii in length and noticeably surpassing it in breadth and
robustness. Its general appearance is the least typically Myotis-
like of any European member of the genus, a peculiarity
heightened by the crowding of the small premolars and conse-
quent shortening of anterior portion of tooth-row. Allowance
being made for the great difference in size the skull is not very
unlike that of Myotis dcmbentonii, but the rostrum is relatively
shorter and the brain-case broader and more depressed. Sagittal
crest barely indicated : auditory bull* proportionately as in the
smaller animal, less inflated than in 3/. bechsti'ini! ; lachrymal
ridge well developed, its lower extremity separated from lachrymal
foramen by an evident notch.
Teeth. ^In form the teeth resemble those of the small Myotis
dauhentonii ,■ molars decidedly more robust than those of M.
berlisteinii, the width of the crown diminishing much less rapidly
toward inner border ; small premolars much crowded, the second
barely or not visible from outer side, the diameter of its crown,
however, not much less than that of anterior tooth ; upper
molars as in M. dauhentonii and M. capaecinii ; mandibular teeth
with no special peculiarities, the premolars and outer cusps of
molars more slender than in M. hechsteinii and 31. dnuhentonii.
Measurements. — Two adult males from Maastricht, Holland :
head and body, 57 and 58 : tail, 49 and 51 ; tibia, 19-8 and
20-2; foot, 11-8 and 12 ; forearm, 43-6 and 44 ; thumb, 9-6
and 9 ; third finger, 72 and 77 ; fifth finger, 57 and 62 ; ear
from meatus, 17 and 17-2; width of ear, 10-6 and 11. Two
adult females from the same locality : head and body, 60 and 61 ;
tail, 47 and 46 ; tibia, 18 and 18 ; foot, 11-4 and 11 ; forearm,
44 and 44 ; thumb, 9 and 9 : third finger, 76 and 75 ; fifth
finger, 61 and 60 ; ear from meatus, 16-6 and 17 ; width of ear,
11 and 11. For cranial measurements .see Table, p. 191.
Specimens cxcanincd. — Six, from tlie following localities: —
Holland: Leyden, 1; Maastricht, 4 (U.S.N.M.).
Belgium : Near Namur, 1.
1. Leyden, Holland. Tomes Collection. 7. 1. 1. 501.
(H. Schlcgel.)
6 al. Namur, Belgium. Re%-. D. B. Lebbe 9. 1. 11. 1.
(c & p).
MYOTIS MYOTIS Borkhausen.
1775. Vespertilio murinus Schreber, Saugthiere, i, i>. 1G5 (Not of Linnfeus,
1758).
1797. Vespertilio myotis Borkhausen, Deutsche Fauna, i, p. 80 (Germany).
1797. Vespertilio myosotis B[orkhause]n, Der Zoologe (Compendiose
Bibliothek gemeinniitzigsten Kenntnisse fiir alle Stiinde, pt. xxi),
Heft v-viii, p. 46 (Germany).
1800. Vespertilio myosotis Bechstein, Pennant's Allgemeine Uebersicht der
Vierfiissigen Thiere, p. C32 (Germany).
MYOTIS 193
1801. Vespertilio viyotis Bechstein, Gemeinn. Naturgesch. Deutschl., i,
2ud ed., p. 1164 (Described but not named in 1st ed., 1789, x'. 1G4)
Thiiriugen, Germany.
1827. Vespertilio subvmrinus Brehm, Ornis, Heft iii, p. 24 (Renthendorf,
Tliiiringcn, Germany).
1844. V[espertilio] latipinnis Crespon, Faune Meridionale, i, p. 17 (Near
Nimes, Gard, France).
1857. Vespertilio murinus Blasius, Saugethiere Deutschlands, p. 82 (Not of
Linnteus, 1758).
1863. [Myotus tnurinus] var. typus Koch, Jahrb. des Vereins fur Natur-
liunde ini Herzogthum Nassau, xviii, p. 415 (Wiesbaden, Nassau,
Germany).
1863. [Myotus vmriuus] var. alpinus Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 415 (St. Gothard, Uri,
Switzerland).
1878. Vespertilio murinus Dobson, Catal. Chiropt. Brit. ]\Ius., p. 309 (Not
of Linnseus, 1758).
1886. Myotis murina var. spelxa Bielz, Verhandl. u. Mittheilungen des
Siebenbiirgischen Vereins fiir Naturwissensch. in Hermannstadt,
XXXVI, p. 83 (Homorod-Almas cave, Hungai-y).
1897. Myotis myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 383,
October, 1897.
1909. Myotis myosoiis Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S., iii,
No. 3, p. 1, April 26, 1909.
1910. Myotis myotis and M. m. spelasa Trouessart, Fauna Mamm. d'Europe,
p. 32.
Type locality. — Thliringen, Germany.
Geographical distribution. — Central and southern Continental
Europe, west to Portugal, north to southern Sweden, eastward
into Asia. One record of its occurrence in England."'
Diagnosis. — Largest species of European Myotis (forearm,
57 to 64 mm.; longest linger, 100 to 110 mm.; condylobasal
length of skull, 22 to 23 '6 mm.) ; form heavy, membranes thick
and leathery ; ear moderately long, extending about 5 mm.
beyond tip of muzzle when laid forward, its posterior margin
scarcely or not emarginate above middle ; foot slightly more than
half as long as tibia ; wing membrane extending to base of
outer toe.
External characters. — Although one of the largest European
bats Myotis myotis does not differ conspicuously in form from
the small M. mystacinus, except that its tail and legs are relatively
shorter. The general build is not I'emarkably heavy as compared
with other European species of approximately the same size, but
the ears and membranes are rather thick and leathery. Ear
moderately long, extending about 5 mm. beyond nostril when
laid forward ; anterior mai'gin moderately convex from base
nearly to rather narrowly rounded-off tip ; posterior border wath
shallow ill-defined concavity above ; antitragus low and long,
marked off posteriorly by a well-deHned notch and not continuous
with posterior border of conch ; tragus about half as high as
* Bell, Hist. British Quadrupeds, p. 88, 1836 : "But in England it . . .
has hitherto only been taken in the gardens of the British Museum."
O
194
CHIROPTERA
conch, its greatest width (slightly above level of antei'ior base)
contained about '2h times in length of anterior margin, the
posterior border convex to just below rather bluntly rounded tip,
the anterior margin straight, posterior basal lobe small but well
developed ; inner surface of conch with seven or eight ill-defined
transverse ridges near posterior border. Wing rather broad but
with no special peculiarities of form ; metacarpals somewhat
elongated relatively to jahalanges as compared with the smaller
species, the thii-d, fourth and fifth slightly but evidently
graduated, the third scarcely shorter than forearm ; membrane
inserted at side of metatarsus, but with a narrow strip extending
to base of outer toe as in M. nattereri. Foot slightly more than
half as long as tibia ; calcar heavy at base but tapering rapidly
and terminating obscurely, its posterior border with slightly
indicated keel, its length about equal to that of free border of
interfemoi-al membrane. Tail about as long as body without
head, only the extreme cartilaginous tip free from membrane.
Fur and colour. — Relatively to size of animal the fur is rather
short (longest hairs of back about 10 mm.); in distribution it
shows no peculiarities ; free border of uropatagium without
fringe. Colour an indefinite brown much like that of Mi/otis
nattereri, the exact shade intermediate between the wood-brown
and broccoli-brown of Ridgway, usually paler on head and neck
than on back, and in immature individuals than in adults, the
hairs slate-black through basal half, then light wood-brown
followed by a darker though not strongly contrasted terminal
area. Underparts strongly contrasted greyish white with a
slight bufly tinge ; a well-defined
line of demarcation along sides
of neck to ear, emphasized in
region of shoulder by a slight,
diffused blackening of edge of
dark area. Muzzle and cheeks
dusky. Ears and membranes an
indefinite brown.
Skull. — Though much larger
than that of any of the other
European .species of Myotu, M.
oxi/gnathus excepted, the skull of
Ml/of is myotis is one of the most
slender in general outline. . The
brain-case is longer proportion-
ately to its breadth and is less
contrasted with rostrum than in
the small members of the group ;
greatest breadth of brain-case
about one-half distance from
lambda to posterior margin of nares. Rostrum relatively deep
and interorbital concavity relatively shallow ; occipital region
Fig. 32
Myolis inyoth.
195
about on level with main portion of brain-case, and very slightly
overhanging foramen magnum ; \ antral profile scarcely elevated
posteriorly ; palate rather narrow (essentially as in M. mystaciiins) ;
width of posterior extension of palate less than its length, median
spine short but well developed ; posterior border of anteorbital
foramen over anterior root of first molar ; mandible with coronoid
process relativelv higher and narrower than in the small species,
its posterior border much more oblique.
Teeth. — The dentition is of a less primitive type than in the
small European species of Mijotls. This is indicated by the
general tendency to reduction shown especially in the lower
incisors, the small premolars and the posterior lower molar.
Upper incisors rather high and slender but not essentially
different in form from those of M. mystacinus, the cingulum of
inner tooth obsolete, that of outer slightly developed. Lower
incisors very strongly im-
bricated, the outline of the
row U-shaped or broadly
V-shaped ; cutting edge of
/j and /^ trifid, but decidedly
oblique owing to reduction
in size of outer cusp ; a
minute cingulum cusp usu-
ally present at extreme outer
edge ; i^ with small postero-
internal tubercle ; ^3 sub-
terete, 4-tuberculate, the me-
dian outer tubercle largest.
Canines relatively smaller
than in M. mystacinus and
with less developed posterior
cutting edge, their form
essentially as in M. nnttenri
and M. emarr/inatus. Small
upper premolars much
crowded, the second usually
forced inward from line of tootli-row, sometimes so much so that
the first is practically in contact with large premolar. In form
they show no special peculiarities ; shaft subterete ; cingulum
well developed. Large upper premolar more reduced than in the
small species, the inner margin of crown a mere cingulum at base
of main cusp, and without trace of secondarj' cusps. Lower
premolars not peculiar in form, but middle tooth usually more
crowded between first and third than in the small members
of the group. Molars .showing no special peculiarities of form ;
thickening that represents hypocone in iit^ and m- barely
indicated ; »(^ more reduced than in the small .species, its meta-
cone scarcely more than a slight widening of terminal portion of
third commissure. First and second lower molars essentially as
Fig.
Mijotis myotis.
196
CHIROPTERA
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197
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198
CHIROPTERA
in the small species, except that cingulum does not form a postero-
internal cusjj behind entoconid ; ^3 with hypoconid and ento-
conid more reduced than in the small species, the hypoconid
displaced further inward, so that second triangle is barely half
as large as first and conspicuously difierent from it in form.
Measurements. — Average and extremes of four adult females
from Tagerwilen, Thurgau, Switzerland: head and body, 75 '7
(72-79); tail, 56-3 (54-60); tibia, 26-1 (25 -4-26 -6); foot,
14-7 (13-16); forearm, 63 (63); thumb, 13-1 (12 -8-13 -4) ;
third finger, 107-7 (106-109); fifth finger, 85-2 (84-86); ear
from meatus, 27-6 (27-28) ; width of ear^lS-l (17-6-19). Two
adult females from Mte. Generoso, Ticino, Switzerland : head
and body, 72 and 76 ; tail, 50 and 51 ; tibia, 25 and 25 '4 ; foot,
15 and 14-4; forearm, 62 and 63-6 ; thumb, 13 and 12 ; third
finger, 104 and 107 ; fifth finger, 83 and 84 ; ear from meatus,
27 and 28; width of ear, 18 and 18-6. , Adult male from
Florence, Italy : head and body, 68 ; tail, 55 : tibia, 25 ; foot,
13; forearm, 61 ; thumb, 12; third finger, 100 ; fifth finger, 78 ;
ear from meatus, 27-6; width of ear, 17-6. For cranial
measurements see Table, p. 196.
Spec'uncns examined. — Ninety-five, from the following localities : —
France : Cadillac, Gironde, 1 (U.S.N.M.) ; Nimes, Gard, 2 (B.M. and
Nimes ; rexjresenting latiinnnis Crespon, but not type).
Germaky : Hamburg, 1 ; Niesiiy, Silesia, 11 ; Strass, near Burgheim,
Bavaria, 8 ; Heidelberg, Baden, 1.
Austria-Hungary : Herkulesbad, 1 ; Fiinfkircben, S.W. Hungary, 2.
RouMANiA : Bustenari, 3; Sinaia, 1 (U.S.N.JI.).
Switzerland : Geneva, 7 (Mottaz) ; Grotte de Vallorbe, Vaud, 1
(Mottaz) ; Boudry, Neucbatel, 2 (Mottaz) ; St. Moritz, 1 ; Thayngen, Schafif-
hausen, 2 (U.S.N.M.); Canton Thurgau, 5; Tiigerwilen, Thurgau, 10
(U.S.N.M.); Andermatt, Uri, 1 (U.S.N.M.); St. Gothard, Uri, 3 (B.M.
and U.S.N.M.) ; Jilonte Generoso, Ticiuo, 4 (U.S.N.M.).
Italy: Domodossola, 2 (U.S.N.M.); Fiualborgo, Liguria, 1 (Genoa);
Florence, 1 (U.S.N.M.) ; Rome, 2; Ostia, Rome, 2 ; ]Marsala, Sicily, G.
Sardinia : Oristano, Cagliari, 10.
Spain : Seville, 2.
Portugal: Cintra, 2.
Meniarhs. — Mijotis rityofis diflers strikingly from the other
European members of the genus, M. oxygnatJms excepted, in its
much larger size. From the large Vespertilionklse of other genera
it is immediately recognizable by its long ears, extending notice-
abl}' beyond nostril when laid forward, and by the greyish white
colour of the underpiarts. In the MediteiTanean region Myotis
myotis is associated with M. oxygnathus ; but it is the only large
species known to occur north of the Alps.
1 al.
Nimes, Gard, France.
G. E. Dobson (e).
80.
12. 14. 2.
1.
Hamburg, Germany.
Dr. J. E. Gray (p).
2 6, 8 V,
Niesky, Silesia. {Dr.
Dr. E. Hamilton (p).
97.
12. 4. 7-17.
9 juv.
W. Baer.)
9.
Strass, Burgheim, Ba-
varia. (Korbitz.)
Lord Lilford (p).
11.
1. 1. 10-15.
9.
Strass, Burgheim, Ba-
Lord Lilford (p).
11.
1. 1. 124.
varia. {Korbitg.)
11.
1. 1. 131.
MYOTIS
199
?. Heidelberg, Baden. Hon. N. C. Eoths- 10. 5. 29. 1.
child (p).
i juv. Herkulesbad, Hungary. Hon. W. Rothschild 7. 9. 16. 8.
[F. J. Cox.) (p).
2al. Fiintkirchen. BudapestMuseum(E). 94.7. 18. 11-12.
2 <J, 9. Bustenari, Prahova, Lord Lilford (p). 4.4.6.8-10.
840 m. Roumania.
(W. Dodson.)
1. St. Moritz.Grisons, Swit- Leon O. Galliard (p). 75. 9. 20. 4.
zerland.
5 ?, Thurgau. [E. H. Zolli- 0. Thomas (p). 4. 4. 5. 9-13.
kofcr.)
2 9. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 26-27.
(!>)■
2. Ostia, Rome. Dr.L. Sambon(c&p). 1. 1. 2. 1-2.
6 9. INIarsala, Sicily. (.4. O. Thomas (p). 6. 8. 4. 17-20.
Bobcrt.) 8. 9. 1. .3-4.
10 al. Oristano, Cagliari, Sar- Hon. N. C. Roths- 7. 5. 24. 1-10.
dinia. child (p).
2 S al. Seville, Spain. (Dr. A. Lord Lilford (p). 73. 1. 8. 1-2.
Eitiz.)
2 i. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 4-5.
MYOTIS OXYGNATHUS Monticelli.
1SS5. Vespcrtilio oxygnathus Monticelli, Ann. Accad. 0. Costa de Aspir.
Nat., I, p. 82. Type in Naples Museum.
1909. Mijotis oxygnathus Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S.,
III, No. 3, p. 1, April 26, 1909.
1910. Myotis myotis oxygnathus Trouessart, Paune Mamm. d'Europe, p. 32.
Type locality. — Matera, Basilicata, Italy.
Geographical distribution. — Mediterranean region from Spain
to Greece, north to Italian vSwitzerland ; Sardinia ; Malta ;
Tunis.
Diagnosis. — Similar to Myotis myotis but smaller and with
shorter, narrower ears; condylobasal length of skull, 18 "6 to
21 • 4 instead of 22 to 23 • 6 mm. ; mandible, 15-2 to 17-2 instead
of 17 "8 to 19 mm.; maxillary tooth-row, 8' 2 to 9' 4 instead of
9-8 to 10-6 mm.
Measurements. — Type (adult male) : head and bod}', 63 ; tail,
54; tibia, 24-4; foot, 13; forearm, 57; thumb, 11-4; third
finger, 98 ; fifth finger, 76 ; ear
(Dartmoor, Devonshire, England).
1834. Vespertilio brachyotos Baillon, Mem. Soc. Royale d'Emulation
d'Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1839. ? [Vespiertilio pipistrellus'] var. nigra de S61ys-Longchamps, Etudes
de Micromamm., p. 140 (nomen nudum).
1839. ? [Yespertiiio pipistrellus'] var. rnfescens de S61ys-Longchamps, Etudes
de Micromamm., p. 140 (nomen nudum).
1840. Y[esp)ertilio] pusillus Schinz, Europ. Fauna, i, p. 9 (Synonym of
pipistrellus; Brehm cited as authority).
1840. Y[espertilio] melanopterus Schinz, Europ. Fauna, i, p. 9. Brehm
cited as authority, but name apparently published here for first
time (Rhentendorf, Tliiiriugen, Germany).
1840. Y\_cspertilio] stenotus Schinz, Europ. Fauna, i, p. 9. Brehm cited as
authority, but name apparently publislied here for first time
(Rhentendorf, Thiiriugen, Germany).
1840. Vespertilio mintitissimus Schinz, Europ. Fauna, i, p. 9 (Ziirich,
Switzerland).
1845. P[ipistrellus'] nigricans Bonaparte, Atti dclla sesta Riunione degli
Scienziati Italiani, Milano, 1844, p. 340. Described but not named
in Atti della seconda Riunione degli Scienziati Italiani, Torino,
1840, p. 247, 1841. (Sardinia.)
1845. Pipistrellus genci Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, Milano, 1844, p. 340 (Alternative name for nigricans).
1845. P[ipistrcUus] typus Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, ililano, 1844, p. 340 (Substitute for pipistrellus
Schreber).
1867. Yesperugo pipistrellus Blasius, Siiugethiere Deutschlands, p. 61.
1862. Yesperugo pipistrellus var. macropterus Jeitteles, Verhandl. der k. k.
Zool. Bot. Gesellsch., Wien, xii, p. 250 (Kaschau, Hungary).
1863. \_Nannugo pipistrellus] var. typjis Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 490. Not of
Bonaparte, 1845 (Wiesbaden, Hessen-Nassau, Germany).
PIPISTRELLDS 205
1863. [Nannugo pipistrellus] var. flavcscens Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 491 (Nassau,
Germany).
1863. \_Nannugo piiiistrelhis] var. nigricans Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 491. Not of
Bonaparte, 1845 (Nassau, Germany).
1868. {Nannngo pipistrellns] var. limbafiis Koch, Jahrb. des Vereins fiir
Naturkunde im Herzogthum Nassau, xviii, p. 491 (Siegen, Nassau,
Germany).
1878. Vcsjierugo jnpistreUus Dobson, Catal. Chiropt. Brit. Mus., p. 223.
1897. Pipistrellns pipistrcllus Miller, Ann. and Mag. Nat. Hist., 6th ser.,
XX, p. 384, October, 1897.
1904. Pipistrellus pipistrcllus mediterraneus Cabrera, Mem. Soc. Espaii.
Hist. Nat., II, p. 273 (Valencia, Spain).
1910. Pipistrellus piipistreUus and P . pipistrellus mediterraneus Trouessart,
Faune Mamm. d'Europe, pp. 14-15.
Type locality. — France.
Geographical distribution. — Europe from the Mediterranean
north to Scotland and Scandinavia, west to Ireland and the
Hebrides, east into Asia.
Diagnosis. — Smallest European member of the genus (forearm,
27-6 to 32 mm. ; condylobasal length of skull, 11 to 12 mm.);
outer upper incisor more than half as high as inner incisor ;
large upper premolar separated from canine by a distinct space,
the small tooth visible from outer side, its crown area about
equal to that of outer incisor ; anterior lower premolar with
crown area equal to more than half that of succeeding tooth ;
lower canine robust, the length of base along cingulum about
equal to length of anterior border of shaft ; tragus with greatest
width less than length of anterior border ; thumb short, its
length about equal to width of wrist ; length of fifth finger about
40 mm. ; posterior edge of wing membi-ane usually dark.
External characters. — General form robust, the tail and legs
rather short, the membranes relatively thick and opaque. Muzzle
with very noticeable glandular swellings extending back to
beneath eye. Ear extending about to nostril when laid forward,
its general form rather short and broad, though with narrowly
rounded tip ; anterior border abruptly convex at base, then
essentially straight almost to tip ; posterior border faintly and
irregularly concave above, evenly convex below, the antitragal
lobe represented by a thickened ridge extending along margin of
ear and turning abruptly inward without producing any notice-
able break in outline of conch. Inner surface of conch slightly
rugose, but without evident cross ridges. Tragus erect, scarcely
half as high as conch, its tip broadly rounded, its greatest width
(slightly above level of anterior base) about half length of anterior
border ; except near tip, both borders are nearlv straight or very
slightly convex ; posterior basal lobe small but well defined.
Wing rather nari'ow, with no special peculiarities of form, the third,
fourth and fifth metacarpals sub-equal (fifth slightly shorter than
206 CHIROPTERA
the others) and extending nearly to point of elbow ; fifth finger
extending beyond elbow to a distance equal to less than one-
third length of forearm ; thumb short, its length about equal to
width of wrist ; membrane inserted at base of outer toe. Foot
about half as long as the short, robust tibia ; calcar considerably
longer than free border of interfemoral membrane, robust at base,
but tajjering rapidly and terminating without lobe, its keel well
developed, with evenly convex margin. Tail about as long as
body without head and 2J times as long as tibia, the short
terminal vertebra free from membrane.
Fa7- and coluur. — The fur is closely confined to the bod}^
showing no tendency to spread on membranes. On wing it
extends, both above and below, to line joining knee and basal
third of humerus ; lower surface of interfemoral membrane
essentially naked except at extreme base, upper surface furred
nearly to middle. Colour of upper parts a uniform brown, in
most specimens nearly intermediate between the wood-bi"own and
cinnamon of Rid way but sometimes darker, with a strong tinge
of jirouts-brown or raw umber, this especially noticeable in
immature specimens, though occasionally evident in adults ;
under parts essentially like back though slightly less dark : hairs
everywhere slaty brown at base, those of upper parts with tips
darker than sub-terminal band, but not enough so to produce a
definitely tricolor effect. Ears and membranes blackish.
Skull. — Notwithstanding its small size, less than that of any
other European bat, the skull is robust and heavily built as
compared with that of the small species of Myotis. Dorsal profile
rising gradually from nares to lambda, with slight concavity in
interorbital region and slight convexity over middle of brain-case ;
occipital region scarcely produced backward except for a median
swelling between foramen magnum and lambda,
on each side of which a condyle is just visible
when skull is viewed from above ; ventral profile
nearly flat except for a slight upward bend pos-
teriorly. Brain-case ovate in general outline,
its region of greatest breadth distinctly beliind
middle, its surface smooth or with faintly indi-
cated sagittal crest and lateral portion of
lambdoid crest ; greatest breadth of brain-case
noticeably exceeding that of rostrum and slightly
Piput^l^'pivis- though evidently more than half greatest length
treiius. Nat. size, of skuU ; floor of brain-casc flat, without vacui-
ties ; a distinct groove between cochlea and
median portion of floor, this groove bounded antero-externally
by a slight though usually evident longitudinal ridge ; auditory
bulla' moderately large, not peculiar in form ; interorbital region
l^roadly hour-glass shaped, its least breadth about equal to
breadth across roots of canines ; between constriction and ante-
orbital foramen the orbital margin is slightly but evidently
PIPISTRELLUS
207
inflated, the inflated I'egion with a median angle suggesting
a rudimentary poster bital process ; rostrum short and broad,
narrowing gradually in fi'ont, a sliglit concavity at each side
bortlering lachrymal inflation, and an evident median longi-
tudinal groove, most noticeable posteriorly ; nasal emargina-
tiou slightly deeper than wide, extending less than half way to
interorbital constriction ; anteorbital foramen small, over point of
contact between large premolar and first molar ; palate broad,
distinctly concave both longitudinally and laterally ; anterior
emai'gination small, wider than deep, its posterior border on line
with posterior edge of canine ; mesopterygoid fossa squarish,
encroached on anteriorly by broadly triangular median palatal
spine ; hamulars slightly turned inward. Mandible robust, the
ramus much deeper at symjjhysis than behind tooth-row, the
coronoid process so low that upper edge of posterior portion of
mandible is squarely and horizontally truncate, parallel with
alveolar line ; angular process short but well developed, on level
with alveolar line, its extremity slightly bent inward.
Teeth. — Relatively to size of skull the teeth are rather large
and robust, though inclined to be low, tendencies especially
noticeable in the canines. Inner upper incisor robust, its shaft
nearly half as high as that of canine, and directed strongly
forward and slightly inward, its crown iiTegularly elliptical-oval
in outline, with main axis nearly in line of tooth-row ; secondary
cusp large and conspicuous, about half as high as main shaft,
from the postero-external surface of which it projects ; cingulum
well developed, often forming a minute postero-basal cusp. Outer
upper incisor slightly but evidently smaller than inner, its shaft
more than half as high as that of inner, to secondary cusp of
which its extremity is closely approximated ; crown outline
essentially as in inner tooth but main axis lying at right angles
to tooth-row ; posterior surface of shaft broadly concave ; inner
margin with small though distinct secondary cusp ; cingulum
moderately well developed. The main cusps of the two teeth lie
in line of general curve of anterior portion of tooth-row. Space
between outer incisor and canine about equal to greatest diameter
of incisor. Lower incisors forming a continuous, broadly (J'shaped
row between canines, their crowns very slightly imbricated ;
crowns much longer than high, trifid, that of i■^ narrowest,
longest and lowest, that of /._, and (3 widened posteriorly but
without additional cusps or tubercles. Upper canine robust,
the greatest diameter of its crown about three-quarters length
of anterior border of shaft, the cross section of shaft broadly
triangular with longest side formed by nearly flat postero-internal
surface ; a sharply defined antero-exteral longitudinal groove, and
less definite postero-external concavity ; anterior edge narrow-
but not strictly trenchant ; posterior edge trenchant, with well
marked angle slightly below middle, this angle frequently becom-
ing a distinct secondary cusp ; cingulum well developed but not
CHIROPTERA
forming true basal cusps. Mandibular canine low and heavy,
its apex scarcely rising above level of highest molar cusps, its
greatest diameter measured along cingulum nearly or quite equal
to length of anterior border of shaft ; cingulum well developed,
foi'ming a distinct antero-basal cusp, the apex of which rises to
level of middle of posterior border of shaft. Anterior upper
premolar with area of crown approximately equal to that of upper
incisor and about one-hfth that of canine. It is somewhat
crowded inward from tooth-row, though about half of its crown is
visible from outer side in space between canine and large
premolar ; main cusp short but well developed, lying somewhat
in front of middle of crown, the general form of the tooth much
like that of canine but proportionally lower. Large upper
premolar with crown area about equal to that
of canine or slightly greater, the inner portion
narrow and flattened-concave, with evident
elevated rim, the posterior border strongly
concave, the anterior border usually convex
' but occasionally a little concave ; height of
main cusp slightly greater than that of highest
Fig. 35. molar cusps and about equal to length of
AnterioTteeth X 5"* tooth along outer cingulum, posterior cutting
edge AVell developed ; secondary cusp low but
evident, rising from cingulum at antero-internal base of main
cusp. Lower premolars with crown areas not conspicuously
unequal, though that of second perceptibly greater than that of
first ; outline of crown of each tooth rhombic, the outer border
somewhat convex, the anterior border of second relatively shorter
than that of first ; cingulum well developed, forming a slight
antero-internal basal cusp ; main cusp triangular in outline when
viewed from the side, that of second as high as molar cusps, that
of first shorter, the antero-external surface of each tooth convex,
the internal and posterior surfaces concave. First and second
upper molars sub-equal, though transverse diameter is relatively
greater in latter than in former ; inner border rather narrowly
rounded, the region of greatest convexity a little in front of
middle ; anterior and posterior borders straight or slightly
concave ; protocone robust though rather low ; hypocone small
but well developed, though not completely distinct from posterior
commissure of protocone ; metacoue higher than paraccme ; styles
well developed ; VV'P^ttern normal ; in'^ with crown area about
two-thirds that of vi^, the hypocone absent, the metacone smaller
than paracone ; no trace of metastyle or fourth commissure.
Lower molars with no special peculiarities ; protoconid higher
than liypoconid in all thi'ee teeth ; hypoconid with greater basal
area than protoconid in j/^j and m.^, but with less in m.^ ; cingulum
well developed, forming a slight posterointernal cusp behind
entoconid.
Measureinenfs. — Adult male from Henley-on-Thames, Oxford-
PIPISTRELLUS
209
shire, England : hccad and body, 44 ; tail, 32 ; tibia, 10-4 ; foot,
6 ; forearm, ."50 -4 ; thumb, 4-4 ; third finger, 52 ; fifth finger,
.■}8. Two adult males from Horrento, Italy : head and body, 42
and 43 ; tail, 29 and 32 ; tibia, 10 -8 and 10 ; foot, 5-8 and 6-2 ;
forearm, 30-4 and 31 ; thumb, 4-8 and 4*2; third finger, 53
and 53 ; fifth finger, 39 and 38 ; ear from meatus, 11*4 and
1 1 • 4 ; width of ear, 8 • 2 and 8 • 2. Two adult females from the
same locality : head and body, 39 and 40 ; tail 32 and 33 ; tibia,
10 "8 and 10-6 ; foot, 6 and 6 ; forearm, 32 and 30-2 ; thumb,
5 and 4*4 ; third finger, 58 and 52 ; fifth finger, 41 '6 and 40 ;
ear from meatus, 1 2 and 1 1 " 2 ; width of ear, 8 • 4 and 8. Adult
female from l^urgos, Spain, and adult female from Ciudad Real,
Spain : head and body, 49 and 39 ; tail, 31 and 30 ; tibia, 10 '6
and 9-6 ; foot, 4-8 and 5-6 ; forearm, 32 and 28-8 ; thumb, 5
and 4-6 ; third finger, 57*6 and 51 ; fifth finger, 42 and 38 ; ear
from meatus, 10 '4 and 10 '4. Extremes of twenty males from
Florence, Italy : head and body, 33-38 ; tail, 26-31 ; tibia, 9'2-
9-6; foot, 5 • 0-5 • 2 ; forearm, 27 ■ 6-30 ; thumb, 5 • 0-5 • ; third
finger, 49-53 ; fifth finger, 34 ■ 6-40. For cranial measurements
see Table, p. 210.
Speciviens examined. — Two hundred and seventy-nine, from the following
localities : —
Ireland : Co. Longford, 1 ; Co. Antrim, 1.
England : Alnwick, Northumberland, 1 ; Bowdon, Cheshire, 1 ; Great
Grimsby, Lincolnshire, 1 ; Henley-on-Thames, Oxfordshire, 1 (U.S.N.M.) ;
Tring, Hertfordshire, 4 : Lilford, Northamptonshire, 2 ; Aberia, Merioneth-
shire, 1; Chelmsford, Essex, 2; London, 1; Wimbledon, Surrey, 1; Twig-
worth, Gloucestershire, 1 ; New Forest, Hampshire, 1 ; Netley, Hamp-
shire, 1 ; Loddiswell, Devonshire, 1.
Sweden : Upsala, 6.
Denmark : Hillerod, Zealand, 1.
France : Boulogne -sur-Mer, Pas-de-Calais, 2 ; Etupes, Doubs, 1
(Mottaz) ; Nimes, Gard, 3 (Mottaz and Nimes, the last wrongly marked
type of nigrans Crespon) ; St. Geuies, Gard, 1 (Mottaz) ; Marseilles, 1
(U.S.N.M.)
Germany : Bonn, 4 ; Ingelhcim, Rheinhessen, 1 ; ^Magdeburg, Saxony, 1 ;
Berlin, 1 ; Rudolstadt, Bavaria, 3 ; Niesky, Silesia, 1.
Austria-Hungary : Transylvania, 1 ; Zara, Dalmatia, 1.
Switzerland : Geneva, 15 (ilottaz) ; Buchillon, Vaud, 4 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Neuchatel, 1 (Mottaz) ; Cortivallo, Ticino,
1 (U.S.N.M.); Mt. San Salvatore, Ticino, 13 (U.S.N.M.).
Italy: Campiglio, Tirol, 1; near GVenoa, 10 (U.S.N.M. and Genoa);
Isola Giglio, 2 (CJenoa) ; Florence, 103 (U.S.N.M. and Mottaz) ; Rome, 2
(U.S.N.M.); Sorrento, 21 (U.S.N.M.); Mondulo, Sicily, 1 (U.S.N.M.);
Palermo, Sicily, 1 (U.S.N.M.) ; Sicily, no exact localitv, 1 ; Ustica Island, 1
(U.S.N.M.). ^ ^ ^-
Sardinia : Cagliari, 17 (B.IM. and Genoa) ; no exact locality, 2.
Greece: Athens, 1 (U.S.N.M.); Tatoi, near Athens, 11; Kephissia,
near Athens, 4.
Spain : Villalba, Lugo, 1 ; Burgos, 3 ; Silos, Burgos, 2 ; La Granja,
Segovia, 4; Ciudad Real, Madrid, 2; Granada, 1; Seville, 1; Aloala, near
Seville, 1.
Jiemarl's. — PipigtrelluH pipisttellns is the smallest as well as
one of the commonest and most generally distributed of European
p
210
CHIROPTERA
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PIPISTEELLUS
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212
CHIROPTEEA
bats. Superficially it may be distinguished from the almost
equally small Mijotis miistacinua by its smaller ears and shorter
legs ; but for positive discrimination from the members of the
genus Pijjiatrellus recourse to the more technical characters of
skull and teeth is necessary. On superficial examination it
may usually be recognized among its congeners by its small size,
and by the shortness of the fifth finger.
<5al.
Co. Longford, Ireland.
Dr. G. E. Dobson (p).
76. 2. 12. 1.
9 St.
Co. Antrim.
Hon. N. C. Rothschild
P)-
1. 9. 3. 6.
?.
Alnwick, Northum-
berland, England.
W. E.de Winton (p).
11. 1. 3. 390.
2<5, 2 9.
Tring, Hertfordshire.
Hon. N. C. Rothschild
P)-
9. 2. 19. 1-4.
9.
Lilford, Northampton
shire.
Lord Lilford (p).
11. 1. 1. 125.
2 9.
Chelmsford, Essex.
M. Christy and E.
Thompson (p).
L.
11. 1. 3. 23-24.
9 juv. al.
London.
Dr. A. Giinther (p).
74. 7. 6. 1.
i.
Wimbledon, Surrey.
C. H. B. Grant (c & p)
11. 1. -6. 27.
juv.
New Forest, Hamp-
shire.
Col. J. W. Yerbury (c &
'p)-
11. 1. 3. 389.
9al.
Netley, Hampshire.
Dr. G. E. Dobson (c &
P)-
76. 11. 3. 1.
9.
Loddiswell, Devon-
shire.
Col. J. W. Yerbury (c &
P)-
11. 1. 3. 25.
3 9.
Up sal a, Sweden.
(Kolthoff.)
Lord Lilford (p).
11. 1. 1. 26-28.
S.
Hillerod, Zealand,
Denmark.
0. Thomas (c & p).
96. 6. 7. 1.
6, 9.
Boulogne, Pas - de -
Calais, Franco.
0. Thomas (c & p).
98. 1. 9. 1-2.
6, 9 al
Bonn, Rhineland,
Germany.
Dr. A. CUinther (p).
9.
Ingelheim, Rhein-
hessen.
C. H. Hilgert (c).
8. 11. 2. 3-4.
9al.
JMagdeburg, Saxony.
Dr. W. WolterstorfE (p
).
92. 12. 1. 2.
Sal.
Berlin.
Dr. A. Giinther (p).
66. 2. L22.
9 al.
Transylvania, Hun-
gary.
G. G. Danford and J.
Brown (c & p).
A.
74. 7. 4. 6.
6.
Zara, Dalmatia.
Lord Lilford (p).
11. 1. 1. 29.
2 al.
Ciudad Real, Spain.
X. Cabrera (p).
8. 7. 23. 2-3.
9.
Burgos, Prov. Burgos
. G. S. Miller (c).
8. 8. 4. 14.
9.
Silos, Burgos.
Ct. S. Miller (c).
8. 8. 4. 15.
1 al.
Campiglio, Tirol,
G. C. Champion (c & p).
96. 8. 7. 1.
Italy.
i juv. al
14 al.
Sicily.
Aristano, Cagliari,
46. 6. 15. P.
7. 5. 24. 11-24
Hon. N. C. Rothschild
(P)
Sardinia. (C.
Erausse.)
2 juv. al
Sardinia. (P. Bonomi.
) E. N. Buxton (p).
95. 4. 16. 4-5.
3 9.
Tatoi, Athens, Gi-reece
(C. Mottaz.)
Hon.N. C.Rothschild
(P)-
8. 10. 2. 19-21
6, 3 9.
Kephissia, Athens.
C. Mottaz (c).
8. 11. 3. 4-7.
PIPISTRELLUS 213
PIPISTRELLUS NATHUSII Ke3'.serling and Blasius.
1839. V[espciiilio] naflMsii Keyserling and Blasius, Wiegmann's Archiv
fiir Naturgesch., 1839, i, p. 320 (Berlin, Germany).
1857. Vesperngo nathusli Blasius, Saugethiere Deutschlands, p. 58.
1878. Vesperngo abranms Dobson, Catal. Chiropt. Brit. Mus., p. 226 (Part :
not of Temminck).
1900. Pipistrellus nathusii M6hely, Monogr. Chiropt. Hungariae, p. 276.
1905. Vesp[eritgo] nathusii var. unicolor Fatio, Arch. Sci. Phys. et Nat.,
Geneve, 4th ser., six, p. 510, May, 1905 (Geneva, Switzerland).
Type in Geneva Jluseum.
1910. Pipistrellus ahramus Trouessart, Faune Mamm. d'Europe, p. 16.
Type locality. — Berlin, Germany.
Geographical distribution. — Central and southern Continental
Europe ; exact limits of range not known.
Diagnosis. — Not so small as PipistreUus lApistrellus (forearm,
32 to 3-5 mm. ; condylobasal length of .skull, 12 '6 to 13 "4 mm.),
which in general it resembles, but : small upper premolar better
developed, the greatest diameter of its crown nearly half that of
canine ; canines both above and below much more slender, the
length of base of lower tooth measured along cingulum slightly
more than half length of anterior border ; tragus more slender,
its greatest width much less than length of anterior border ;
thumb long, its length much greater than width of wrist ; length
of fifth finger about 46 mm. ; posterior edge of wing membrane
always pale, though never sharply defined white.
E.dernal characters. — In general and apart from the animal's
less diminutive size, the external characters are essentially as in
Plj}i.^treUus pipistrellus. Ear larger and broader with more
obtuse apex, more evadently concave posterior border, the inner
surface of conch more rugose and with about four irregular cross
striations behind tragus ; antitragus small but well defined,
projecting distinctly beyond border of conch ; tragus about as
high as in P. pipistrellus and similarly blunt . at tip, but with
posterior border more evidently convex ; posterior basal lobe
small, usually le.ss well defined than in P. 2}!pistreUus. Wing
larger and relatively broader than in P. pipistrellus, the meta-
carpals as in the smaller animal, but fifth finger extending beyond
elbow to a distance equal to decidedly more than one-third length
of forearm ; thumb less shortened than in the other European
members of the genus, its length noticeably greater than width
of wrist ; membrane inserted at base of outer toe. Foot, calcar
and tail as in P. pipistrellus.
Fur and colour. — Fur slightly more loose in texture than that
of P. jripistrelhis, the individual hairs somewhat longer, those at
middle of back about 7 mm. in length. In distribution it shows
no peculiarities, though it extends perhaps less widely on dorsal
surface of interfemoral membrane. Colour essentially like that
of Pipistrellus pipistrellus, though usually distinguishable by a
214
CHIROPTERA
tendency aAvay from the cinnamon and raw-umber tints toward
a clearer brown more resembling Eidgway's mars-brown. Mem-
branes less blackish than in P. pipistrdlus, the wing from foot
nearly to fifth finger with a noticeable pale border about 1 mm.
in width, similar to that present in P. kultlii, but less sharply
defined and less nearly white.
Skull. — The skull is less diminutive than that of Pipistrellus
pipistrellus, its general size about as Myotis mystnciuus. General
form less robust than in P. pijristreUus, the width of brain-case
barely one-half greatest length, but more contrasted with that of
rostrum. Dorsal profile as in the smaller sjaecies, but with more
evident anterior concavity and posterior convexity, the anterior
edge of interparietal indicated bj'' a slight transverse constriction.
Other details of form essentially as in P. pipistrellus.
7'eetJi. — As compared with those of P'qnstrelhts pipistrellus the
teeth throughout show a tendency toward slenderness and height.
Inner upper incisor noticeabh' more slender than that of P.
pipistrellus, and with less well developed secondary cusp ; outer
upper incisor distinctly larger than inner, its apex extending
noticeably beyond secondary cusp of inner tooth, its general form
essentially as in P. pipistrellus, but inner margin without evident
secondary cusp. Lower incisors less crowded than in P. j^iipia-
trdlus, a slight space usually present in
median line, another between /^ and i^, and
another between i^ and canine ; outer edge
of /j slightly ovei'lapping i.^ ; in form the
teeth are not peculiar. Upper canine like
that of P. piinstreUus, except that the
greatest diameter of its crown is only about
half length of anterior border of shaft.
Mandibular canine high and slender, its apex
Pipistrcllus nathusii. rising distinctly above that of highest molar
Anterior teeth x 5. cusps, its greatest diameter measured along
cingulum equal to a little more than half
length of anterior border : apex of anterior cingulum cusp not
rising above level of basal third of posterior boi-der. LIpper
premolars as in P. pipistrelhis, except that the small tooth is
relatively higher and more perfectly in the tooth-row, and the
posterior border of its shaft usually shows some indication of an
angular secondary cusp corresponding to that of canine. Lower
premolars with crown area more nearly equal than in the
smaller species, but without special peculiarities of form. Molars
both above and below essentially similar to those of P. piplstreUus.
Measurements. — Adult male from Berlin, Germany (topotype) :
head and body, 45 ; tail, 35-4 ; tibia, 13 ; foot, 6*8 ; forearm,
33; thumb, 5*2; third finger, 65; fifth finger, 47; ear from
meatus, 12; width of ear, 11. Average and extremes of six
adults from Buchillon, Vaud, Switzerland : tibia, 1 2 • 9 ( 1 2 ■ 6-14) ;
foot, 7-3 (6 -8-7 -6); forearm, 33 (32 • 4-34 -G); thumb, 5-7
PIPISTRELLUS 215
(5-2-6); third finger, 61-3 (58-65); fifth finger, 44-6 (41-47).
Adult male and female from Florence, Italy : head and body,
46 and 47; tail, 40 and 38; tibia, 12-8 and 12-6; foot, 6-8
and 7 ; forearm, 35 and 33 ; thumb, 6 ■ 2 and 6 • 6 ; third finger,
63 and 62; fifth finger, 47 and 46; ear from meatus, 12-6 and
12-6; width of ear, 1 1 and 11-4. For cranial measurements see
Table, p. 222.
Specimens examined.— Thinx-thxee, from the following localities :—
Feance : St. Gilles, Gard, 1.
Geemany: Berlin, 1; Bavaria, 1 (U.S.N.M.) ; Ingelheim, Ehein-
hesseu, 1.
Switzerland: Geneva, 6 (Mottaz and Geneva, including type of
unicolor Fatio) ; Montreux, Vaud, 1 (Mottaz) ; Buchillon, Vaud, 6 (Mottaz) •
Neuchatel, 1 (U.S.N.M.) ; Canton Uri, 1 ; St. Gothard, Uri, 1 (U.S.N.M.).
Austkia-Hungary : Palics, Bacser, southern Hungary, 2.
Italy : Siena, 1 (U.S.N.M.) ; Florence, 3 (U.S.N.M.) ; Rome, 5 (B.M.
and U.S.N.M.) ; Borzoli, Liguria, 1 ; Catanzaro, Calabria, 1 (U.S.N.M.).
Bemarks. — Though readily distinguishable from the other
European members of the genus by its cranial and dental
characters, Pipistrelliis nathusii is superficially muoh like P. 2J'pis-
trcllus. It is usually recognizable, however, by its slightly less
diminutive size, more robust form, and by the constant presence
of an ill-defined light (though never actually whitish) border to
the wing.* As pointed out by Mehely in 1900 it has no very
near relationship to the Oriental P. abramus.
6. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4. 128.
6 al. Berlin, Germany. Dr. Giinther (c & p). 66. 2. 1. 22.
?. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 3.
2 al. Palics, Bacser, Hungary. Budapest :Museum (e). 0. 4. 9. 1-2.
S. Canton Uri, Switzerland. Tomes Collection. 7. 1. 1. 398.
6,9. Rome. {C. Coll.) G. Barrett-Hamilton (p). 11.1.2.20-21.
9. Borzoli, Liguria. Italy. Marquis G. Doria (c & p). 5. 12. 15. 7.
PIPISTRELLUS KUHLII Kuhl.
1819. Vespertilio kuhlii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
{- Neue Ann., i), pt. 2, p. 199 (Triest).
1835. Yesp[ertiUo] albolimbatiis Kiister, Isis, p. 75 (Cagliari, Sardinia).
1837. Vespertilio vispistrellus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx
(near Rome, Italy). Type in British Museum.
1837. Vespertilio alcythoc Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Sicily.) Type in British Museum.
1838. Romicia calcarata Gray, Mag. Zool. and Bot., ii, p. 495 (locality
unknown).
1841. Pipistrellus jnarginatus Bonaparte, Iconogr. Fauna. Ital., Indie.
distrib., nomencl. mod. (Substitute for albolivibatus).
1844. Vespertilio 7narginatus Wagner, Schreber's Siiugthiere, Suppl., i,
p. 503, pi. Lv A. No description. Name occurs in synonymy of
kuhlii with ^Michahelles as authority, and on plate. Apparently
not previously published.
* A light border sometimes occurs in P. pipistrellus, but is rare. That
constantly present in P. kuhlii is more sharply defined and more truly
whitish than in P. nathusii.
216
CHIROPTERA
1840. V[es2)ertilio] Ursula Wagner, Schreber's Saugthiere, SuppL, i, p. 505
(Morea, Greece).
1857. Vcsperugo kuhlii Blasius, Saugethiere Deutscblands, p. 63.
1878. Vesperugo kuhlii Dobson, Catal. Cbiropt. Brit. Mus., p. 230.
1886. [Vesperugo kuhlii] var. albicans Monticelli, Atti Soc. Ital. Sci. Nat.,
Milano, xxvii, p. 200, Marcb, 1886 (Caivano, Naples, Italy).
1886. [Vesperugo kuJilii'} v&t . imllatiis Monticelli, Atti Soc. Ital. Sci. Nat.,
Milano, xxvii, p. 200, March, 1886 (Bella Vista, near Portici,
Naples, Italy).
1900. Pipistrellus kuhlii M6hely, Monogr. Cbiropt. Hungarise, p. 261.
1910. Pipistrellus kuhli Trouessart, Faune Mamm. d'Europe, p. 17.
Type locality. — Trieste, Austria-Hungary.
Geographical distribution. — Mediterranean region and eastward
into Asia.
Diatjnoms. — Size about as in Pipistrellus natlmsii (forearm, 31
to 35 mm.; condylobasal length of skull, 12-0 to 13-2 mm.);
outer upper incisor less than half as high as inner incisor ; large
upper premolar almost or (}uite in contact with canine, the small
premolar forced inward from tooth-row and scarcely or not visible
from outer side, its greatest diameter about equal to that of
outer incisor ; canines less robust than in P. pipistrellHs, less
slender than in P.natluisii ; tragus with greatest width less than
length of anterior border ; thumb short, its length about equal to
width of wrist ; posterior edge of wing membrane with sharply
defined whitish border.
External eharacters. — General form very similar to that of
Pipistrellus pipistrellus, the wing similarly narrow as compared
with that of P. natlmsii. Ear narrowly rounded at tip, the posterior
border slightly concave ; antitragus slightly developed, producing
an evident break in contour of conch ; inner surface of conch
somewhat rugose, without well defined transverse striations ;
tragus essentially as in P. natlmsii, the posterior border noticeably
convex. Wing, foot, calcar and tail as in P. p>ip)istrellus.
Fur and colour. — Quality and distribution of fur essentially
as in Pipistrellus pipjistrellus, but dorsal surface of interfemoral
membrane haired scarcely beyond basal third. Colour not very
different from that of Pipistrellus pipistrellus, but somewhat
lighter and more yellow, often approaching raw-siena. Mem-
branes blackish, the wing between foot and fifth finger with a
sharply defined very narrow (less than 1 mm.) nearly white
border.
Skull. — The skull resembles that of Pipistrellus natlmsii in
size, but its form is even more robust than that of P. pipnstrellus.
Dorsal profile with very slight interorbital concavity and barely
perceptible convexity over middle of brain-case. Breadth of
brain-case about half greatest length of skull. Dorsal surface
of rostrum less rounded off" at sides than in P. pipistrellus and
P. natlmsii, but not .so much flattened as in P. savii. Narial
emargination more abruptly narrowed posteriorly than in the
PIPISTEELLUS
217
other European species. Mesopterygoid space slightly longer
than wide. Mandible with coronoid process distinctly higher
than articular process, so that upper edge of posterior portion is
oblique and not pai'allel with alveolar line.
Teeth. — Inner upper incisor essentially as in PipistreUus
yi pistrellus except that secondary cusp is reduced to a minute,
sometimes obsolete, projection from cingulum at posterior base of
shaft ; outer incisor very small, less than half as high as inner,
its apex about on level with highest point of cingulum of larger
tooth, its crown area about two-thirds that of latter, its structure
essentially as in P. pipistreUus, though with very small secondary
cusp ; small tooth situated directly exterior to
large, so that a line perpendicular to main
axis of skull would pass through middle of
all four incisors ; lower incisors essentially as
in P. pipistrellus, though relatively larger and
more strongly imbricated. Canines both abo^e
and below intermediate in form between those
of P. lAjjistrellus and P. nathusii. Anterior
upper premolar crowded inward from tooth- 1/ y,M -ry
row and closely wedged between canine and
large premolar which are nearly or quite in pipistrTiii^'kuhm.
contact ; the small tooth is usually though not Anterior teeth x .'>".
always invisible from outer side, its cusp is
very low, nearly terete, its crown area about equal to that
of outer incisor ; large premolar with no special peculiarities.
Lower premolars as in the related species, but disproportion in
size more marked, the crown area of first a little more than
half that of second. Molars both above and below essentially
as in P. piinstrelhis and P. natliusii, but somewhat more robust.
Measiiremeiifs. — Two adult males from near Genoa, Italy :
head and body, 43 and 46 ; tail, 37 "4 and 38 ; tibia, 12-4 and
13 ; foot, 6 and 6 ; forearm., 33-6 and 34 ; thumb, 5 and 5-2 ;
third finger, 60 and 60 ; fifth finger, 45 and 44 ; ear from meatus,
12 6 and 13; width of ear, 10-4 and 10. Two adult females
from the same locality : head and body, 44 and 47 ; tail, 39 and
40 ; tibia, 1 2 and 13 ; foot, 6 ■ 2 and 6 • 2 ; forearm, 33 • 4 and 35 ;
thumb, 5-2 and 5-4; third finger, 60 and 63 ; fifth finger, 45
and 47 ; ear from meatus, 13 and 13 ; width of ear, 10 and 10.
Adult male and female from Palermo, Sicily : head and body,
42 and 44 ; tail, 36 and 35-4 : tibia, 12-4 and 12-4 ; foot, 5-6
and 5 • 4 ; forearm, 3 1 and 34 ; thumb, 5 • 4 and 5 • 2 ; third finger,
56 and 61 ; fifth finger, 41 and 43 -6 ; ear from meatus, 12-4 and
13 ; width of ear, 10 and 10. Adult male and female from
Cagliari, Sardinia : head and body, 44 and 45 ; tail, 35 and 37
tibia, 12-6 and 12-4 ; foot, 6-2 and 6-8 ; forearm, 33 and 33
thumb, 5 and 5 ; third finger, 58 and 62 ; fifth finger, 43 and 45
ear from meatus, 13 and 12 ; width of ear, 10 and 10. For
cranial measurements see Table, p. 218.
218
CHIROPTERA
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Balearic Islands : San Cristol
France : St. Gilles, Gard
Italy : Perti, Finalborgo
Siena
Kome
Sorrento .
Catans^aro, Calabria
Sicily : Palermo .
Ustica Island .
Sardinia: Wt. Gennargcntu
Greece : Corfu
Argostoli, Cephaloni
PIPISTRELLUS 219
Specimens examined. — One hundred and eight, from the followino
localities : — "^
Feaxce: St. GiUes, Gard, 2; Nimes, Gard, 3 (Mottaz) ; St. Genies
Gard, 4 (Mottaz) ; Marseilles, 1 (U.S.N.M.).
Switzerland: Coremmo, Ticino, 1 (Mottaz) ; Lugano, Tioino, 1 (Mottaz).
Italy; Near Genoa, 14 (U.S.N.M. and Genoa); Siena, 6 (B.M. and
U.S.N.M.); Florence, 1 (U.S.N.M.); Rome, 6 (B.M. and U.S.N.M.);
near Rome 1 (type of vispistrellus Bonaparte) ; Sorrento, 3 (U.S.N.M.) ;'
Catanzaro, Calabria, 6 (U.S.N.M.) ; Palermo, Sicily, 23 (U.S.N.M ) '
Corleone, Sicily, 1 (U.S.N.M.); Ustica Island, Sicily, 4 (U.S.N.M.);
Sicily, 1 (type of alcythoe Bonaparte).
Sardinia: Cagliari, 4 (U.S.N.M.); Mt. Gennargentu, 7 (U.S.N.M.);
no exact locality, 2.
Greece : Corfu, 2 ; Cephalonia, 10 ; Patras, 3.
Spain : San Cristobal, Minorca, Balearic Islands, 2.
Bemarks. — PipistreUus huhlii is easily recognizable by the
form and relative size of the upper incisors. Externally it may
usually be known by the sharply defined whiti.sh border to the
wing membrane, though too much reliance should not be placed
on this character alone. Many specimens from Sardinia are
lighter in colour than those from the mainland. These represent
the albolimhatus of Kiister. Normally coloured examples also
occur ; and in the absence of adequate material it has seemed
preferable for the time being not to attempt to define the insular
form.
6. St. GiUes, Gard, G. S. MiUer (c). 8. 8. 4. 127.
France.
<^. Siena, Italy. Dr. E. Hamilton (p). 98. 10. 2. 2.
(S. Brogi.)
3 6, ?. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. IG-IP
(P)-
skeleton with- Sicily. {Prince Bona- Tomes Collection. 7. 1. 1. 780.
out skull. 2^a'-tc.) {Type of V. alcythoe Bonaparte.)
skeleton with- Near Rome, Italy. Tomes Collection. 7. 1. 1. 7"29.
out skuU. {Prince Bonaparte.) {Type oi F. vispisfeZ^MS Bonaparte.)
i, 9. Corfu, Greece. J. I. S. Whitaker (p). 8. 10. 1. 4-5.
(C. Mottaz.)
3 <;. Argostoli, Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 1-:3.
(C. Mottaz.)
6, 9. Patras. {C. Mottaz.) Hon. N. C. Rothschild 8. 10. 2. 17-18.
2 c?. San Cristobal, O. Thomas and R. I. 0.7.1.29-30.
Minorca, Balearic Pocock (c & p).
Islands.
PIPISTRELLUS SAVII Bonaparte.
1837. Vespertilio savii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx (Pisa)
Type in British Museum.
1837. Vespertilio aristippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Sicily).
1837. Vespertilio leucippe Bonaparte, Iconogr. Faun. Ital., i,. fasc. xxi
(Sicily). Type in British Museum.
1833. Vespertilio bonapartii Savi, Nuovo Giorn. de' Lettcrati, Pisa, sxxvii
p. 226 (Tuscany).
220
CHIROPTEEA
18i4. Vesp[ertilio] nigrans Crespon, Faune Meridionale, i, p. 24 (Nimes,
Gard, France).
1853. Vesperugo manrus Blasius, Wiegmann's Arcliiv fiir Naturgesch.,
1853, I, p. 35 (Central chain of the Alps).
1857. Vesperugo maurus Blasius, Siiugethiere Deutschlands, p. 67.
1872. V[espertilio] agilis Fatio, Faune Vert. Suisse, Append, au vol. i,
p. iii (Alternative name for V. savii Bonaparte, ex Savi MS.).
1878. Vesperugo maurus Dobson, Gatal. Chiropt. Brit. Mus., p. 218.
1904. Vespertilio ochromixtus Cabrera, Mem. Soc. Espaii. Hist. Nat., ii,
p. 267 (Sierra de Guadarrama, Madrid, Spain).
1910. PipistreUus savii and P. savii oclironiixtus Trouessart, Faune Mamm.
d'Europe, pp. 13-14.
Tiipc locality. — Pisa, Italy.
Geographical distribution. — Southern Europe, west to the
Iberian Peninsula, north to the Alps ; also the Canary Islands,
northern Africa and southern Asia. Limits of range very
imperfectly known.
Diagnosis. — Largest European member of the genus (condy-
lobasal length of skull, 13 to 14 mm. ; forearm, 31 to 33 mm.) ;
outer upper incisor more than half as high as inner ; large
premolar broadly in contact with canine, the small tooth very
minute, crowded inward from axis of tooth-row, invisible from
outer side and occasionally covered by the gum, its diameter
much less than that of outer incisor ; anterior lower premolar
with crown area less than half that of succeeding tooth ; lower
canine robust ; tragus with greatest width nearly equal to length
of anterior border ; thumb short ; hairs of back usually with
contrasting light brown tips.
External characters. — Ear broad, its general form about as in
PipistreUus nathusii, the posterior border slightly but evidently
concave above middle, the inner surface of conch noticeably
rugose and with faint, irregular transverse ridges behind tragus ;
antitragus small aiid ill-defined, but producing an evident break
in outline of conch ; tragus less than half as high as conch, very
wide (greatest width, at level of middle of anterior border, nearly
equal to length of anterior border), the anterior border nearly
straight, the posterior border strongly and evenly convex from
tip to notch above small basal lobe. Wing, foot, calcar and tail
as in P. pipistrellus.
Fur and colour. — The fur resembles that of Piplstrellus Tcuhlil
in quality and distribution. Colour differing from that of the
other European species in the evident contrast between light
tips of hairs of back and darker ground tint. It is also the only
species in which there is much individual variation in colour.
Four specimens from the neighbourhood of Genoa are coloured as
follows : male, not fully adult, uniform very dark vandyke-brown,
the extreme tips of hairs of back faintly lighter, underparts a
light brown faintly overlaid on blackish under colour ; adult male :
light tips on back well developed, giving general colour to region,
between raw-umber and clay-colour ; adult male : light tips very
PIPISTRELLUS 221
conspicuous, a peculiar dull brownish ochraceous-bufF; adult female :
light tips as conspicuous as in last, dull brownish cream-buft".
SJatll. — The skull is slightly larger than that of Pipistrellus
nafhiisii and P. kuhVu, and is immediately distinguishable among
the European species by the flatness of dorsal surface of rostrum,
prominence of ridge along edge of orbit, and
relatively small size of narial emargination,
characters the first two of which suggest
Verpcrtilio murinus. Dorsal profile of skull as
in P. jiipistrellus and P. Icnldii, but brain-case
slightly more depressed. Breadth of brain-case
about half greatest length of skull. Rostrum
relatively broader than in the other European
species, its dorsal surface more flattened and
orbital ridges more prominent ; narial emar-
gination scarcely larger than in P. pipistrellus.
Mesopterygoid space about as wide as long, its „• V",,""
general outline, aside irom the notch caused Nat. size,
by median spine of palate, broadly barrel shape,
the hamulars distinctly turned inward. Mandible with coronoid
process and upper border of posterior portion as in P. hihlii,
but with angular process less curved and relatively longer than
in the other European species.
Tt'eth. — Incisors both above and below essentially as in
P/jjitifrellus pipistrellus, except that inner upper tooth has the
secondary cusp somewhat better developed.
Canines with no .sj^ecial peculiarities, not
essentially different from those of P. pipi-
strellus. Anterior upper premolar very
minute, sometimes hidden in the gum or
occasionally absent, its crown area never
much more one-sixth that of outer incisor ;
large premolar always strongly in contact
. .. with canine, its form peculiar in the absence
Anterior teeth, x's. Or slight development of the antero-internal
cusp. Lower premolars strongly contrasted
in size, the crown area of first decidedly less than half that
of second, its cusp relatively lower and less developed than in
anj' of the other European species. Molars both above and
below with no special peculiarities.
Measurements. — Two adult males from Palermo, Sicily : head
and body, 43 and 47 ; tail, 34 and 35; tibia, 12-6 and 12-8;
foot, 6-4 and 7; forearm, 31 and 32-6; thumb. Sand 5-6;
third finger, 54 and 56 ; fifth finger, 42 and 41 ; ear from meatus,
12-4 and 12-6; width of ear, 'l2 and 12. Two adult females
from the same locality : head and body, 46 and 47 ; tail, 35 and
39 ; tibia, 13 and 13-4 ; foot, 6-6 and 7 ; forearm, 33 and 33 ;
thumb, 5 and 5-4 ; third finger, 56 and 57 ; fifth finger, 42 and
43 ; ear from meatus, 12-4 and 13 ; width of ear, 12 and 11-6.
CHIROPTEEA
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224
CHIROPTEKA
Adult male from Escorial, Spain (paratype of ochromi.vtus Ca.hrern) :
head and body, -iS-G ; tail, 34 ; tibia, 13-2; foot, 5-6; forearm,
57 ; thumb, 5 "4; third finger, 43 ; ear from meatus, 13 ; width
of ear, 11. For cranial measurements see Table, p. 223.
Specimens examined. — Twenty-four, from the following localities : —
Switzerland : St. Gothard, Uri, 1 ; no exact locality, 1.
Italy: Near Genoa, 10 (B.M. and Genoa); Florence, 3 (Mottaz) ;
Sorrento, 1 (U.S.ISr.M.) ; no exact locality, 1 (type) ; Palermo, Sicily, 8
(U.S.N.^I.); Sicilv, no exact locality, 1 (type of leucippe Bonaparte);
Ustica Island, Sicily, 1 (U.S.N.M.).
FRA^'CE : Near Nimes, Gard, 2 (U.S.N.M. and Nimes ; the latter
agreeing with description of nigrans Crespon, though not marked type) ;
St. Gilles, Gard, 1 ; no exact locality, 1.
Spain: El Escorial, iladrid, 1 (paratype of ochromixtus Cabrera).
Be marks. — This species is readily distinguishable among the
European members of the genus Plpi-s-freUus by the peculiar form
of the tragus, apart from its very pronounced cranial and dental
characters. Its colour gives it a superficial resemblance to
Ejjtesicus nilssoni, a likeness that is so heightened by the exces-
sively small size of the anterior upper premolar that the animal
has been once and pierhaps twice described as a member of the
genus EptesicKs*
1. St. Gothard, Uri, Swit- Tomes Collection. 7. 1. 1. 397.
zerland.
Switzerland. Purchased (Brandt). 45. 11. 1. 3.
4 al. Genoa, Liguria, Italy. Genoa ^Museum (e). 86.11.3.14-
17.
i. Borzoli, Liguria. !Marquis G. Doria 5. 12. 15. 6.
(c & P).
skeleton with- Italy. {Prince Bona- Tomes Collection. 7. 1. 1. 7-32.
out skuU. parte.) [Type of species.)
skeleton with- Italy. [Prince Bona- Tomes Collection. 7. 1. 1. 731.
out skull. parte.) (Type of F. Z<;«cy)pt' Bonaparte.)
? al. France. Purchased (Lefebre). 46.1.2.12.
6. El Escorial, Madrid, A. Cabrera (p). 8. 7. 23. 1.
Spain. (Paratype of Y. ochromixtus Cabrera.)
Genus EPTESICUS Ratinesque.
1820. Eptesiciis Rafinesque, Annals of Nature, p. 2 [melanops = fitscus).
1S29. Cnephxus Kaup, Entw.-CTesch. u. Natiirl Syst. Europ., Thierwelt, i,
p. 103 [serotinus).
1839. Vespcrugo Keyserling and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, i, p. 312 (part).
1839. Vesp>crus Keyserling and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, i, p. 313 (Sub-genus of Vesperugo, part). Not Yesperns
Latreille, 1829.
* This is certainly the case with the Vespertilio ochromixtus of Cabrera.
In the paratype of this species (B.M. no. 8. 7. 23. 1), which I carefully
examined before removal of the skull, in company with Mr. Knud Andersen,
no trace of the small premolar could be found. Wbeu the skull was cleaned,
however, the presence of the tooth in its normal position was revealed,
thus showing the animal's true identity. It seems not improbable, so far
as can be judged from the original description, that Satunin's Ycspcrugo
caucasicus (Zool. Anzeiger, xxiv, p. 462, August 5, 1901) was based on
similar specimens.
EPTESICUS
225
I
1841. JSoctula Bonaparte, Iconogr. Faun. ItaL, i, fasc. xxi, in account of
Vespcrtilio alcijthoc {serotinus).
1856. Cateonis Kolenati, Allgeni. deutsche Naturhist. Zeituug, Dresden,
neue Folge, ii, p. 131 (serotinus).
1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 (part).
1857. Vesperus Blasius, Saugethiere Deutschlands, p. 51 (Sub-genus of
VesjJcrugo), part.
1858. Amblyotiis Kolenati, Sitzuugsber. kais. Akad. Wissensch. Wien,
Math.-Naturwissensch. Classe, xxix, p. 252 (atratiis — nilssoni).
1863. Aristijipc " Kolenati, Beitriige zur Kenntniss der Phthiriomyiarien,
Petersburg, 1803" (part, included both discolor = vmrinus and
7iilsso?ii).
1866. Pachijomus Gray, Ann. and Mag. Kat. Hist., 3rd ser., xvii, p. 90,
February, 1863 (ixiclii/omus).
1870. Nyctiptenus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 424 (smithii).
1878. Tesj^enis Dobsou, Catal. Chiropt. Brit. Mus., p. 184 (Sub-genus of
Vespcriigo), part.
1892. Adelomjcteris H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1891,
p. 466, January 19, 1892 (part; substitute for Vesperus, pre-
occupied).
1897. Yespcrtilio Miller, Ann. and Mag. Nat. Hist., 0th ser., xx, p. 384,
October, 1897 (part).
1900. Eptesicus Mehely, Monogr. Chiropt. Hungarise, p. 219 (part).
1907. Eptesicus Miller, Families and Genera of Bats, p. 207, June 29, 1907.
Type species. — Eptesicus melanops Rafinesque = Verpertilio
fuscus Beauvoi.s.
Geographical distribution. — Europe, Asia (except Malay region),
Au.stralia, Africa, Madagascar ; America from southern Canada
southward (except Lesser Antilles).
Characters. — Dental formula : / |-^, c J--i, pm j^,, m p", = 32.
Teeth strictly normal throughout, and showing no special pecu-
liarities. Both upper incisors well developed, the inner larger
than the outer and usually with distinct secondary cusp, the
outer separated from canine by a space equal to its greatest
diameter ; m^ variable in form, usually with well developed
metacone and three commissures in the smaller species, but with
raetacone and third commissure obsolete in larger forms. Skull
without special peculiarities of form or structure, the rostrum
tlattish or more usually rounded ofl' above, the nares and palatal
emargination not specially enlarged, the latter at least as deep
as wide. Ear of moderate size, not peculiar in form : wing
broad (normal).
BemarJcs. — Among European bats the members of the genus
Eptesicus may be distinguished by their dental formula combined
with a simple Pij>i.'<trelliis-]\ke ear and not specially modified
skull. The group is nearh' related to PipistrelJus through
P. snvii, in which the small premolar is occasionally absent and
not infreciuently so minute as to be concealed by the gum. About
forty-five species are known, three of which occur in Europe.
226 CHIROPTEEA
KEY TO THE EUBOPEAN SPECIES OF EPTESICUS.
Forearm less than 40 mm. ; coudylobasal length of
skull less than 16 mm. ; a distinct line of demar-
cation between colours of upper and lower surfaces
of neck E. nilssonii, p. 2.34.
Forearm more than 45 mm. ; condj'lobasal length of
skull more than 17 mm. ; no line of demarcation
between colours of upper and lower surfaces of
neck.
Coudylobasal length of skull 19 to 21*6 mm E. serotinus, p. 226.
Condylobasal length of skull about 18 mm E. sodalis, p. 231.
EPTESICUS SEROTINUS Schreber.
1774. Vcspertilio serotinus Schreber, Siiugthiere, i, pi. liii (Description, i,
-p. 167, 1775, under name : Die Blasse Fledermaus). France,
based primarily on "La Serotine " of Daubenton, Hist. Acad.
Royale des Sci., 1759, p. 377. 1765.
1776. Vcspertilio serotine P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 16 (Based on "die Blasse Fledermaus" of Schreber).
1827. Vcspertilio u-iedii Brehm, Ornis, Heft in, p. 24 (Renthendorf,
Thiiringen, Germany).
1827. Vespertilio okcnii Brehm, Ornis, Heft in, p. 25 (Renthendorf,
Thiiringen, Germany).
1844. Vcspjlertiliol incisivus Crespon, Faune M6ridiouale, i, p. 26 (Nimes,
Gard, France).
1857. Vcsperugo serotinus Blasius, Siiugethiere Deutschlands, p. 76.
1863. [Cateorus serotiniis'\ var. typus Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 466 (Wiesbaden, Nassau,
Germany).
1863. \_Cateorus serotinus'] var. rufesccns Koch, Jahrb. des Vereins fiir
Naturkjude im Herzogthum Nassau, xviii, p. 466 (Freiburg,
Breisgau, Germany).
1878. Vesiwrugo serotinus Dobson, Catal. Chiropt. Brit. Mus., p. 191.
1885. Vespierus serotinus var. transylvanus Daday, Orvos-Termeszettudo-
manyi Erteset(5, Kolozsvar, x, p. 275 (Also-Szocs, Szolnok-Doboka,
Hungary).
1886. Vesperus serotinus var. transsylvanus Daday, Verhandl. u. Mittheil-
ungen des Siebenbiirgischen Vereins fiir Naturwissensch. in
Hermannstadt, xxxvi, p. 81.
1900. Eptesic.us serotinus Mehely, Monogr. Chiropt. Hungarise, p. 209.
1904. Vespertilio serotinus insularis Cabrera, Mem. Soc. Espan. Hist. Nat.,
II, p. 263 (Minorca, Balearic Islands).
1904. Vcspicrtilio isnhellinus Cabrera, Mem. Soc. Espan. Hist. Nat., ii,
p. 264 (southern Spain). Not of Temminck.
1904. Vcspertilio hoscai Cabrera, Mem. Soc. Espan. Hist. Nat., ii, p. 265
(Muchamiel, Alicante, Spain).
1910. Eptcsicus serotinus, E. serotinus transsylvanus, and E. boscai Troues-
sart, Faune Mamm. d'Europe, pp. 20-22.
Tj/jye locality. — France.
GeograjMcal distrihution. — Central and southern Europe
from England and Denmark to the Mediterranean ; eastward
into Asia.
Diaynosis. — Size rather large (forearm more than 45 mm.,
EPTESICUS.
227
condylobasal length of skull more than 19 mm.) ; colour of upper
parts a yellowish brown without noticeably contrasted light tips
to the hairs ; under parts essentially similar, so that there is no
line of demarcation along sides of neck.
External characU'rs. — General form robust, though less so
than in Nyctalus, the tail and legs rather short, the membranes
thick and opaque. Muzzle with moderately prominent glandular
swelHngs, its greatest width across this region less than distance
from nostril to ear ; nostrils projecting very slightly, the concavity
between them not conspicuous, the orifice crescentic. Ear
moderately long, extending slightly more than half way from eye
to nostril when laid forward, its breadth when flattened about
equal to height above crown ; anterior border of conch abruptly
convex below, then nearly straight to narrowly rounded olT tip ;
posterior border straight or irregularly concave from just below
tip to level of anterior base, then convex to abrupt angle under
meatus marking posterior limit of small but well defined anti-
tragus, the anterior border of which terminates obscurely about
3 mm. behind angle of mouth ; inner surface of conch obscurely
papillose, the region behind tragus marked by about six faint
cross ridges ; tragus short, its height less than half that of ear
conch, its anterior border straight, its posterior border gently
convex from narrowly rounded tip to upper edge of small but
distinct basal lobe, its greatest width, at level of middle of anterior
border, equal to slightly more than half length of anterior border.
Wing broad, the fifth finger exceeding forearm by one-quarter
to one-third length of forearm, the membrane leathery and
opaque, though perhaps less so than in Nyctahis noctula, joining
leg at base of outer toe ; third and fourth metacarpals sub-equal,
nearly as long as forearm, fifth about 2 mm. shorter. Leg rather
slender, the foot less than half as long as tibia ; calcar about
four-fifths as long as tibia and slightly exceeding length of free
border of uropatagium, its keel ill-detined, its terminal lobe well
developed though small. Tail extending to between shoulders
wlien laid forward, the last vertebra and distal tliird of penulti-
mate vertebra free from membrane.
Far and colour. — Fur soft and dense, the longest hairs on
back about 10 mm. in length, those of underparts shorter. It
is strictly confined to body, only extending as a thin pubescence
on extreme base of membranes and along a narrow line bordering
under surface of forearm ; free edge of uropatagium naked,
(i round colour of upper parts ranging from prouts-brown to a
light wood-brown, the basal portion of the hairs not essentially
different, the tips of the hairs of back behind shoulders ^\•ith
inconspicuous lighter (bufi'y) tips ; underparts slightly paler,
sometimes approaching ochraceous-bufi", but never sufficiently
contrasted to produce a line of demarcation along sides of neck.
Muzzle, cheeks, ears and membranes blackish. The variation
in general colour appears to be strictly individual.
Q 2
228
CHIEOPTERA
SJcuH. — General aspect of skull robust and flattened, with
widely spreading zygomata, but rather narrow brain-case and
rostrum. Dorsal prolile rising gradually from nares to over-
hanging lambda, essentially straight throughout, though with
slight concavity over lachrymal region. Ventral profile very
slightly elevated posteriorly. Brain-case ovate in general outline,
narrower than in Ni/ctahis noctula, the straight, well developed
lambdoid crests which form its posterior border meeting in median
line almost at right angles, depth at middle about half mastoid
breadth ; sagittal crest low but evident, the region at each side
of it not depressed ; floor of brain-case smooth, with no evident
ridges or depressions, a very narrow slit between cochlea and
basioccipital ; auditory bulhe small, the transverse diameter con-
siderably less than distance between bulla?. Interorbital region
moderately constricted, hour-glass shaped, the lachrymal region
decidedly less wide than brain-case,
with slight tubercular projection close
to anterior rim of orbit ; rostrum
flattened, with shallow but evident
lateral concavity on each side, dis-
tinctly narrower anteriorly than pos-
teriorly, the narrowly obovate narial
emargination extending about half
way back to level of lachi'ymal fora-
men ; rostral depth at front of orbit
less than distance from orbit to outer
incisor ; anteorbital foramen less re-
duced than in Ni/ctalus noctula, its
posterior border over region of con-
tact between large premolar and first
molar, lachrymal foramen directly
behind it, on inner side of orbital
rim. Palate long and narrow, slightly concave both laterally and
longitudinally, the anterior emargination small, squarish, extend-
ing back to level of middle of canine. Posterior extension of
palate nearly parallel sided, though narrowing a little posteriorly,
its width at level of posterioi' molar considerably less than its
length ; hamulars slightly turned inward ; median spine well
developed. Mandible robust, but with lower border nearly
parallel to alveolar line ; posterior portion high in front, low
behind, the height of coronoid process above level of alveolar
line about equal to horizontal diameter, the upper border sloping
abruptly from coronoid to articular process ; angular process
moderately long, about on level with alveolar line, its main axis
directed gradually outward and downward, its distal extremity
slightly expanded and hooked upward.
Teeth. — Relatively to size of skull the teeth are large and
robust, rather more so than in Nyctalini noctula. Inner upper
incisor about half as high as canine, its crown area about one-
Fia. 40.
Eptesicus serotinus.
EPTESICUS
229
quarter that of canine, the subterete shaft directed inward and
slightly forward, its secondary cusp large and prominent, situated
on outer side of shaft near tip ; ciugulum well developed, but
without CUSJ3S. Outer upper incisor much shorter than inner
and with about half its crown area, the apex of its shaft slightly
exceeding level of cingulum of larger tooth ; outer and posterior
surfaces flattened or double-concave, the two concavities some-
times separated by a low but evident ridge ; inner margin with
a low secondary cusp on well developed cingulum. A line
perpendicular to main axis of skull would pass through centre
of shafts of all four incisors ; the outer tooth is separated from
canine by a space about equal to breadth of its own crown. Lower
incisors large, much crowded, and very conspicuously imbricated,
i\ and ?.j overlajjping more than half of front surface of the
succeeding tooth, the general outline of the entire series V-shaped ;
front surface of crowns about as high as wide, the edge obliquely
trifid (occasionally a low, rudimentary fourth cusp at outer
margin of i.^) ; cross section of crown somewhat triangular, the
posterior angle occupied by a low tubercle in i., and ^'3. Upper
canine large, its shaft decidedly the highest of the upper series,
its cross section sub-triangular, the posterior cutting edge well
developed, the anterior less trenchant than in Nyctalns noctula ;
inner surface divided by a low ridge into two shallow con-
cavities, the posterior of which is the larger ; antero-outer surface
convex ; postero-outer surface with deep longitudinal groove ;
cingulum narrow but complete, without cusps. Lower canine
very robust, the diameter of crown noticeably greater than least
distance between canines, the shaft decidedly higher than main
cusps of molars, smoothly rounded in front, flattened-concave
behind and on inner side, the cingulum narrow but complete
except at point of contact with jj, where it becomes abruptly
obsolete, terminating in a slight tubercle corresponding to the
cusp present in Ni/ctalus and Pipistrellus. Upper j^remolar with
crown area about ecjual to that of canine and about two-thirds
that of first molar, its main cusp robust, nearly as long as canine
(measured along cingulum), sharply trenchant posteriorly, flcxt-
tened-concave on inner side, a well developed external and antero-
internal longitudinal groove ; crown with slight anterior and
more marketl posterior emargination, the inner side narrow, with
slight concave crushing surface and well developed cingulum,
which rises to a small cusp anteriorly. Lower premolars closely
crowded, the crown area of first about half that of canine, that
of second nearly four-fifths that of canine ; cusp of first a little
more than half as high as second, which slightly exceeds main
cusps of molars ; cingulum of each tooth well developed, tending
to form a slight postero-internal tubercle. First and second
upper molars sub-equal in crown area, the second wider but more
coustricted at middle ; protocone robust, not very high ; no true
hypocone, but region which it would occupy indicated by slight
230
CHIEOPTERA
columnar thickening of posterior base of protocone ; paracone
lower and smaller than metacone, the contrast unusually notice-
able ; styles and commissures well developed, the W-pattern
normal ; m'' with crown area less than half that of 711^, its longi-
tudinal diameter through metacone much less than half transverse
diameter, the mesostyle, metacone, and second and third com-
missures greatly reduced, though not sufficiently to lose their
identity. Lower molars with no special peculiarities ; angles in
commissures between outer and inner cusps rather wide and
shallow, especially that between protoconid and metaconid ; area
of second V in "'3 scarcely half that of first.
Measurements. — Adult female from Herrnhut, Saxony : head
and body, 62 ; tail, 54 (its free tip, 6-6) ; tibia, 20 ; foot, 10-4 ;
forearm, 50-4; thumb, 9 ; third finger, 84 : fifth ringer, 63; ear
from meatus, 18; width of ear, 15. Adult male from Barsac,
Gironde, France : head and body, 69 ; tail, 54 (free tip, 7 ■ 6) ;
tibia, 22 ; foot, 11 -4 ; forearm, 51 ; thumb, 9 ; third finger, 91 ;
fifth finger, 67 ; ear from meatus, 18 '4 ; width of ear, 16. Adult
male and female from Seville, Spain : head and body, 67 and 64 ;
tail, 46 and 51 ; tibia, 21 -6 and 22-4 ; foot, 11 and 10'4 ; fore-
arm, 49 and 50 ; thumb, 9 • 2 and 8 • 4 ; third finger, 89 and 90 ;
fifth finger, 64 and 68; ear from meatus, 17 '6 and 18. Adult
male and female from Rome, Italy : head and body, 72 and 73 ;
tail, 54 and 54 ; tibia, 21-4 and 21 ; foot, 11 and 11 -4 ; fore-
arm, 51-6 and 53 ; thumb, 9 • 6 and 9 ■ 8 ; third finger, 91 and 92 ;
fifth finger, 66 and 67 ; ear from meatus, 19 and 19 ; width of
ear, 15 '6 and 15 -4. For cranial measurements see Table,
p. 232.
Specimens examined. — Seventy-eight, from the following localities: —
England: Kenley, Surrey, 1 ; Hawkhurst, Kent, 1; Whitstable, Kent, 2 ;
Wingbam, Dover, Kent, 1 ; Yalding, Kent, 5 (B.M. and U.S.N.JI.) ; Isle
of Wight, 5.
France : Barsac, Gironde, 1 (U.S.N.M.) ; near Nimes, Gard, 2 (Nimes ;
type of inc'isivus Crespon, and a specimen wrongly marked type of palusiris
Crespon).
Geemany : Ingelheim, Rheinhessen, 2 ; Magdeburg, Saxony, 2 (B.M.
and U.S.N.M.); Herrnhut, Saxony, 1 (U.S.N.M.); Strass, near Burgheim,
Bavaria, 3; Bavaria, no exact locality, 2 (U.S.N.jNI.) ; Tubingen, Silesia, 2.
Austria-Hungary : Moravia, 1 ; Csallokoz-Somorja, Pressbnrg, Hun-
gary, 1 ; Budapest, Hungary, 2 ; Transylvania, 5 ; Zara, Dahuatia. 2.
Boumania : Bustenari, Prahova, 3.
Greece: Patras, 14 ; near Athens, 1.
Italy : Siena, 3 (B.M. and U.S.N.M.) ; Florence, 2 (B.M. and U.S.N.M.) ;
Vallombrosa,! (U.S.N.M.); Volterra, 1 (U.S.N.M.) ; Rome, 3 ; San Martino
al Cimino, Rome, 1 (Genoa) ; Ustica Island, Sicily, 1 (U.S.N.^I.).
Sardinia : Cagliari, 2 (Genoa).
Spain : Pajares, Leon, 1 ; Seville, 3 ; Muchamiel, Alicante, 1 (Madrid ;
type of boscai Cabrera).
Bemarks. — Among the bats of Europe this species is recogniz-
able by its rather large size, noticeable free tip to the tail,
moderately long, narrow ear, and straight, erect tragus. With
the material now available for study it seems impossible to
EPTESICUS
231
distinguish any local geographical forms. Specimens from Seville
representing the isabelUnus of Cabrera I am unable to separate
from true serotlHuii ; the type of huscai Cabrera is a young of the
same animal ; insularis I have not seen, but there is nothing in
the original description to indicate that it is distinct.*
2 9.
$al.
1 St.
3 9.
9al.
2.
1.
1.
6,9.
2 J, 9.
2al.
lal.
2.
2al.
5 6 a.].
6, 9.
6, 9.
1 9.
9.
9.
1 al.
6.
9.
9al.
lal.
1.
Kenley, Surrey, England.
Whitstable, Kent.
Wingham, Kent.
Yalding, Kent.
Yalding, Kent.
Isle of Wight.
Freshwater, Isle of Wight.
Freshwater, Isle of Wight.
(F. Bond.)
Bembridge, Isle of Wight.
Ingelheim, Bheinhessen,
Germany.
Magdeburg, Saxony.
(Wolterstorff.)
Strass, Burgheim, Bavaria.
(Korbitz.)
Tiibingen, Silesia.
Moravia, Hungary.
Csallokoz-Somorja, Press-
burg.
Budapest.
Transylvania.
Zara.Dalmatia. (K.Blos.)
Bustenari, Prahova, 840 m.
Roumania. ( W. Dodson.)
Patras, Greece. {C.Motfaz.)
Patras.
Athena. (C. Mottaz.)
Florence, Italy.
Rome. (C. CoU.)
Pajires, Leon,
(N. Gonzalez.)
Seville.
Seville.
Europe.
Spain.
W. R. Ogilvie-Grant
(c & p).
C. H. B. Grant (c & p).
G. Donker (c & p).
H. Reid (c & p).
W. R. Ogilvie-Grant
(c & p).
Rev. C. Bury (c & p).
F. Bond (c & p).
Tomes Collection.
Tomes Collection.
C. Hilgert (c).
Lord Lilford (p).
Lord Lilford (p).
Dr. A. Giinther (p).
Purchased (Parreys).
Budapest Museurq. (e).
Budapest Museum (e).
C. G. Danford and J. A.
Brown (c & v).
Lord Lilford (p).
Lord Lilford (p).
Hon. N. C. Rothschild
C. Mottaz (c).
Hon. N. C. Rothschild
Florence Museum (e).
G.Barrett-Hamilton (p).
O. Thomas (p).
Dr. V. L. Seoane (p).
Seville Museum (e).
Leyden Museum (e).
11. 1. 3. 45.
11.1.3.46-47.
90. 4. 17. 1.
97. 8. 27. 1.
11.1.3.42-44.
44. 6. 16. 7.
61. 11. 5. 1-2.
7. 1. 1. 352.
7. 1. 1. 353.
8. 11. 2. 1-2.
11. 1. 1. 38.
11. 1. 1. 33-.35.
66. 2. 1. 7-8.
46. 6. 15. 54.
94. 3. 1. 12-13.
94. 7. 18. 6-7.
74. 7. 4. 1-5.
11. 1. 1. 36-37.
4. 4. 6. 2-3.
8. 10. 2. 1-13,
15.
8. 11. 8. 3.
8. 10. 2. 14.
85. 7. 6. 13.
11. 1. 2. 28.
8. 2. 9. 1.
94. 1. 1. 6.
94. 5. 8. 1.
37. 4. 28. 58.
EPTESICUS SODALIS Barrett-Hamilton.
1910. Vespertilio sodaU.v Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
8th ser., v, p. 291, March, 1910. Type in British Museum.
1910. Eptesicus sodalis Trouessart, Faune !!Mamm. d'Europe, p. 22.
Tj/pc locality. — Bustenai'i, Prahova, Koumania (in Carpa-
thians, alt. 840 m.).
(h'0(iraphical (listrihufidu. — Known only from the type locality
and St. Gothard, Switzerland.
* Mr. Cabrera has come to the same conclusion (Bol. Real Soc. Espafi.
Hist. Nat., VI, p. 449, December, 1908).
232
CHIEOPTERA
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EPTESICUS
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234
CHIROPTERA
Diagnosis. — Similar to Eptesims serotinus but smaller, condylo-
basal length of skull about 18 mm. instead of 19 to 21-6 mm.
Measurements. — Type (young-adult male) : head and body,
63; tail, 42; tibia, 18-6; foot, 9-8; forearm, 45-4; third
finger, 79-0; fifth finger, 58-0 ; ear (fresh), 18. Adult from
St. Gothard, Switzerland : tibia, 19 ; forearm, 48. For cranial
measm-ements see Table, p. 233.
Specimens examined. — Two, from the following localities : —
Switzerland: St. Gothard, 1 (U.S.N. M.).
RouMANiA : Bustenari, Prahova, 1 (type).
RemnrJcs. — The two specimens on which this species is based
indicate the existence of an animal bearing much the same
relationshij) to Eptesirus serotinus as Nyctalus nocfula to N.
maximus.
S. Bustenari, Prahova, 840 m. Lord Lilford (p). 4. 4. 6. 1.
Roumauia. (W. Dodson.) [Type of species.)
EPTESICUS NILSSONII Keyserling and Blasius.
1836. Vespertilio kuhlii Nilsson, Ilium. Fig. Skand. Fauna, pt. 17, pi. .34
upper figure. Not of Kuhl, 1819.
1888. Vespert\ilio'] borealis Nilsson, Ilium. Fig. Skand. Fauna, pt. 19,
pi. 34 (renumbered 36) upper figure. Not of P. L. S. Miiller, 1776
(Scandinavia).
1839. V[csp}crfilio] nilsso7iii Keyserling and Blasius, Wiegmann's Archiv
fiir Naturgesch., 1839, p. 315 (Mountains of Scandinavia. Based
on the V. kuhlii of Nilsson, 1836).
1857. Vesperugo nilssonii Blasius, Saugethiere Deutschlauds, p. 70.
1858. Amhlyotas atratus Kolenati, Sitzungsber. kais. Akad. Wissensch.
Wien, Math.-Naturwissensch. Classe, xxix, p. 252 (Altvater,
Austrian Silesia, alt. 2400-4600 ft.).
1878. Vesperugo burealis Dobson, Catal. Chiropt. Brit. Mus., p. 203.
1894. Vesperugo nillsoni (sic) Rhoads, Reprint Ord's N. Amer. Zoology,
Append., p. 3.
1907. E['ptesicus'] nilsso7ii Miller, Families and Genera of Bats, p. 209,
June 29, 1907.
1910. Eptesicus nilssoni Trouessart, Faune Mamm. d'Europe, p. 23.
Type localiUj. — Sweden.
Geographical distribution. — Continental Europe, from northern
Norway to the Alps.
Diagnosis. — Size medium (forearm less than 40 mm., condylo-
basal length of skull less than 16 mm.) ; colour of upper parts a
rich dark brown, the hairs of back with noticeably contrasted
light tips ; underparts light yellowish brown ; a well defined
line of demarcation along sides of neck.
External form. — In general the external form agrees with
that of Eptesicus serotinus, due allowance being made for the less
robust stature of the smaller animal. Ear relatively Ljnger,
EPTESICUS
235
extending to nostril when laid forward, though of the same
general form as in E. serotinus ; tip less narrowly rounded oft',
and flattened or concave portion of posterior border less con-
spicuous ; tragus relatively shorter and wider, though not
essentially difterent in form, its greatest width nearly equal to
length of anterior border ; transverse striations on inner surface
of conch oljsolete. Wings and feet essentially as in E. serotinus.
Tail slightly longer than in the related animal, extending nearly
to head when laid forward, its terminal vertebra free.
Fur and colour. — In quality the fur resembles that of E.
serotinus except that it is softer and more silky in texture, the
hairs fullj- as long as in the larger animal. In distribution it
shows no special peculiarities ; dorsal surface of uropatagium
thinly furred to about middle instead of on extreme base only.
Colour above a rich daik brown, ranging from burnt-umber nearly
to seal-brown, the hairs everywhere with slightly darker, faintly
slaty bases, those of median dorsal region from crown to base of
tail tipped with light glossy ochraceous-buft' in evident contrast
with ground colour, the light tips most numerous behind shoulders,
and forming a noticeable mantle over middle of back ; under-
parts rather strongly contrasted light yellowish brown, between
the wood-brown and ochraceous-buiF of Ridgway, the basal
portion of the hairs similar to ground colour of back, the yellowish
brown area extending over sides of head and completely encircling
base of ear, the line of demarcation between it and the dark
brown of upper parts sharply defined along sides of neck. Muzzle,
cheeks, ears and membranes blackish.
Skull. — In general the skull difiers from that of Eptesicus
serotinus, apart from its smaller size, in a general tendency to
greater depth, less elongation, and smoother, more evenly
rounded surfaces. Dorsal profile with evident convexity at middle.
Lambda not overhanging ; low, ill-defined
lambdoid crests curving slightly toward point
of contact at middle, the rounded 2:)osterior
outline of the occiput plainly visible behind
them when skull is viewed from above ;
sagittal crest essentially absent. Brain-case
sub-spherical or broadly ovate in outline,
its depth slightly more than half mastoid
breadth ; floor of brain case marked by a
wide lateral groove between each cochlea
and the median line ; auditory bulla? slightly
larger than in E. ,sr, o//hhv. Interorbital fig. 41.
region relatively less constricted than in Epte^icus nU:<xonu.
E. serotinus, but of essentially the same form ; ^at- si^e.
lachrymal swelling present but less noticeable
than in the larger animal. Rostrum rounded off" at sides, with
only the faintest trace of lateral concavities ; narial and palatal
emarginations essentially as in E. serotinus. Palate showing no
236
CHIROPTERA
noteworthy i^eculiai'ities, its general outline less narrow than in
the related species.
Teeih. — Inner upper incisor as in Epfesicus serotinus, but more
robust in proportion to its height ; outer incisor noticeably higher
than in the related species, its apex reaching level of secondary
cusp of larger tooth, its secondary cusp more prominent ; no
marked contrast between crown areas of the two teeth ; each
pair in line of general curve of anterior jjortion of palate, instead
of at right angles to main axis. Lower incisors less crowded
than those of E. serotinus, their imbrication distinct but not
unvisual, the general outline of the row broadly V-shaped ; in form
the individual teeth show no special peculiarities ; /^ without
postero-internal tubercle. Canines and premolars with no special
peculiarities. First and second upper molars essentially as in
Eptesicus serotinus, but disproportion between paracone antl
metacone less evident ; m^ with crown area about two-thirds that
of wi', its longitudinal diameter through metacone slightly more
than half trans^■erse diameter, the mesostyle and metacone well
developed, the second and third commissures more than half as
long as first ; lower molars like those of E. serotinus in form, but
angles in commissures between outer and inner cusps deeper ;
area of second V ^^ '"3 nearly equal to that of first.
Measurements. — Adult male from Grotte de Vallorbe, Vaud,
Switzerland: head and body, 68 • 5 ; tail, 47; tibia, 17; foot, 10;
forearm, 38 • 2 ; thumb, 9 • 8 ; third finger, 68 ; fifth finger, 49.
Forearm in adult male from Upsala, Sweden, 38-4. Three
females from the same locality : forearm, 39, 39, and 39 ■ 6 ;
third linger in the four specimens from Upsala : 66, 68, 70 and
68. For cranial measurements see Table opposite.
Specimens examined. — Fifteen, from the following localities : —
Norway : No exact locality, 1.
Sweden : Upsala, 8 (B.M. and U.S.N.M.) ; Upland, 2 (U.S.N.M.).
Germaky : Wernigerode, Saxony, 2.
Austria-Hungary : Csallokoz-Somorja, Pressburg, 1.
Switzerland : Grotte de Vallorbe, Vaud, 1 (Mottaz).
Itemarlcs. — This species is readily distinguishable from Epfesicus
sn-otinus by its smaller size and by the conspicuous pale tips to
the hairs of back. From Vespertilio murinus, which resembles it
in colour jiattern, the narrow ear, more yellowish (less whitish)
hair-tips on hack, and slightly smaller size distinguish it super-
ficially. Pipistrellus sarii, which often has almost exactly similar
coloration, is a much smaller animal (forearm, 31 to 33 instead
of 38 to 40).
1. Norway. {Collctt.) E. E. Alston (p.) 81. 6. 9. 2.
4 9. Upsala, Sweden. {EoUhoff.) Lord Lilford (p). 11.1.1.39-42.
6, ?. Wernigerode, Saxony, Ger- Dr. W. WolterstorfE 0. 2. 8. 3-4.
many. (c&p).
1. Csallokoz-Somorja, Press- Budapest IMuseum (e.) 94.3.1.14.
burg, Hungary.
EPTESICUS
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238 CHIROPTERA
Genus VESPERTILIO LinnEeus.
1758. Vespertilio Linnseus, Syst. Nat., i, lOtb ed., p. 31 (nmriims by
tautonymy).
1839. Vesperugo Keyserliug and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, i, p. 312 (part).
1839. Vesperiis Keyserling and Blasius, Wiegmann's Archiv fiir Naturgesch.,
1839, I, p. 313. Sub-genus of Vesperugo (part).
1856. Meteorus Kolenati, Allgem. deutscbe Naturbist. Zeitung, Dresden,
neue Folge, ii, p. 131 (part).
1857. Vesperus, misprinted Vesperugo Blasius, Saugetbiere Deutscblands,
p. 69 (Sub-genus of Vesperugo).
1863. '^ Aristippe Kolenati, Beitrage zur Kenntniss der Pbtbiriomyiarien,
Petersburg, 1863" (part).
1872. Marsipolxmus Peters, Monatsber. k. preuss. Akad. Wissenscb.,
Berlin, p. 260 (Sub-genus of Vesperugo for albigularis = murinus).
1878. Vesperus Dobson, Catal. Cbiropt. Brit. Mus., p. 183. Sub-genus of
Vesperugo (part).
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., Ctb ser., xx, p. 384,
October, 1897 (part).
1900. Vespertilio Mebely, Monogr. Cbiropt. Hungariaj, p. 219 (part).
1907. Vespertilio Miller, Families and Genera of Bats, p. 209, June 29, 1907.
Type species. — Vespertilio 'murinus Linn^^^us.
Geograpliical distribution. — Forested northern portion of the
Paltearctic region from the Atlantic coast eastward through
Continental Asia.
Characters. — Like Eptesicus, but ear much shortened and
broadened, rostrum flattened above, with deep concavity on
each side between nares and lachrymal region ; nares very large,
extending back nearly half way to interorbital constriction, and
palatal emargination extended so far laterally that its width is
distinctly greater than its depth.
Remarlcs. — The genus Vespertilio as now restricted contains
only two species, V. murinus Linnaeus, and V. superans Thomas,
the former European, the latter Asiatic. The form of the skull
.suggests in certain respects that of Nyctalus, and in others the
North American Lasionj/cteris.
VESPERTILIO MURINUS Linn;eus.
1758. [Vespertilio] murinus Linnaeus, Syst. Nat., i, 10th ed., p. 32 (Sweden).
1819. Vespertilio discolor Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 187 (Vienna, Austria).
1857. Vesperugo discolor Blasius, Saugetbiere Deutscblands, p. 73.
1872. Vesperus [Marsipolsemus) albigularis Peters, Monatsber. k. preuss.
Akad. Wissenscb., Berlin, p. 260 (Type supposed to have been
taken in Mexico ; for its reference to this species see Mebely,
Monogr. Cbiropt. HungariEe, pp. 229, 341, 1900).
1878. Vesperugo discolor Dobson, Catal. Cbiropt. Brit. Mus., p. 204.
1885. Vesperus siculus Daday, Orvos-Termeszettudomduyi Erteseto, Kolozs-
var, X, p. 275 (Homorod-Almas cave, Hungary). See Mebely,
Monogr. Cbiropt. Hungarise, pp. 229, 346, 1900.
VESPERTILIO
239
}
188G. Vesperus siculus Daday, Verhandl. u. Mittheilungen des Siebenbiirg-
ischen Vereinsfiir Naturwissensch. in Hermauustadt, xxxvi, p. 82.
1897. Vcs2)crtilio murinus Miller, Ann. aud Mag. Nat. Hist., 6th ser., xx,
p. 382, October, 1897.
1910. Eptesicus siculus and Vcspertilio murinus Trouessart, Paune INIamm.
d'Europe, pp. 23 and 25.
Type locality. — XJpsala, Sweden.
Oeographical distribution. — Northern and central Europe,
from southern Norway and central Sweden south to the Alps.
One record of its occurrence in England.*
Diagnosis. — General characters as in the genus ; length of
forearm about 43 to 45 mm.
External characters. — General appearance much as in Eptesicus
nilssonii but slightly larger and more robust. Ear extending
nearly to nostril when laid forward, its width when flattened
about li times height from crown ; anterior basal lobe so much
reduced as to suggest the keel of the Molossidse, the anterior
border of conch nearly straight from forehead to broadly rounded
tip ; posterior border shallow ly sinuous-concave from just below
tip nearly to level of meatus ; here it turns forward almost at
right angles and extends nearly to angle of mouth, where it
terminates at a well developed wart, the terminal portion often
forming a slight fold or pocket ; antitragus low and keel-hke,
nearly 2 mm. from margin of conch ; inner surface of conch
slightly papillose, without evident cross ridges ; tragus low,
scarcely rising above level of outer angle of anterior lobe, its
greatest width (slightly above middle) nearly equal to length of
straight or slightly concave anterior border, its tip broadly
rounded, the convexity continuous behind with that of upper
portion of posterior border ; posterior basal lobe obsolete. Wing
rather narrow, the fifth finger exceeding forearm by one-sixth to
one-fifth length of forearm, the membranes thin, extending to
base of outer toe ; third and fourth metacarpals sub-equal, nearly
5 mm. shorter than forearm, fifth about 3 mm. shorter than
third. Leg rather robust ; foot about half as long as tibia ;
calcar slightly longer than tibia and free border of uropatagium,
its keel ill defined, its termination obscure. Tail extending to
between shoulders when laid forward, the last vertebra free.
Fur and colour. — The fur is rather short and dense, less silky
than that of Eptesicus nilssonii, the hairs at middle of back about
7 mm. in length. In distribution it shows no special peculiarities ;
upper surface of interfemoral membrane furred on basal third.
Colour above essentially as in Eptesicus 7iilssonii, but liglit tips to
hairs very pale, almost whitish crcam-bufl", producing a decidedly
" frosted " appearance ; underparts creain-buft", the dusky under
colour showing through on chest and anterior portion of belly ;
line of demarcation on sides of neck sharply defined ; muzzle and
chin dusky ; ears and membranes blackish.
* A single specimen, undoubtedly a straggler, taken at Plymouth.
240 CHIROPTEEA
ShiU. — The skull combines the short, smooth brain-case of
Ej^tesicHS nihsonii with the broad, flat rostrum of E. serotinus, but
differs conspicuously in aspect from both in the very deep,
N>/cfahis-like narial and palatal emarginations. Dorsal profile
essentially straight from nares to rounded, not overhanging
lambda. Sagittal crest obsolete ; lambdoid crest low but evident.
Depth of brain-case at middle decidedly more than half mastoid
breadth ; floor with wide conspicuous slit between cochlea and
basioccipital : a flattened pit-like depression at front of each slit ;
auditory bullae moderately large, their trans-
verse diameter about equal to space between
them. Interorbital region broadly hour-glass
shaped, flattened above ; edge of orbit from
lachrymal region nearly to most constricted
portion distinctly and irregularly swollen.
Dorsal surface of rostrum flattened, with broad,
shallow lateral depressions ; narial emargina-
tion broadly triangular, its apex extending
back to level of lachrymal region ; rostral depth
at front of orbit equal to distance from orbit
"'r' '"" . to front of inner incisor ; anteorbital foramen
\ es'perlilio murvaux. . „
Nat. size. moderately large, over point ot contact between
large premolar and first molar ; lachrymal fora-
men directly behind it on inner rim of orbit. Palate broad and
short, noticeably concave both laterally and longitudinally, the
anterior emargination large, much wider than deep, but extend-
ing back to level of middle of premolar ; posterior extension of
palate parallel sided, its length slightly greater than width
behind molars ; hamulars slightly bent inward ; median spine
well developed. Mandible robust, much deeper at symphysis
than behind tooth-row, the coronoid process low (height less than
least width of posterior section) with gradually sloping upper
border ; angular process rather slender, its extremity slightly
curved inward.
Teeth. — While in general resembling those of the European
species of Eptesicus, especially those of E. nilssonii, the teeth of
Vesjpertilio murinus show certain notable peculiarities. Inner
upper incisor similar to that of Epfesicus serotinus when viewed
from in front, but with well developed postero-basal cusp which
in some specimens rises nearly to half the height of main cusp,
l)roducing a distinctly trifid tooth ; outer incisor with crown area
slightly less than that of inner, the cusp rising nearly to middle
of shaft of inner tooth ; secondary cusp slightly developed ;
position of incisors relatively to tooth-row about as in E. nilssonii.
Lower incisors as in E.nil.'^.'ionii. Upper canine with transverse
diameter slightly greater than longitudinal diameter, the reverse
of the condition in the two species of Ej^tesicus ; lower canine
with same peculiarity in outline of crown though to a less degree.
Upper premolar with longitudinal diameter of crown relatively
VESPERTILIO
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less than in the European species of Eptesicus, and antero-interual
basal cusp better developed : lower premolars more crowded and
compressed than in E. nihsonii, and crown area of first relatively
smaller. Molars as in E. nilssonii except that columnar hypocone
of m^ and m? stands out more prominently from outline of
protocone.
3Ieasiirements. — Three adult males from Denmark : head and
body, 62, 59 and 62; tail, 43, 40 and 44 ; tibia, 17-4, 16-8 and
16 • 8 ; foot, 9 • 6, 9 • 2 and 10 ; forearm, 43 • 4, 44 and 43 ; thumb,
7, 7 '4 and 7' 6; third linger, 73, 76 and 74; fifth finger, o2, 54
and 53; ear from meatus, 15, 15 and 15' 6; width of ear, 16,
16 and 17. Adult female from the same locality: head and
body, 63 ; tail, 41 ; tibia, 16 • 4 ; foot, 9 ; forearm, 45 ; thumb, 8 ;
third finger, 78; fifth finger, 54: ear from meatus, 16; width
of ear, 17. Adult male from Morat, Fribourg, Switzerland:
head and body, 56 ; tail, 42 ; tibia, 16 : foot, 8*8 ; forearm, 45 :
thumb, 5'4 ; third finger, 76 ; fifth finger, 55 ; ear from meatus,
12; width of ear, 12 "4. For cranial measurements see Table,
p. 241.
Specinieyis examined. — Twelve, from the following localities : —
England : Plymouth, 1.
Sweden : No exact locality, 1.
Denmark: No exact locality, 4 (U.S.N.M.).
Germany: Gross Hennersdorf, Saxony, 1; Bavaria, 2 (U.S.N.M.).
AnsTRiA-HuNGARY : Csallokoz-Somorja, Pressburg, 1.
Switzerland : Morat, Fribourg, 1 (Mottaz).
Locality unknown: One; type of aifcir/aiaris Peters (Berlin).
Remarks. — Superficially this bat somewhat resembles Ejjtesicns
nilssoni. It is readily distinguishable, however, by its slightly
larger size (forearm, 43 to 45 instead of 38 to 40), low, rounded
ear, and l)y the whitish rather than yellowish wash on back.
1. Plymouth, Devonshire, Dr. Leach (p). 37. a.
England.
— Sweden. Stockholm Museum (e). 46. 6. 2. 21.
?. Gross Hennersdorf, Lord Lilford (p). 99.1.9.5.
Saxony {W. Baer.)
1. Csallokoz-Somorja, Press- Budapest ^luseum (e). 94. 3. 1. 1-5.
burg, Hungary.
Genus NYCTALUS Bowdich.
1825. Nijctalus Bowdich, Excursions in ^Madeira and Porto Santo, p. 36
(verrucosus).
1829. Pten/gistes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt,
I, p. 99.
1839. Vesperucjo Keyserling and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, i, p. 312 (part).
1842. Noctulinia Gray, Ann. and Mag. Nat. Hist., x, p. 258 (part ; contained
inoterus = noctula and fulvus = Scotophilus kulilii).
1856. Fanugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 {noctula and leisleri).
1857. Vesperugo Blasius, Siiugethiere Deutschlauds, p. 49 (part).
NYCTALUS 243
1878. Vesperugo Dobson, Catal. Chiropt. Brit. Mus., p. 183 (part).
1893. Noctulinia II. Allen, Proc. U.S. National Museum, xvi, p. 30,
June 13, 1893.
1897. Pterygistes Miller, Ann. and Mag. Nat. Hist., Gth ser., sx, p. 384,
October, 1897.
1899. Euvcspenigo Acloque, Faune de Prance, Mammiferes, p. 32 (part,
included twctula, leisleri, maurus, kuhlii, pipistrellus, and
abramus).
1907. Pterygistes Miller, Families and Genera of Bats, p. 207, June 27,
1907.
1908. Nyctalus Andersen, Ann. and Mag. Nat. Hist., 9th ser., i, p. 434,
May, 1908.
Type species. — Nyctalus verrucosus Bowdicli.
Geographical distribution. — Northern portion of Eastern Hemi-
sphere from the Azores and Madeira to Japan.
Characters. — Dental formula as in Pipistrellus ; skull with
nares extending unusually far back, half way to interorbital con-
a b c d
Fig. 43.
Xi/ctahis maximus (a), N. noctula (b), N. leiilen (c), and X. azoreum (A). Jfat. size.
striction, and with large anterior palatal emargination ; fifth finger
much shortened, scarcely exceeding metacarpal of fourth or third.
Hemarhs. — This genus is well diflerentiated from allied groups
by the peculiar narrowing of the wing due to the shortness of the
fifth finger. Three of the half dozen known species occur on the
continent of Europe, while a fourth inhabits the Azores. The
most obvious character by which the European forms are dis-
tinguished is the size of the skull (see fig. 43).
KEY TO THE EUROPEAN SPECIES OP NYCTALUS.
Condylobasal length of skull more than 17 mm. ; fore-
arm more than 45 mm. ; hairs of back without
dark bases.
Condylobasal length of skull 22 to 23 mm. ; forearm
64 to 68 mm. (Southern Europe) N. maximus, 'p. 244.
,Condylobasal length of skull 17 ■ 4 to 19 • 4 mm. ; fore-
arm 47 to 55 mm. (Distribution general) N. noctula, p. 245.
Condylobasal length of skull less than 17 mm. ; fore-
arm less than 45 mm. ; hairs of back with notice-
able dark bases.
Condylobasal length of skull 15 to 16 mm. ; forearm
39 to 43 mm. (Distribution general) N. leisleri, -p. 252.
Condylobasal length of skull 13 to 14-2 mm. ; fore-
arm 35 to 42 mm. (Azores) N. asorcum, p. 254.
R 2
244
CHIROPTERA
NYCTALUS MAXIMUS Fatio.
1781. ? Vespe^'tilio lasioptcrns Schrober, Siingethiere, pi. lvii b. See
Thomas, Ann. and Mag. Nat Hist., 8th ser., viii, pp. .379-380,
September, 1911.
1827. ?■? Vespertilio ferrufjineus Brehm, Ornis, Heft iii, p. 26 (Renthendorf,
Thiiringen, Germany).
1869. [Vcsperugo noctula] var. viaxima Fatio, Faune Vert. Suisse, i, p. 67.
Co-type in Geneva Museum.
1900. Pterygistes maximus Miller, Proc. Biol. Soc. Washington, xiii,
p. 156, June 13, 1900.
1910. Nydalus maximus Trouessart, Faune JIamm. d'Europe, p. 19.
Ty2)e locality. — Amsteg, Uri, Switzerland.
Geographical distribution. — At present known from a few
localities in Switzerland and Italy.
Diagnosis. — Essentially similar to Nyctalus noctula but much
larger; condylobasal length of skull, 22 to 23 mm. ; forearm, 64
to 68 mm.
External cltarncters. — Except for the conspicuously greater
size and consequent more robust form, there appears to be no
tangible character by which the animal can be distinguished
from N. noctula.
Fur and colour. — In the few specimens examined the furred
area on under surface of wing membrane behind forearm appears
to be better defined than in N. noctula. Colour as in the common
species.
Skull and teeth. — The skull and teeth so closely resemble those
of Nyctalus noctula that in general they differ in their greater
size only. The posterior portion of occiput is, however, more
elevated above base of cranium than in the smaller animal, so
that lower edge of condyle is about on level with anteorbital
foramen and alveolus of canine instead of distinctly below them.
Correlated with this character is a more abrupt I'ising of lambdal
region above level of anterior portion of brain-case, and a less
nearly horizontal truncation of occipital region. Palate appar-
ently more concave longitudinally than in any of the smaller
European species.
Measurements. — Adult male and female from Pisa, Italy :
head and body, 78 and 87 ; tail, 59 and 66 ; tibia, 23 • 6 and 24 ;
foot, 12 and 13 ; forearm, 65 and 68 ; thumb, 11 and 11 ; third
linger, 119 and 123 ; fifth fing(!r, 76 and 77 ; ear from meatus,
21 and 22 ; width of ear, 23 and 24. Adult male from Venice : *
head and body, 92 ; tail, 65; tibia, 26; foot, 14 ; forearm, 67.
Adult female from Amsteg, Uri, Switzerland : head and body, 90± ;
tibia, 21-6 ; foot, 13 ; forearm, 64 ; thumb, 9-4 ; third finger,
114 ; fifth finger, 72. For cranial measurements see Table, p. 250.
Specimens examined. — Five, from the following localities : —
Switzerland: Amsteg, Uri, 1 (Geneva; co-type).
Italy : Pisa, 3 (B.M. and U.S.N.M.) ; Ravenna, 1.
Ninni, Atti Soc. Ital. Sci. Nat., Milano, xxvi, p. 109, 1883.
NYCTALUS 245
Remarks. — Ni/ctalus maximus is readily distinguished among
the bats of Eui-ope by its large size. It is at present a rare and
little known animal.
9 al. Pisa, Italy. Dr. A. Senna (e). 94. 6. 3. 3..
6 al. Ravenna. Florence iluseura (e). 85. 7. C. 9.
i al. Lidth dc Jeude Coll. 67. 4. 12. 337.
NYCTALUS NOCTULA Schreber.
1774. Vcspcrtilio nochda Sohreber, Saugthiere, i, pi. lii ; description, i,
p. 16G, under name : Die Speckmaus (France ; based primarily
on " La Noctule " of Daubenton, Hist. Acad. Royale des Sci.,
Paris, 1759, p. 376. 1765).
1776. Vespertilio lardarius P. L. S. MiiUer, Natursyst. Suppl. u. Regist.-
Band, p. 15 (France; based primarily on Schreber's Speckmaus).
1789. [Vespertilio] magnus Borkenhaut, Syn. Nat. Hist. Great Britain and
Ireland, i, p. 1 (Cambridge, England; based on Pennant, Brit.
Zool., No. 38).
1789. Vespertilio altivolans White, Nat. Hist, and Antiquities of Selborne,
p. 93 (Selborne, Hampshire, England).
1816. ? Vespertilio major Leach, Syst. Catal. Spec. Indig. Mamm. and
Birds Brit. ]Mus. (Willoughby Society reprint, 1882), p. 5 Nomen
nudum : " Great Bat."
1818. Vespertilio proterus Kuhl, Ann. Wetterau, Gesellsch. Naturk, iv
(= Neue Ann., i), pt. 1, p. 41 (Substitute for noctula).
1829. Vespertilio rufesccns Brehm, Isis, p. 643 (Jena, Thiiringen, Germany).
1841. Vespertilio noctula Bonaparte, Iconogr. Faun. Ital., i, lud. Distrib.
(sp. iUustr.).
1841. Noctula serotina Bonaparte, Iconogr. Faun. Ital., i, Ind. Distrib.
(uomencl. moderna).
1844. Vcsp[ertilio] palustris Crespon, Faune M6ridionale, i, p. 22 (marshes
near Nimes, Gard, France).
1857. Vesperugo noctula Blasius, Saugethiere Deutschlands, p. 53.
1869. [Vesperugo noctula] var. minima Fatio, Faune Vert. Suisse, i, p. 58
(Geneva, Switzerland).
1878. Vesperugo noctula Dobson, Catal. Chiropt. Brit. Mus., p. 212.
1897. Ptenjqistes noctula Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,
p. 384, October, 1897.
1910. Nyctalus noctula Trouessart, Faune Mamm. d'Europe, p. 18.
Tjipe locality. — France.
Geographical diMrihutian. — Europe from southern Norway
and central Sweden to the Mediterranean, and from England
eastward into Asia.
Diagnosis. — Condylobasal length of skull, 17 '4 to 19 '4 mm. ;
forearm, 47 to 55 mm. ; general colour dark j'ellowish brown, the
hairs not darker at base.
External characters. — General form robust and heavy. Muzzle
broad, with conspicuous glandular swelling between eye and
nostril, the greatest width across this region decidedly more than
distance between nostril and ear ; nostrils projecting forward and
outward with evident median concavity between them, the orifice
crescentic. Ear short, extending when laid forward about half
246 • CHIROPTERA
way from eye to nostril, its breadth when flattened decidedly
greater than height above crown ; anterior border of conch
abruptl}' convex below, then nearly straight to broadly rounded
off extremity ; posterior border convex throughout, most strongly
at middle ; antitragus long and low, well marked off posteriorly,
its anterior border extending to just below angle of mouth ; inner
surface of con^h finely papillose, without evident cross ridges ;
tragus very short, scarcely rising above level of anterior base of
conch, much wider above than below, its greatest width about
equal to height ; anterior and posterior borders concave, the
anterior more so than posterior, their length about equal to that
of expanded upper portion, which is more abruptly rounded
anteriorly than posteriorly. Wing long and slender, the iifth
finger exceeding forearm by only ^ to ^ length of latter, the
membrane leathery and opaque, joining leg at ankle ; third and
fourth metacarpals sub-equal, the third slightly the longer and
about equal to forearm, fifth abruptly shorter by slightly more
than one-fifth forearm. Leg short and strong, the broad foot
about one-half as long as tibia ; calcar 1 ^ to twice as long as
tibia or as free border of interfemoral membrane, its distal
termination obscure but basal portion strong, well defined and
with keel about 2-5 mm. wide supported by a well defined
thickening and terminating in an abrupt convexity on side
nearest heel. Tail rather short, extending to between shoulders
when laid forward, the tip of last vertebra (about 2 ram.) free.
Fiu- and colour. — Fur dense and velvety, the hairs at middle
of back only about T) mm. in length ; on dorsal surface of wing
it extends to line joining knee with middle of humerus, on ventral
surface to line from elbow to knee ; beyond this line it spreads
thinly on under surface of antebrachial membrane and also
behind forearm and across bases of metacarpals along an area
about 10 mm. wide. Interfemoral membrane furred both above
and below at extreme base only. Colour of upj^er parts a rather
dark yellowish brown, near the wood-brown and cinnamon of
Ridgw ay, the hairs showing a faint clouding of prouts-brown in
certain lights, their basal portion a light dull isabella-colour ;
underparts scarcely difierent from back, though usually a little
lighter and more dull ; muzzle and cheeks dusky ; ears and
membranes blackish.
Skull. — General aspect of skull broad and robust, rather high
posteriorly, low anteriorly, with conspicuous narial emargination.
Dorsal profile rising rather rapidly from incisors to lambda, with
slight convexity over lachrymal region and a slighter though
more abrupt concavity just in front of lambda. Ventral profile
elevated posteriorly. Brain-case ovate, but with wide mastoid
and paroccipital region and squarely ti-uncate occiput which
together produce a distinctly truncate-cuneate outline, slightly
though evidently wider than long ; depth at middle distinctly
more than half mastoid breath : sagittal crest low but evident,
247
I
Nyctalus noctiUa.
Nat. size.
the region on each side of it flat or sHghtly depressed ; lambdoid
crest high, curved abruptly forward at point of junction with
sagittal crest ; floor of brain-case with small but evident lateral
pits, well defined anteriorly but communicating posteriorly with
vacuity between cochlea and basioccipital ; auditory bulhe well
developed but of moderate size, the transverse diameter al>out
equal to distance between bulhe. Interorbital region deeply
constricted, short hour-glass shaped, the lachiymal region widen-
ing abruptly to a breadth nearly equal to that of brain-case, and
forming a slight though evident tuber-
cular projection over anterior rim of orbit ;
rostrum squarish, somewhat narrower
anteriorly than posteriorly, the oval or
ovate narial emargination extending back
to level of lachrymal foramen ; rostral
depth at front of orbit about equal to
distance from orbit to front of inner
incisor ; anteorbital foramen small, over
space between large jiremolar and first
molai', lachrymal foramen slightly above
and behind it, on orbital rim. Palate
rather short, owing to the large size of
the anterior emargination, the posterior
edge of which is on level with middle of
large premolar, rather evidently concave
laterally, less so longitudinally. Posterior extension of palate
nearly parallel sided, its width at level of posterior edge of third
molar slightly less than length ; hamulars barely turned inward ;
median spine large, acute. Mandible robust, noticeably deeper
at symphysis than immediately behind tooth-row ; posterior
portion rather low and long, the height of coronoid process
above alveolus less than least length, the upper border sloping
gradually from coronoid to articular jsrocess ; angular process
moderately long, about on level with alveolar line, its main
axis directed rather abruptly outward, its distal extremity
oblicjuely widened.
Ti/efh. — Relatively to size of skull the teeth are large, though
the cusps are rather low. Inner upper incisor slender, with
slightly developed cingulum, its crown area barely one quarter
that of canine, the nearly terete shaft directed obliquely inward,
its apex extending slightly beyond level of cingulum of canine,
its .secondary cusp small but evident, situated on postero-external
surface of shaft ; outer upper incisor with crown area distinctly
greater than that of inner tooth, the shaft deeply concave, its
concavity directed outward and backward, the main cusp lying close
against secondary cusp of inner tooth, the well-defined se-.-ondary
cusp relatively much larger than that of inner incisor, cingulum
M-ell developed, bearing a small but distinct antero-external cusp
on opposite side of concavity from secondary cusp. The two
248
GHIROPTERA
incisors are closely crowded against each other ; outer tooth
separated from canine by a very narrow space. Lower incisors
rather strongly imbricated, forming a very broadly U'Stiap^d
row between canines, the crowns of /^ and /., compressed, trilid,
that of i^ sub-terete, flattened anteriorly, with two low posterior
tubercles in addition to the three cusps corresponding to those of
other teeth. Upper canine robust, with well developed anterior
and posterior cutting edge, the shaft triangular in cross-section ;
cingulum well developed but without true secondary cusps.
Lower canine with posterior surface of shaft strongly concave,
the well developed cingulum forming a postero-internal basal
cusj) and a conspicuous secondary cusp near middle of antero-
internal border of shaft ; diameter of crown distinctly less than
distance between canines. Anterior upper premolar minute, closely
wedged in space between canine and posterior premolar, its cross-
section less than half that of incisors, its crown reniform in
outline, its cusp low but distinct ; posterior upper premolar with
crown area somewhat more than half that of first molar, the
protocone well developed, nearly as high as in molars, the main
cusp rather high but much shorter than canine (measured along
cingulum), its two cutting edges well developed. Lower pre-
molars closely crowded, their ci-owns about equal in cross-section
and somewhat more than half that of canine, the shaft of first
lower and more robust than that of second and about half as
high as canine ; cingulum well developed, in each tooth forming
a small but evident antero-internal and postero-internal cusp.
First and second upper molars with crowns rather broad on
inner side, the posterior emargination slight ; protocone robust
but low ; hypocone small but well developed and distinct from
commissure of protocone ; outer cups without sjiecial peculiarities,
the styles low but well developed, the VVP'ittern normal, //t^ with
crown area distinctly more than half that of m}, its longitudinal
diameter through metacone about half transverse diameter, its
three cusps, two styles and three commissures well developed,
but proportion between paracone and metacone reversed as
co)npared with other molars. Lower molars with unusually
robust hypoconid and entoconid, producing a strong contrast in
size between the anterior and posterior V in «*i and in.^ and
approximate equality in m-^ ; cingulum well developed but not
forming basal cusjd behind entoconid.
Measurements. — Adult female from Kew Gardens, Surrey,
England: head and body, 71 ; tail, 51 ; tibia, 18; foot, 10;
forearm, 52 ; thumb, 9 ; third finger, 94 ; fifth finger, 58 : ear
from meatus, 15; width of ear, 16. Adult female from Herrn-
hut. Saxony, Germany : head and body, 79 ; tail, 53 ; tibia, 19 ;
foot, 10' 6; forearm, 53; thumb, 9*4; third finger, 97; fifth
finger, 60 ; ear from meatus, 17 ; width of ear, 17. Two adult
males from Pisa, Italy : head and body, 70 and 71 ; tail, 49 and
52-6 ; tibia, 18 and 19 ; foot, 10 and 11 , forearm, 51 and 53 :
NYCTALUS 249
thumb, 8 and 9 ; third finger, 92 and 97 ; fifth finger, 56 and 59 ;
ear from meatus, 14 "6 and 16 ; width of ear, 14 and 15 "6. Two
adult females from the same locahty : head and body, 74 and 75 ;
tail, 50 and 53 ; tibia, 18 '6 and 18 ; foot, 10 and 11 ; forearm,
51-6 and 52 ; thumb, 9 and 10 ; third finger, 93 and 97 ; fifth
finger, 57 and 58; ear from meatus, 16 and 16; width of ear,
16 and 16. Two adult females from Corinth, Greece: head and
body, 69 and 76; tail, 51 and 54; tibia, 18 and 19; foot, 11
and 10 ; forearm, 52 and 54 ; thumb, 9 • 4 and 9 • 4 ; third finger,
96 and 102 ; fifth finger, 58 and 62 ; ear from meatus, 15-6 and
16 ; width of ear, 15 and 16. For cranial measurements see
Table, p. 250.
Specimens examined. — One hundred and fourteen, from tlie following
localities : —
England : Bowdon, Cheshire, 1 ; Stoke, Staffordshire, 1 ; Oundle,
Northamptonshire, 4 ; Bedfordshire, 1 ; Trumpington, Cambridgeshire, 1 ;
Cambridgeshire, no exact locality, 1 ; Henley-on-Tliames, Oxfordshire, 1 ;
Sandringham, Norfolk, 1 ; Kingsbury, Middlesex, 1 ; Chelmsford, Essex, 1 ;
Wcllyn, Hertfordshire, 1 ; Tring, Hertfordshire, 3 ; Wandsworth Common,
Surrey, 1 ; Shalford, Surrey, 2 ; Earlsfield, Surrey, 5 ; Kew Gardens,
Surrey, 1 (U.S.N.M.) ; Pulham Park, London, 1 ; Eastwell, Kent, 2 ; St.
Leonards, Sussex, 1 ; Parnborough, Hampshire, 1 ; Wareham, Dorset-
shire, 1.
Holland: Oosterbeek, Guelderland, 1.
GEEJtANY: Moritzburg, Saxony, 3 (U.S.N.M.); Herrnhut, Saxony, 2
(B.M. and U.S.N.M.) ; Niesky, Silesia, 1 ; Schwarzburg, Thiiringen, 2 ;
Strass, near Burgheim, Bavaria, 3 ; Bavaria, no exact locality, 8 (U.S.N.M.) ;
Stuttgart, 1.
Austbia-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 2.
RouMANiA : Gageni, 1 ; Bustenari, 1 ; no exact locality, 1 (U.S.N.M.).
Switzerland : Geneva, 4 (Geneva and Mottaz) ; St. Margarethen,
Appenzell, 8 (B.M. and U.S.N.M.).
Spain : Seville, 4 ; no exact locality, 2.
Italy : Pisa, 18 (U.S.N.M.) ; Plorence, 4 (Mottaz) ; Vallombrosa, near
Florence, 1 (U.S.N.M.) ; Siena, 1 ; Rome, 2.
Greece : Corinth, 12 (U.S.N.M.).
Bemarks. — This is one of the most widely distributed and
abundant of the larger European bats. From Eptesicits serotinus,
the only species with which it is likely to be confused, it is easily-
distinguished by the narrow wing, short, recurved tragus, and
low, rounded ear.
6. Aberia, Merionethshire, Col. Yerbury (c & p). 11. 1. 3. 26.
Wales.
9. Bowdon, Cheshire, Eng- G. Barrett-Hamilton 11. 1. 2. 100.
land. (T. A. Coivard.) (p).
jll.l. 1. 12G-
2 c5, 2 9. Oundle, Northampton- Lord Lilford (p). 127.
shire. 11. 1. 1. 129-
[ 130.
9. Bedfordshire. J. S. Elliot (c & p). 11. 1. 8. 391.
1. Trumpington, Cambridge- G. Barrett-Hamilton 11. 1. 2. 101.
shire. (p).
9 al. Cambridgeshire. Rev. L. Jenyns (c & p).
250
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CHIKOPTERA
Henley-on-Thames, Ox- Heatley Noble (c & p).
fordshire.
6 st. Sandringham, Norfolk.
?. Chelmsford, Esses.
?. Wellyn, Hertfordshire.
2 (5, ? st. Tring, Hertfordshire.
2 S. Shalford, Surrey.
3 d, 2 9. Earlsfield, Surrey.
<5st.
(5,9.
Fulham Park. Loudon.
Eastwell, Kent.
St. Leonards, Sussex.
Farnborough, Hampshire.
Wareham, Dorset.
Oosterbeek, Guelderland,
Holland.
Niesky, Silesia, Germany.
{W. Baer.)
Schwartzburg, Thtiringen,
200 m.
St r ass, Burgheim,
Bavaria. {Korhitz.)
Stuttgart, Wurtemburg.
Csallokoz-Somorja, Press-
burg, Hungary.
6. Gageui, Prahova, Kou-
mania. {W. Dodson.)
9. Bustenari, Prahova. (IT.
Dodson.)
2(5,3 9. St. Margaret hen,
Appenzell, Switzer-
land. (E.H.ZoUikofcr.)
Seville, Spain. {A. Ruiz.)
Spain,
3 6.
1 al.
4al.
2al.
i juv. al. Rome.
H.M. King Edward VII.
il. Christy and L. E.
Thompson (c & v).
C. H. B. Grant (c & p).
Hon. N. C. Rothschild
(c & p).
W. R. Ogilvie - Grant
(c & p).
C. H. B. Grant (c & p).
J. Saunders (c & p).
C. H. Caton - Haigh
(c & P).
R. Butterfield ^c & Pi.
H. S. Scott (c & p).
W. T. Blanford (c & p).
0. Thomas (c & p).
Lord Lilford (p).
Lord Lilford (p).
Lord Lilford (p).
Dr. A. Giinther (p). .
Budapest Museum (e).
Lord Lilford (p).
Lord Lilford (p).
0. Thomas (p).
Seville Museum (e).
Lord Lilford (p).
Florence Museum (e)
11.
3.
40.
96.
27
. 1.
11.
3.
41.
11.
3.
33.
1.
5.
22.
1-3.
11.
3.
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11
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3.
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32.
87.
7.
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3.
392.
11
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39.
11
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38.
98
2.
1.
5.
99
1.
9.
7.
95
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11
1.
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32.
66
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6.
94
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4.
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NYCTALUS LEISLERI Kuhl.
1818. Yc&pcrtilio leisleri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 46.
1818. Vcspcrtilio dasycarpos Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 49 (Alternative name for leisleri).
1839. Vcspcrtilio pachygnathus MichaheUes in Wagner, Schreber's Saug-
thiere, Suppl.', i, pi. lv b (Dalmatia. See Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe. lxii,
Abth. I, p. 222, 1870).
1857. Vesperugo leisleri Blasius, Saugethiere Deutschlands, p. 66.
1878. Vesperugo leisleri Dobson, Catal. Chiropt. Brit. Mus., p. 215.
1910. Nyctalus leisleri Trouessart, Faune Mamm. d'Europe, p. 19.
Tijjte localUii. — Hanau, Hesseu-Nassau, Germany.
Geographical distribution. — Central Europe, west to Ireland.
Diagnosis. — Like Nyctalus noctula but not so large ; condylo-
NYCTALUS 253
basal length of skull, 15 to 16 mm. ; forearm, 39 to 43 mm. •
basal portion of fur conspicuously darkened. '
Cohmr.— The colour is usually a brown, darker than in JV
nociula, nearly the prout-brown of Ridgvvay, with or without -i
lighter, wood-brown tinge, and usually showing drab or isabelk
reflections in certain lights, the underparts not so dark as back •
hairs everywhere blackish (seal-brown) through basal half Eai-s
and membranes (dry) blackish. Two skins are a pale bufty wood-
brown throughout except for the usual dark bases of the hairs
Slcull and teeth.—Except for its smaller size and more delicate
structure the skull does not differ appreciably from that of
Nyctahis noctula. Teeth essentially as in the larger species but
crown area of upper incisors nearly equal, and lower incisor row
forming a deeper, almost V-shaped convexity.
Measurements.— Externiil measurements of adult male from
Co. Wicklow, Ireland : head and body, 60 ; tail, 39 • tibia 16 • "^ ■
foot, 8-2 ; forearm, 42 ; third finger, 78 ; fourth finger, 48 • elv
from meatus, 13. Adult female from Co. Armagh, Ireland •
head and body, 63 ; tail, 42 ; tibia, 16-6 ; foot, 8 ; forearm 4"' •
third finger, 76 ; fifth finger, 49 : ear from meatus, 14 • width
of ear, 14. Two adults from Welford, Warwickshire, Ent^knd •
forearm, 40 ■ 4 and 41 • 4. Adult from Meiringen, Bern, Switzer-
land : forearm, 40. For cranial measurements see Table
p. 255. '
Spcciviens examined. ~^\e\m, from the following localities •—
Ieeland: Belfast, 1; Bray, Co. Wicklow, 2- Co Armacrh ">
England: Mexbro', Yorkshire, 1; Welford, Warwickshire! 2 ■ Cleave
Priory, 1 ; no exact locality, 1. ' ^^^"'^'^
RouMANiA : Bustenari, Prahova, 1.
Switzerland : Meiringen, Bern, 1.
JJemarfc.— Though its smaller size is the most obvious
character of this species as compared with JV. noctida the
bicolored fur, the difierent relative sizes of the upper incisors
and the more abrupt convexity of the mandibular incisor series
would by themsehes be quite sufficient to distinguish it.
9. Belfast Ireland. G. Barrett-Hamilton (p.) 1. 3. 15 1
6 a . Bray, Wicklow. J. E. Harting (c & p).' gO. 2. 14 1
9a. Armagh. Dr. G. E. Dobson (c & p). 89.11.12.5
2 9 wTf?!'^ w •,.,• R-M.Barrington(c&P). 74.5.28.8.'
2y. Welford, Warwickshire, Tomes CoUection 7 1 1 SSfi-
England. 'sS7
6. Cleeve Priory, Warwick- Tomes Collection. 7. 1. 1. 388.
1. England. Dr. Leach (p). 63 a
6. Bustenari, Prahova, 840 m. Lord Lilford (p) 44fifi
Roumauia. (ir.Z>o(/.>,o«.) *• ^. o. o.
1. Meiringen, Bern, Switzer- Tomes Collection. 7 1 1 sqq
land. I. J.. J., ooy.
254
CHIROPTERA
NYCTALUS AZOREUM Thomas.
1901. Pferygistes azoreum Thomas, Ann. and Mag. Nat. Hist., 7th ser., viii,
p. 33, July, 1901. Type in British Museum.
Type locality. — St. Michael, Azores.
Geograjjhical description. — Azores.
Diognosis. — Smaller than Nyctaliis leisleri (condylobasal length
of skull, 13 to 14' 2 mm. ; forearm, 35 to 42 mm.) ; colour darker
than in the Continental European species.
External characters. — Except for its conspicuously smaller
size Nyctalus azoreuni does not differ appreciably in external
features from N. noctula. The fifth finger shows a tendency to
be less shortened relatively to length of forearm, but this is appa-
rently not constant enough to be regarded as a specific character.
Colour. — Upper parts jjrout-brown, the hairs with darker,
ill-defined slaty basal area, and usually with lighter tips, the
lighter colour ranging in diflerent individuals from raw-umber
through wood-brown to a dull light buft', and usually producing
a noticeable wash from shoulders backward, particularly near
edge of membranes ; underparts not essentially different from
back ; muzzle, cheeks, ears, and wings blackish.
Skull and teeth. — The skull does not differ appreciably in form
from that of N. noctula, though it is perhaps slightly narrower
and deeper, and the surface is more smoothly rounded off;
lambdoid crest slightly developed. Teeth essentially as in the
larger species, but transverse diameter of m^ relatively greater,
crown area of upper incisors nearly equal, and lower incisor row
forming a more abrupt convexity.
Measurements. — Type (adult male, St. Michael) : head and
body, 54 ; tail, 42 ; tibia, 17 ; foot, 7 -7 ; forearm, 37 ; thumb, 6 ;
third finger, 62; fifth finger, 43 • 7 ; ear from meatus, 12. Adult
male from Terceira : head and body, 50 ; tail, 43 ; tibia, 1 6 ■ 6 ;
foot, 7 ; forearm, 42 : thumb, 6 ; third finger, 72 ; fifth finger,
48 ; ear from meatus, 12 ; width of ear, 1 1. For cranial measure-
ments see Table, p. 255.
Specimens examined. — Seventeen, from the following localities in the
Azores :— Terceira, 4 (B.M. and U.S.N.M.) ; above Magdalena, Pico, 5 ; St.
Michael, 6 (B.M. and U.S.N.M.) ; St. George, 2.
Remarks. — The Azorean Nyctalus is slightly more differentiated
from N. leisleri than the Continental forms are among themselves,
since it differs from the other members of the genus rather
noticeably in colour as well as in size. The specimens show
considerable variation in the extent of the light wash on upper
parts caused by the pale tips to the longer hairs, but this appears
to be independent of locality.
6 al. St. Michael, Azores. F. Du Cane Godman 65. 10. 2. 1.
(c & p). {Type of species.)
2d, 2 9. St. Michael. (W. R. 0. Hon. W. Rothschild 3.6.5.1-3,7.
Grant.) (i").
255
a
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„ slightly worn.
,, not worn.
„ slightly worn.
„ not worn.
„ not worn.
,, slightly worn.
,, not worn.
,, slightly worn.
„ not worn.
,, slightly worn.
,, moderately worn.
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256 CHIROPTERA
6,29. Reguinho, Terceira. Hon. W. Rothschild 3.6.5.4-6.
{W. R. O. Grant.) (p).
2^,3?. Magdalena,Pico. (T7.B.0. Hon. W. Rothschild 3.6.5.10-14.
Grant.) (p).
6,9. Calheto, St. George. Hon. W. Rothschild 3.6.5.8-9.
(TF. B. 0. Grant.) (p).
Genus PLECOTUS Geoffroy.
1816. ? Macrotiis Leach, Catal. Spec. Indig. INIamm. and Birds Brit. Mus.
(Willoughbv Society reprint, 1882), p. 1. Nomen nudum : "Euro-
pean Longear, Macrotiis eiirojuieus."
1818. Plecoiits Geofiroy, Description de I'Egvpte, ii, p. 112. For date see
Sherborn. Proc. Zool. Soc. London,' 1897, p. 288.
1857. Plccotus Blasius, Saugethiere Deutschlands, p. 38.
1878. Plecotus Dobson, Catal. Chiropt. Brit. Mus., p. 177 (part).
1907. Plecotus Miller, Families and Genera of Bats, p. 224, June 29, 1907.
Type species. — Vespertilio auritus Linnajus.
Geographical disfrihution. — Temperate Europe, Asia and
northern Africa.
Characters. — Dental formula : * ^^^, c Jf?, jj»i 70,, m ^^ = 36.
Skull with large, elongate and rounded brain-case, and slender
but normally formed rostrum ; auditory bulliB larger than in any
other European member of the family, their greatest diameter
equal to about three times the distance between them. Ears
very large, much longer than head, joined across forehead ;
no.strils opening upward, their orifices continued backward by
slit-like prolongations, the wart-like outgrowths on muzzle not
specially prominent.
Bemarks. — The genus Plecotus is at once recognizable among
European bats by the very large ears, joined across forehead.
In the only other European genus with 36 teeth {Miiiiopterus)
the ears are unusually short. Seven species are known, one of
which occurs in Europe.
PLECOTUS AURITUS Linnseus.
1758. Vespertilio auritus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden).
1816. ? Macrotus europieus Leach, Catal. Spec. Indig. Mamm. and Birds
Brit. Mus. (Willoughby Society reprint, 1882), p. 1. Nomen
nudum: " European Longear, Macrotus europpeus."
1818. Plccotus auritus Geoffrey, Description de I'Kgypte, ii, p. 118.
1825. Vcsiwrlilio otus Boie, Isis, p. 1206 (Copenhagen, Denmark).
1826. Vespcrtilio cornutus Faber, Isis, p. 515 (Jutland, Denmark).
1827. Plecotus communis Lesson, Man. de Mammal., p. 95 (France).
1828. P[lecotus'] brevivianus Jenyns, Trans. Linn. Soc, London, xvi, p. 55
(Grunty Fen, Isle of Ely, Cambridgeshire, England).
1829. Plecotus vulgaris Desmarest, Faune Frau(jaise, p. 18 (Prance).
PLECOTUS 257
1829. [VcsjKrtilio am-itus] y auslrincus Fischer, Synops. ]\Ianiin., p. 117.
Based on the "second variety" of Desmarest, Diet, des Sci. Nat.,
LViii, p. 51, 182'J (Vienna, Austria).
1832. riccotus velatus I. Geoffroy, Gu^rin's Mag. do Zool., ii, CI. i, pi. 2,
p. 5 (not numbered), footnote (Name api)licd by lapsus calami to
tlie common Plccotus of England).
1810. Plrcotiis megalotos Schinz, Europ. Fauna, i, p. 19 (Synonym of
auritus ; lirehm cited as authority).
1857. I'lecotus auritus Blasius, Siiugethiere Deutschlaiids, p. 39.
18G0. Plccotus kirschbanmii Koch, Achter Ber. Oberhess. Gesellsch.
Natur.- u. Heilkunde, Giesson, p. 40, May, 1860 (Dillenburg,
Oberhessen, Germany).
1863. [Plccotus auritus] var. ttjinis Koch, Jahrb. des Vereius fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 406 (Wiesbaden, Nassau,
Germany).
1863. [Plccotus auritus] var. montanus Koch, Jahrb. des Vercins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 406 (Westerwald, Nassau,
Germany).
1863. [Plccotus auritus] var. brcvipes Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 407 (Substitute for kirscli-
bauniii Koch).
1878. Plccotus auritus Dohson, Gatal. Chiropt. Brit. Mus., p. 178.
1910. Plccotus auritus Trouessart, Faune Mamm. d'Europc, p. 12.
Type locality. — Sweden.
Geographical distribution. — From Ireland ea.stward into Asia,
and from the Mediterranean north to the Scandinavian Peninsula.
Diagnosis. — Characters as in the genus ; auditory bulhe
not excessively enlarged ; forearm about 37 mm.
External characters. — General form slender and delicate, the
most conspicuous feature the greatly enlarged ears, the super-
ficial area of which together when outstretched is considerably
greater than that of entire body. Muzzle rather nari'ow, its
width at middle about equal to distance from tip to point on
forehead at middle of base of joining membrane of ears ; glandular
swellings not conspicuous, but extending up over sides to median
line behind nostrils, and backward to eye. Nostrils opening
upward and slightly foi'ward, their orifices crescentic with slit-
like posterior elongation ; space between nostrils flat, crossed
at middle by a transverse groove with overhanging anterior edge.
Lips moderately full, smooth. Ear very large, extending about
20 mm. beyond tip of muzzle when laid forward, its length fi-om
meatus nearly equal to that of forearm : general outline of conch
a simple, rater elongate oval, the anterior and posterior borders
evenly convex, the tip rather narrowly rounded ; anterior bases
joined across forehead, the membrane at point of junction about
2 • 5 mm. high ; outline of anterior border broken about 3 mm.
above base by conspicuously projecting, evenly rounded lobe
2 mm. long by 3 mm. wide at base ; antitragus scarcely indicated,
the posterior border of conch terminating abruptly a little behind
level of eye ; inner surface of conch with about twenty faint
cross striations ; joining membrane thinly furred ; a hairy line
s
258 CHIROPTERA
parallel with anterior border, and minute scattered tufts on other
parts of inner surface ; tragus simple, erect, about half as high
as conch, the anterior border straight below, slightly convex
above, the tip narrowly rounded, the posterior border faintly
concave above, distinctly convex below, greatest width (through
anterior base) ecjual to a little less than half length of anterior
border ; posterior basal lobe well developed though not large.
Wing broad, the fifth finger exceeding forearm by more than
t)ne-third length of latter, the membrane thin and semi-ti'ans-
parent, joining foot at base of outer toe ; third and fourth meta-
carjials equal, about 2 mm. shorter than forearm, fifth slightly
shorter than fourth ; leg slender ; foot about half tibia ; calcar
slender but sharply defined, equal in length to tibia and to free
Ijorder of inteiiemoral membrane, its distal termination marked
by a slight lobe, its keel obsolete. Tail extending about to point
of juncture of ears when laid forward, the extreme tip (about
1 • 5 mm.) free.
ShuU. — Among the European members of the order the
skull of Plecotus is noticeable for its large, rather low, smoothly
rounded brain-case, relatively small, weak rostrum, and much
inflated auditory bulliie. Dorsal profile rising gradually from front
of nares to middle of anterior portion of brain-case, with slight
concavity at interlachrymal region, then falling gradually to
low, strongly convex, slightly overhanging lambda ; ventral
profile scarcely elevated posteriorly. Brain-case large but rather
low, the depth at middle only a little more
than half mastoid breadth, the general out-
line when viewed from above squarish with
rounded occipital projection and abrupt
anterior constriction, the surface smoothly
rounded, with slightly indicated sagittal ridge
anteriorly and very low, short lambdoid crest,
the latter situated unusually close to upper
edge of foramen magnum ; an irregular area
in supramastoid region thickly sprinkled with
small vacuities ; base of brain-case smooth,
without sj^ecial features, a narrow groove at
„, V" ' ■. each side of basioccipital ; auditory bulla
Nat. size. relatively larger than in any other European
bat, its diametei- about three times the
space between Inilhe. Interorbital constriction narrow, scarcely
hour-glass shaped, the anterior upper rim of orbit with narrow,
slightly jirojecting edge ; lachrymal region scai'cely half as wide
as brain-case. Rostrum both narrow and short, the narial
emargination, though not unusually large, extending slightly
more than half way back to lachrymal level, its general outline
oval, sometimes flattened posteriorly ; dorsal surface of rostrum
with slight median crease, the sides evenly rounded off ; rostral
depth at front of orbit about equal to distance from orbit to
PLECOTUS 259
aKoulus of inner incisor ; auteorbital foramen small, close to I'ini
of orbit, its orifice over parastyle of first molar ; lachrymal fora-
men slightly above and behind it, on inner edge of orbit. Palate
rather broad and short, strongly concave both laterally and
longitudinally, the anterior emargination small, scarcely extend-
ing back to level of middle of canine ; posterior extension of
palate squarish, slightly longer than wide, with slightly developed
median spine ; mesopterygoid space somewhat wider than deep.
Zygoma slightly but evidently expanded at middle. Mandible
slender, but noticeably deeper at symphysis than behind tooth-
row ; coronoid process moderately high, the upper margin of
posterior section of mandible oblique-concave ; angular process
ratlier heavy, scarcely or not expanded at tip.
Teeth. — The teeth are rather small relatively to size of skull.
Inner upper incisor about half as high as canine, with large
secondary cusp slightly above middle of shaft, the outline of
crown oval, its long axis in line of tooth-row ; outer upper incisor
about half as high as inner, its apex a little below secondary cusp
of larger tooth, its secondary cusp well developed, on inner side, the
outline of crown ovate, with long axis perpendicular to tooth-row
and narrower extremity directed inwards. Lower incisors closely
crowded but slightly imbricated, the row broadly (J shaped ; the
crown area increases regularly from first to third, by addition to
posterointernal portion ; cutting edge obscurely trifid ; /g with
well developed postero-internal tubercle. Upper canine small,
scarcely higher than main cusps of molars, its shaft nearly terete,
with slightly developed posterior cutting edge, on each side of
which lies an evident groove ; cingulum complete, without cusps.
Lower canine small, slightly exceeding molars in height, its
shaft concave posteriorly, flattened interiorly, evenly convex
antero-externally ; cingulum well developed, foi'ming a prominent
antero-internal secondary cusp. Anterior upper premolar
perfectly in tooth-row, about as large as outer incisor, though
more robust, its crown sub-terete, about one-third that of canine
in basal area, its shaft with evident posterior cutting edge ; large
premolar with crown area about two-thirds that of first molar,
the inner side very narrow and without crushing surface, the
antero-internal cusp slender but well developed. Crown area of
anterior lower premolar abovit half that of canine, that of
posterior premolar nearly ecjual to that of canine, that of middle
premolar a little more than half that of first ; first sub-terete,
second oval with long diameter of crown lying across tooth-row,
third scjuarish with antero-external corner rounded oflf ; cusp of
first about half as high as canine, that of second a little shorter,
that of third a little longer ; cingulum well developed but not
forming true secondary cusps. Upper molars narro\\' internally,
the jirotocone with rather short base, the hypocone absent or
barely indicated ; metacone decidedly higher than paracone ;
styles and commissures well developed, though mesostyle of j«'^
s 2
260
CHIKOPTERA
does not extuiul outward to level of parastyle and nietastyle ;
Wpattern normal ; m-' with three cusps and three commissures,
its crown area about half that of m'-', its transverse diameter
through metacone about half length of anterior border. Lower
molars with no special peculiarities except that protoconids and
inner cusps are unusually high and slender ; ciugulum forming a
barely indicated postero-internal cusp behind entoconid ; second
triangle of riu much narrower than tirst, but about equal to it
in area.
3Icnsurcmeiiis. — Adult female from Chelmsford, Essex,
England: head and body, 42; tail, 45; tibia, 17; foot, 9*6;
forearm, 39 ; thumb, 7 ; third hnger, 69 ; fifth finger, 52 ;
ear from meatus, .'56 ; width of ear, 23 ; tragus, 19. Two adult
males from Silos, Burgos, Spain: head and body, 50 and 51 ;
tail, 46 and 46 ; tibia, 18 and 17 ; foot, 8 and 8-4 ; forearm,
41 and 38; thumb, 7' 2 and 7 •4; third finger, 72 and 70; fifth
finger, 57 and 53 ; ear from meatus, 35 and 37 ; width of ear,
26 and 26 "4 ; tragus, 17 and 18. Adult female from La Granja,
S(•go^ ia, Spain : head and body, 45 ; tail, 44 ; tibia, 20 ; foot, 9 ;
forearm, 40 ; thumb, 9 ; third finger, 72 ; fifth finger, 53 ; ear
from meatus, 37 ' 4 ; width of ear, 23 ; tragus, 17 ■ 6. Adult female
from I'lorence, Italy : head and body, 49 ; tail, 48 ; tibia, 19" 2 ;
foot, 9; forearm, 39-4; thumb, 7-8; third finger, 69; fifth
finger, 54 ; ear from meatus, 34 ; width of ear, 23*4 ; tragus, 17.
For cranial measurements see Table, p. 262.
Specimens examined. — Eighty, from the following localities : —
Scotland : Montrose, Forfar, 4.
Ireland : Antrim, 1 ; Co. Longford, 2.
England: Alderley, Cheshire, 1; Bowdon, Cheshire, 1; Diss, Norfolk, 1;
Winfarthing, Norfolk, 1 ; Woburn Sands, Bedfordshire, 1 ; Pembrokeshire,
no exact locality, 1 ; Chelmsford, Essex, 1 (U.S.N.M.) ; Tring, Hertford-
shire, 1 ; Boxniore, Hertfordshire, 1 ; Hoddesdon, Hertfordshire, 1 ;
London, 4 ; Gtodalming, Surrey, 1 ; Surrey, no exact locality, 1 ;
Bonchurch, Isle of Wight, 1; near Honiton, Devonshire, 2 ; Devonshire,
no exact locality, 8 ; no exact locality, 1.
Holland : Oosterlieek, Guelderlaud, 4.
France : Near Barcelonnette, Basses- Alpes, 1 (Mottaz).
Germany : Wernigerode, 3 ; Ummerstadt, Thiiringen, 2 ; Magdeburg, 2 ;
Niesky, Silesia, 4 ; Damsdorf , Silesia, 1.
Austria-Hungary : Csallokoz-Somorja, Pressburg, 7.
Switzerland : Geneva, 2 (Mottaz) ; Lausanne, Vaud, 1 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Boudrv. Neuchatel, 2 (Mottaz) ; Teufen,
Appenzell, 1 (U.S.N.M.); Thurgau, St. Gallen, 1; St. Gothard, Uri, 3
(U.S.N.M.) ; no exact locality, 1.
Italy: i3orzoli, near Genoa, 1 (Genoa); Florence, 1 (Mottaz) ; Kimini,!;
Rome, 1 ; no exact locality, 1 ; Sicily, 1.
Spain : Silos, Burgos, 2 (U.S.N.M.) ; La Granja, Segovia, 1.
IteniarJcs. — Plecotus auritus is so readily distinguished from all
other European bats by the great size of the ears that no sj^ecial
comparisons are required.
6, 3 ?. Montrose, Scotland. J. H. Coward (c & p). 11. 1. 3. 48-51
6 St. Antrim, Ireland. Hon. N. C. Rothschild 1. 9. 3. 5.
PLECOTUS 261
2 6 al. Longford. Dr. G. E. Dobson (c & p). 76. 2. 12. 2.
6. Pembroke, Wales. W. E. dc Winton (c & p). 11. 1. 3. 393.
1 St. Winfarthing, Norfolk, E. Markham (c & p). 9G. 7. 27. 3.
England.
5. Diss, Norfolk. (IF. R. C. H. B. Grant (p). 11. 1. 3. 394.
Slwrrin.)
1 al. Woburn Sands, Bed- A. Death (c & p). 7. 9. 2. 1.
fordshire.
9. Boxmore, Hertford- H. Piffard (c & p). 11. 1. 3. 55.
shire.
1 St. Hoddesdon, Hertford- A. Chittenden (c & p). 80. 9. 27. 1.
shire.
6. Tring, Hertfordshire. Dr. E. Hartert (c & p). 11. 1. 3. 50.
9 al. London. No history.
i. London. Sergt. Brown (c & p). 11. 1. 3. 57.
6. S.Kensington, Middle- 11.1.3.54.
sex.
i. Godalraing, Surrey. W. T. Blanford (c & p). 11. 1. 3. 58.
1. Bonchurch, Isle of Rev. C. A.Biuy (c & p). 41.3.29.0.
Wight.
S,9. Honiton, Devonshire, G. C. Shortridge (c & p). 11.1.3.52-53.
300 ft.
4 (5, 4 9 al. Devonshire. No history.
2 .5, 2 9. Oosterbeek, Guelder- O. Thomas (c & p). 98. 2. 1. 1-4.
land, Holland.
9. Wernigerode, Harz, W. Wolterstorff (c & p). 0.2.8.1.
Germany.
6, 9. Wernigerode, Harz. Lord Lilford (p). 11. 1. 1. 45-46.
{Wolterstorff.)
5. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 47.
(Wolterstorff.)
<J, 9. Ummerstadt, Thiirin- Lord Lilford (p). 11.1.1.43-44.
gen. (Schuchardt.)
i. Niesky, Silesia. (IF. Lord Lilford (p). 99. 1. 9. 1.
Baer.)
(< al. Damsdorf, Silesia. Dr. A. Giinther (p). GO. 8. 1. 10.
7. C sail 6 k 6 z-Somorja, Budapest ^Museum (e). 94.3.1.5-11.
Pressburg, Austria-
Hungary.
6. Thurgau, St. Gallen, 0. Thomas (p). 4. 4. 5. 14.
Switzerland. {Zolli-
kofer.)
1. Switzerland. Leon O. Galliard (p). 75. 9. 20. 2.
9 al. Rimini, Italy. Florence IMuseum (e). 85. 7. 6. 2.
6. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 15.
Skeleton Italy. {Prince Bona- Tomes Collection. 7. 1. 1. 728.
(without parte.)
skull).
9 al. Sicily. Purchased (Parzudaki). 52. 2. 26. 18.
1 al. La Granja, Segovia, jM. de la Escalera (c). 8. 7. 30. 5.
Spain.
1. Europe. Leyden ]\Iuseum (e). 37. 4. 28. 25.
262
CHIROPTERA
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BARBASTELLA
2G3
Genus BARBASTELLA Gray.
1821. Barbastella Gray, London Med. Repos., xv, p. .300, April 1, 1821.
1825. Barbastdlns Gray, Zool. Journ., ii, p. 243, July, 1825.
1839. Synotus Keyserling and Blasius, Wiegmann's Archiv fiir Naturgescb.,
1889, I, p. -305.
1857. Synofiis Blasius, Saugethiere Deutschlands, p. 42.
1878. Synotus Dobson, Catal. Cbiropt. Brit. Mus., p. 175.
1897. Barbastella Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 375,
October, 1897.
1907. Barbastella Miller, Families and Genera of Bats, p. 223, June 29,
1907.
Type species. — VespertiUo harhasteUiis Bchreber.
Geographical distribution. — ^Northern Africa ; central and
southern Europe ; west-central Asia to the Himalayas.
Characters. — Dental formula : i '^J, c ^~, pm '—, m || = 34.
Skull with rather long, rounded brain-case and weak rostrum, the
upper surface of which is occupied by a shallow, flattened-concave
area extending from nares to faintly developed supraorbital
ridges ; auditory bullaj not specially enlarged. Ears broad and
short (laid forward they reach slightly beyond tip of muzzle),
joined across forehead ; nostrils opening upward and outward
on a flat median space between two high lateral swellings and
behind a prominent median pad.
Bemarhs. — The short broad ears joined together across fore-
head distinguish this genus from the other European membei-s of
the family Vespertilionidse. Two species are known, one peculiar
to the Himalayan region, the other occurring in Europe.
BARBASTELLA BARBASTELLUS Schreber.
1774. VespertiUo barbastellus Schreber, Siiugtbiere, I, pi. lv (description, i,
p. 168, under name: Das Kurzmaul). Based primarily on "La
Barbastelle " of Daubenton, Hist. Acad. Royale des Sci., Paris,
1759, p. 377. 1765.
1776. VespertiUo barbastelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 17 (Burgundy).
18.36. Barbastellus daubentonii Bell, Hist. Brit. Quadr., pt. i, p. 63
(Burgundy ; based primarily on Daubenton).
1838. Barbastellus communis Gray, j\Iag. Zool. and Bot., ii, p. 495,
February, 1838 (Renaming of barbastellus).
1857. Synotus barbastellus Blasius, Saugethiere Deutschlands, p. 43.
1878. Synotus barbastellus Dobson, Catal. Cbiropt. Brit. Mus., p. 176.
1897. Barbastella barbastellus Miller, Ann. and IMag. Nat. Hist., Gth sor.,
XX, p. 385, October, 1897.
1910. Barbastella barbastellus Trouessart, Faune Mamm. d'Europe, p. 11.
Type locality. — Burgundy, France.
Geographical distribution. — Central and soutliern Europe,
264
CHIROPTERA
west to England, north to southern Norway and Sweden, east
into Asia.
Diagnosis. — Characters as in the genus ; foi'earm about 38
to 40 mm.
External cliaraders. — General form slender and delicate, the
legs long, the tail about equal to head and body. Muzzle short
and broad, its width considerably greater than distance from tip
of snout to joining membrane of ears, each side with a very large
glandular mass rising above level of flat median dorsal surface
and extending downward to involve most of iipper lip behind
nosti'il pad ; glandular masses densely hairy in noticeable contrast
with the finely pubescent nostril pad and essentially naked
median region between nostrils and inner bases of ears ; nostrils
crescentic in outline, opening upward, somewhat crowded between
glandular lateral masses ; nostril pad well defined, rounded off"
above, continued downward into median portion of upper lip, the
edge of which, between glandular masses and separated from
them by evident grooves, is distinctly convex, fitting like a
valve, when mouth is closed, over bare median callosity of lower
lip. Ear large and broad, though not specially elongated, the
tijj extending about 5 mm. beyond nostril when laid forward ;
antei'ior basal lobe very small, appearing like a rudimentary
Molossine keel, the portion of anterior border usually reflexed in
Vespertilionine bats thrown forward so that its base joins its fellow
of opposite ear, the region of juncture low but sufficiently well
developed to form a distinct pocket on side next forehead ; out-
line of anterior border strongly convex excejit for an evident
flattening just below the rather abruptly rounded off" and some-
what recui'ved tip ; posterior border abruptly concave above, then
nearly straight except foi' an abruptly projecting lobe near
middle (lobe usually about 1 ' 5 mm. long "by 1 mm. wide, its uppei-
border convex, its lower border concave) ; lower border of ear full
and rounded, but with no dift"erentiated antiti'agus ; tragus large,
somewhat ti-iangular in outline, its greatest width (slightly above
level of anterior base) equal to about two-thirds length of anterior
border, its up^ier jjortion I'apidly narrowing to an attenuate tip ;
meatus with well developed keel-like ridge ; outer surface of ear
densely furred except at extreme tip and along posterior border
to a little below level of projecting lobe ; inner surface with a
hairy line marking juncture of anterior border of conch with
portion usually folded backward, elsewhere irregularly sprinkled
with fine hairs. Membranes thin and delicate, the wing broad,
inserted at base of outer toe, the antebrachial membrane con-
tinued outward as a very narrow fold to base of thumb ; third
metacarpal slightly shorter than forearm and a little exceeding
the sub-equal fourth and fifth ; fur soft and loose, the hairs on
middle of back about 10 mm. in length ; on both ui:)per and
under surface of wing the fur extends to a line joining middle of
forearm with knee ; foot less than half as long as tibia ; calcar
BAKBASTELLA
265
about as long as tibia, with small but evident terminal lobe and
posterior keel ; tail with extreme tip projecting beyond inter-
i'emoral membrane.
Coluur. — General coloui' a very dai'k brown between the seal-
brown and vandyke-brown of Ridgway, the hairs everywhere
dark brown to extreme base, those of upper parts tipped with
light glossy wood-brown producing a sharply contrasted wash
throughout I'egion behind shoulders, those of undei'parts tipped
with a paler, more drabby brown, but without producing so
decided a contrast as on back, though the wash is usually evident
along middle of belly ; hairs on l)ase of under side of interfemoral
membrane pale ecru-drab or whitish smoke-grey nearly or quite
to base. Ears and membranes in dry specimens br-own, hardly
so dark as ground colour of body.
Skull. — The skull, though lightly-built and small, scarcely
exceeding that of Myotis mystacinus in length, is rather deep and
robust, with unusually lai'ge bi'ain-case relatively to the short
rostrum. Dorsal profile abruptly convex over anterior half of
brain-case, then nearly horizontal to somewhat overhanging
occiput ; ventral profile nearly horizontal, the floor of brain-case
scarcely elevated above level of tooth-i'ow. Brain-case high and
inflated anteriorly, relatively low posteriorly, its lateral outlines
when viewed from above essentially as in the
species of Myotis, but less rounded and globular
posteriorly ; floor of brain-case with no special
peculiarities ; auditory bulke scarcely more in-
flated than in the small species of Myotis.
Interorbital region slightly constricted, its sur-
face flattened and not well defined from that of
rosti'um ; lachrymal ridges rather prominent,
especially at their lower extremities. Zygoma
nearly straight, scarcely bowed outward beyond
level of outer surface of brain-case. Ante-
orbital foramen relatively large, over anterior narbasteiia harha-
margin of m^. Rostrum short, its upper sur- stdlus. Nat. size,
face flattened-concave, deeply emarginate in
front by the unusual backward extension of the uppei- margin of
anterior nares, the posterior border of which is at level of ante-
orbital foramina, exposing anterior extremity of vomer. Palate
.short, noticeably concave both longitudinally and laterally ; its
anterior emargination small, extending slightly behind level of
middle of canine ; its posterior border slightly behind level of vi^,
double emarginate with short median projection ; mesopterygoid
space slightly wider than long. Mandible slender, with no special
peculiarities.
Teeth. — In general aspect the teeth resemble those of Plecotus
(iHvitus, though they are throughout smaller and more slender.
1 ncisois botii above and below essentially as in Plecotus auritus
except that crown of outer upper incisor is nearly terete, and
266
CHIEOPTERA
space between it and eingulum of canine is veiy narrow ; lower
incisors forming a less broadly (J -shaped row. Canines as in
P. auritus. Anterior upper premolar very minute, crowded
inward completely from tooth-row, the diameter of its crown
barely one-quarter that of outer incisor, its cusp and ciugulum
distinct ; large upper premolar much as in P. auritus but less
narrowed on inner side and with a narrow concave crushing
surface ; no antero-internal basal cusp. Lower premolars almost
exactly similar to middle and posterior lower premolar of Plecoius
auritus, but crown area of larger tooth slightly exceeding that of
canine. Molars not appreciably different from those of Plecotus
auritus except that they are smaller and the concave median
region of the upper teeth is larger relatively to area of crowns.
Measurements. — Adult male from Cheshire, England : head
and body, 48; tail, 49; tibia, 19; foot, 7- 2; forearm, 38-4;
thumb, 6 ; third finger, 69 ; fifth finger, 52 ; ear from meatus,
15 ; width of ear, 14. Adult female from Epping, Essex,
England: head and body, 49*6 ; tail, 46 ; tibia, 18*2; foot, 7 ;
forearm, 36; thumb, 5-8; third finger, 69 ; fifth finger, 51 ; ear
from meatus, 16 ; width of ear, 14. Average and extremes of
five adults from Damsdorf , Silesia, Germany : head and body,
50(47-52); tail, 47-1 (45-49); tibia, 18-5 (18-19); foot, 6-6
(6-7); forearm, 38 (37-4-39); thumb, 5-8 (5 -4-6 -2); third
finger, 70 (68-72) ; fifth finger, 52-6 (51-54) ; ear from meatus,
15-5 (15-16); width of ear, 14-1 (13-6-15). For cranial
measurements see Table opposite.
Specimens examined. — Sixty-one, from fche following localities : —
England: Elton, Huntingdonshire, 1; Ellingham, Norfolk, 1 ; Arrow
Lodge, Warwickshire, 1 ; Alcester, Warwickshire, 1 ; Kugby, Warwick-
shire, 1; Welford, Rugby, 1 ; Pilton, Oundle, Northamptonshire, 1; Epping,
Essex, 1 ; Llanelwedd, Radnorshire, 1 ; Swindon, Wiltshire, 1 ; Milton,
Hampshire, 1.
France: Caterille, Haute-Cxaronne, 1.
Germany: Biichenberg, Harz Mountains, 1; Niesky, Silesia, 4;
Bavaria, no exact locality, 3 (U.S.N.IM.) ; near Wernigerode, Saxony, 1.
Switzerland : Grotte de Vallorbe, Vaud, 40 (Mottaz).
Remarks. — This species is immediately recognizable among
European bats by its short, joined ears, and liy the peculiar form
of the muzzle.
?. Llanelwedd, Radnorshire, H. E. Forrest (c ct p). 4. 7. 3. 1.
Wales.
9. Elton, Huntingdonshire, Lord Lilford (p). 94. 9. 5. 1.
England.
i. Ellingham, Norfolk. E. R. Alston (p). 81. 6. 9. 1.
(Dr. W. M. Crowfuot.)
6. Arrow Lodge, Warwickshire. Tomes Collection. 7.1.1.348.
1. Alcester, Warwickshire. Tomes Collection. 7. 1. 1. 348.
i St. Rugby, Warwickshire. G. Dalgleish (c & p). 4. 10. 13. 2.
i. Oundle, Northamptonshire. Lord Lilford (c & p). 94. 4. IG. 1.
? al. Epping, Essex. H. Doubleday (c & p). 44. 5. 1. 1.
i. Milton, Hampshire. Rev. J. E. Kelsall (p). 11. 1. 3. 59.
BARBASTELLA
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6. Swindon, Wiltshire. Rev. E. A. Goddard 8. 5. 12. 1.
(c & p).
6. Caterille, Haute-Garonne, 0. Thomas (p). G. 4. 1. 7.
1000 ra. France.
{A. Robert.)
6, 9. Niesky, Silesia, 181 m. Ger- Dr. E. Hamilton (p). 97. 12. 4. 2-3.
many. (W. Bacr.)
2 9. Niesky, Silesia. [W. Bacr.) Lord Lilford (p). 99.1.9.2,4.
i. We mi ger ode, Saxony. Lord Lilford (p). 0.2.8.2.
(WoUerstorff.)
Sub-Family MINIOPTERINtE.
1878. Minioptcri Dobson, Catal. Chiropt. Brit. Mus., p. 170 (part).
1907. Miniopterinx Miller, Families and Genera of Bats, p. 227, June 29,
1907.
Geograj^hiral distrihution. — Africa, southern Europe and
southern Asia, eastward to the Malay region, Japan and
Australia.
Characters. — Like the Vespertilioninae, but presternum with
median lobe enormously developed and forming the greater part
of the bone ; scapula with coracoid straight, directed con-
spicuously inward.
BemarJcs. — The sub-family MiniopterhiBP, though widely dis-
tributed in the warmer portions of the Old World, is at present
known to contain the genus Minioidcrus only.
Uenus MINIOPTERUS Bonaparte.
1837. Miniopterus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx, under
Vcspcrtilio cmarginatns (Sub-genus of Ve^pertilio).
1857. Miniopterus Blasius, Siiugethiere Deutschlands, p. 45.
1860. Miniopteris Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 91,
February, 1866.
1878. Miniopterus Dohson, Catal. Chiropt. Brit. I\Ius., p. 347.
1892. Minyoptcrus Winge, Jordfundne og nulevende Flagermus (Chiroptera)
fra Lagoa Santa, Minas Geraes, Brasilien, p. 36.
1900. Minncoptcrus Lampe, Jahrb. Nassau Ver. Naturkundo, Jahrg. 53,
Catal. Saugeth.-Samml., p. 12.
1907. Miniopterus Miller, Families and Genera of Bats, p. 227, June 29,
1907.
Tijjje sperU's. — Vcspertilio urshii Bonaparte = V. srlireihersii
Kuhi
Geographical distribution. — Same as that of the sub-family.
Characters. — Dental formula : i i^'f, c ^J, pm ^~, m t;' = 34.
Skull with unusually high brain-case, and low, flattened rostrum.
Second phalanx of third finger nearly three times as long as fii'st.
Ears short, separate, the upjjer margin (in European species)
appearing almost artificially truncate.
Memarhs. — The peculiar shortening of the ears together with
the remarkable elongation of second phalan.x of thiid linger readily
MINIOPTERUS
l'G9
di,stini,'uisli tliis <((!nus among tlie inenil)ers of the Eui'opefiii fauna.
About a dozen forms have been described, one of which occurs
in soutliern Europe.
MINIOPTERUS SCHREIBERSII Kulil.
1819. Vcspcrtilio schrcibcisii Kuhl, Ann. Wetterau. Gesollsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 185 (Hungary).
1837. Vcspcrtilio ursinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Monto Corno, Ascoli, Italy). Typo in British Museum.
18-11. Vesjxriilio orsinii Temminck, Monogr. do Mamm., ii, p. 179 (modi-
fication of ursinii).
1857. Minioptcrus schrcibcrsii Blasius, Saugethiere Deutschlands, p. 4G.
1878. Miniopterus schrcibcrsii Dobson, Catal. Chiropt. Brit. Mus., p. 348.
1910. Minioptcrus schreibcrsi Trouessarfc, Fauiio Mamm. d'Europe, p. 34.
Type locnlifij. — Kuhnbazer Cave, mountains of soutliern
Bannat, Hungary.
(h'0(jraphic(d distn'hutlon. — Soutliern Europe from the Iberian
Peninsula eastward, north to Switzerland and Hungary. Limits
of range not known.
Dia/jnosis. — Charaters as in the genus ; forearm about
43 mm.
Ejctcrnal characters. — General form rather slender, with long
tail and legs, wing broad at base but conspicuously tapering at
tip, and short ears with a peculiar truncate aspect. Muzzle
rather broad, though without conspicuous glandular swellings,
its greatest width about ecjual to distance from eve to nostril ;
muzzle pad narrow, with slight median emargination, bounded
below by a low horizontal ridge which is continuous with projecting
inner margin of nostril. Eyelids noticeably glandular-swollen ;
a deep horizontal groove in cheek below eye. Ear short, extend-
ing about half way from eye to nostril when laid forward, its
general aspect difierent from that of any other European bat,
owing to the length of the anterior basal lobe, the short, straight
anterior border, and the broadly, evenly convex posterior border
which joins anterior border in such a manner that there is
l)ractically no " tip," the whole anterior border appearing like an
oblicjuely, almost artificially truncate extremity ; antitragus low,
obscurely marked ofli' from posterior border of conch, practically
continuous with lower lip anteriorly ; inner surface of conch
slightly rugose, without evident cross ridges ; tragus about half
as high as conch, a little curved forward owing to .slight con-
cavity of anterior border, the blunt tip and upper half of e.xterior
border forming a uniform, rather noticeable convexity, the
posterior margin straight below to rudimentary basal lobe ;
greatest width of tragus about half anterior border. Wing rather
wide basally, the tifth finger exceeding forearm by about one-tifth
length of latter, the tip unusually slender and elongate owing to
the great length of last bone of third tinger ; third and fourth
"270 CHIROPTERA
metacarpals sub-e(|iial, about 3 mm. shorter tlian forearm ; tifth
metacarpal about 4 mm. shorter than third ; membrane joining
leg at or a little above ankle. Leg rather slender, the foot
scarcely half as long as tibia ; calcar about as long as tibia and
nearly equal to free border of uropatagium, its distal termination
very obscure, its posterior border without trace of keel. Tail
about as long as head and bod\', included in the membrane to
extreme tip.
Fur and colour. — The fur is of a Aery soft, silky texture,
though rather short, the hairs at middle of back only about
7 mm. in length, those of head abruptly nuich shorter (about
4 mm.) in rather noticeable contrast. Above it scarcely extends
on membranes except for a sparse pubescence on basal half of
median portion of uropatagium ; below it reaches line joining-
middle of forearm with knee and continues across extreme base
of uropatagium. General colour of upper parts drab, faintly
lighter and more nearly hair-brown anteriorly ; underparts ecru-
drab. The hairs of back show three evident though not strongly
contrasted colour-bands : (o) at base (3 nnii.), a slaty mouse-grey ;
(h) at middle (tips of shorter hairs, 2 mm.), ecru-drab ; and (c-) at
tip (extremities of longer hairs, 2 mm.), drab. Muzzle, ears and
membranes brown, scarcely darker than body.
Skull. — The skull differs from that of all other European bats
in the great inflation of the anterior portion of brain-case and
the consequent very abrupt angle at which the forehead rises
above the low, flat rostrum. In other respects the general aspect
of skull, especially when viewed from above, is rather slender
and lightly built. Dorsal profile rising gradually from front
of nares to inter-lachrymal region, then
abruptly at an angle of nearly 45 to middle
of anterior portion of brain-case which is
nearly or quite as high as lambda, the region
£<^^ between these two highest points occupied
by a .shallow concavity corresponding to
anterior edge of interparietal ; below and
behind lambda the supraoccipital bulges
noticeably outward, but not sufficiently to
conceal condyles when skull is viewed from
directly above ; ventral profile nearly flat,
*'"'• *^- slightly elevated posteriorly. Brain-case
iliidopterus schreibersU. in i. ii i.T iT i t i
Nat. size. broadly ovate, the outline broken at each
side posteriorly by the slightly projecting
mastoid region ; sagittal crest low Ijut evident anteriorly,
obsolete posteriorly ; lambdoid crest moderately developed at
sides, barely indicated at middle ; general outline of skull when
viewed from behind squarish, the depth through bulla about
equal to breadth of bi'ain-case ; floor of brain-case with no special
peculiarities ; between cochlea and median line a shallow groove
with abrupt pit-like anterior termination a noticeable ridge
MINIOPTERDS
271
at outer side of each pit ; auditory bulla moderately large, the
transverse diameter about equal to distance between bullte.
Interorbital region moderately constricted, slightly hour-glass
shaped, the lachrymal region scarcely more than half as wide
as brain-case, smoothly rounded at sides ; rostrum tapering
gradually, rounded oft" at sides, ftattened concave along median
line, the narial emargination small, squarish, extending backward
about one-third of the distance to front of forehead ; rostral
depth at fi'ont of orbit about equal to distance from orbit to
outer incisor ; anteorbital foramen rather lai'ge, directly above
small pi-emolar, the anteorbital canal half as long as rostrum,
thus much longer than in any other European bat ; lachrymal
foramen just outside of orbit, on level with uppei- border of
anteorbital foramen. Palate long and wide, distinctly concave
both laterally and longitudinally, terminating rather abruptly a
little behind level of last molar, a small but evident foramen at
each side near posterior edge ; median sjiine large ; mesoptery-
goid space squarish, slightly wider posteriorly than anteriorly,
the short hamulars bent inward. Mandible slender, a little
deeper at symphysis than behind tooth-i'ow, the posterioi' seg-
ment vmusually small, with nearly horizontal, slightly concave
upper border ; angular process relatively long, expanded at
outer end.
Teeth. — Relatively to size of skull the teeth are small and weak.
Inner upper incisor low, the crown very oblique, with postero-
external concavity and small posterointernal cusp ; outer upper
incisor considerably larger than inner, the crown flattened in
axis of tooth-row, the width of its flattened-conca^•e posterior
surface about half its height, that of its outer border scarcely
one-fifth height, cingulum obsolete, but forming a minute postero-
external cusp ; the two teeth lie in curve of anterior portion of
tooth-row, and the outer is separated from canine by a space
about equal to its greatest diameter. Lower incisors closely
crowded but not imbricated, the crown-area increasing regularly
from first to third, the outline of the row as a whole broadly
V-shaped ; i^ and i., with crown very low, the cutting edge
t>bscurely tritid, the crown of each tooth wider posteriorly (in line
of tooth-row) than anteriorly, but this more evident in second
tliau in first ; /.j nearly terete, the middle and posterior cusps
eidarged and separated by a deep groove, the anterior cusp
reduced to a mere rudiment. Upper canine slender and weak,
sub-terete, but distinctly flattened on inner side, the shaft with
distinct anterior and posterior longitudinal grooves but without
well developed cutting edge, the cingulum narrow, complete,
but without cusps ; lower canine scarcely higher than main
cusps of molars, its inner and posterior surfaces flattened, its
antei'o-external surface smoothly rounded, cingulum forming a
low but evident antero-internal cusp. Anterior upper premolar
with crown area about equal to that of canine, its general form
272 CUIKOPTERA
essentially as in large premolar except that the antero-internal
basal cusp is barely indicated, the cutting edges of the main cusp
are less developed, and the crown is narrower extei-nally than
internally ; large upper premolar with crown area about equal
to that of first molar and double that of small premolar, its
anterioi' and postei'ior borders very slightly concave, the antei'O-
internal basal cusp small but evident, the main cusp with well
developed anterior and posterior cutting edges, and antero-internal
and postero-external longitudinal groove ; lower premolars
essentially alike in form, the crown area of the posterior tooth
about equal to that of canine, that of the two others successively a
little less, the outline S(iuarish, with antero-external angle tending
to become I'ounded off", jiarticulai'ly in the posteiior tooth ; cusp of
first about half as high as canine, that of second slightly higher
than tii'st, that of third nearly as high as main cusps of molars ;
cingulum well developed, forming in each tooth a small but
evident antero-internal basal cusp. First and second upper
molars with posterior border- deeply and almost angularly
emarginate near iniier edge of tooth, the anterior border nearly
straight ; protocone low but rather broad ; hypocone clearly
indicated but not distinct from posterior commissure of jirotocone ;
pai'acone and metacone nearer outer edge of crown tlian usual,
leaving a wide, pit-like median concavity, metacone slightly the
higher of the two cusps ; styles and commissures well developed,
Wpattern normal ; third upper molar with crown area about
half that of first, its width through metacone slightly less than
half length of anterioi- border, its three cusps and three
commissures normally developed. Lower molars with no special
peculiarities, the second triangle oi vi.^ narrower than first, but
of about the same area.
Measurements. — Adult male and female from Neuchatel,
Switzerland : head and body, 59 and 59 ; tail, 58 and 60 ;
til)ia, 20 and 19 ; foot, 10 and 9-4 ; forearm, 45 and 44-6 ;
thumb, 7 and 7 ; third finger, 92 and 92 ; fifth finger, 53 and 54 ;
ear from meatus, 12-6 and 12 ; width of ear, 12 and 12. Adult
male and female from Corinth, Greece : head and body, 57 and
56; tail, 57 and 55; tibia, 18-6 and 19; foot, 10 and 9-6;
forearm, 44 and 45 ; thumb, 6 • 6 and 7 • 2 ; third finger, 92 and
90 ; fifth finger, 54 and 53 ; ear from meatus, 12 and 12 ; width
of ear, 12 and 12. Two adult males from Silos, Burgos, Spain :
head and body, 60 and 60 ; tail, 60 and 57 ; tibia, 19 and 19-4 ;
foot, 11 and 9*6 ; forearm, 44 and 44 ; thumb, 7-4 and 7 4;
third finger, 93 and 89 ; fifth finger, 55 and 56 ; ear from meatus,
11-4 and 11-6; width of ear^ 13-4 and 12-6. Two adult
females from the same locality : head and bod}^ 59 and 59 ; tail,
60 and 59 ; tibia, 19-6 and 20 ; foot, 9-8 and 10 ; forearm,
45-4 and 46 ; thumb, 8 and 7-4 ; third finger, 93 and 9:! ; fifth
finger, 55 and 55 ; ear from meatus, 12 and 11*6 ; width of ear,
12 '6 and 12-6. For cranial measurements see Table, p. 274.
MINIOPTERUS
273
Siiecimcns examined. — Three huudred and fifty-one, from the following
localities : —
Spain : Silos, Burgos, 9 (B.M. and U.S.N.M.) ; Seville, 1 ; Minorca,
Balearic Islands, 3 ; llajorca, Balearic Islands, 2.
France: Troubate, Hautes-Pjreuces, 2; Dions, Gard, 9; Marseilles, 1
(U.S.N.M.).
Switzerland: Geneva, 235 (Mottaz) ; Neuchatel, 2 (U.S.N.M.).
Austria-Hungary : Ofener Mountains, Hungary, 1 ; Hungary, no exact
locality, 3 (U.S.N.M.).
Montenegro : Beri, 1 ; Velgi, Czolo, 1.
Italy : Western Liguria, 14 (Genoa) ; Finalborgo, Liguria, 5 (Genoa) ;
Maremma, Tuscany, 1 (U.S.N.M.); Pisa, 1; Florence, 1 (U.S.N.M.);
Livorno, 6 (U.S.N.^NI.) ; Spezia, 5 (U.S.N.M.); Monte Corno, Ascoli, 1
(tyipe oi ur. si nii Bonaparte); Rome, 10; Velletri, Rome, 2 (U.S.N.M.) ; no
exact locality, 1 ; Elba Island, 5 (U.S.N. Til.) ; Marsala, Sicily, 2.
Sardinia : Sassari, 3 (U.S.N.IM.) ; Cagliari, 4 (U.S.N.M.) ; Mount
Gennargentu, 2 (U.S.N.M.).
Greece : Corinth, 17 (U.S.N.M.) ; Labyrinth, Crete, 1.
MemnrTcs. — The lengthened second phalanx of the third finger,
the peculiar cropped appearance of the ears, and the short, dense,
velvety fur of the head are the most obvious external character-
istics of Minio])terus schreibersii as compared with other European
bats.
2.
S,9.
3S.
<?, 9.
8 <5, 1 9
1 al.
9.
9.
7 <J, 3 9.
Silos, Burgos, Spain. Rev. S. Gonzalez (c).
Inca, Majorca, Balearic O. Thomas and R. I.
Islands. Pocock (c & p).
San Cristobal, Minorca. 0. Thomas and R. I.
Pocock (c & p).
Troubate, Hautes- 0. Thomas (p).
Pyrenees, 400 m.
Prance. (A. Robert.)
Dions, Gard, 70 m. 0. Thomas (p).
France. (C. Mottaz.)
Ofener Mountains, Budapest Museum (e).
Budapest.
Beri, Montenegro.
(L. Fllhrer.)
Velgi, Czolo.
(L. Fiihrer.)
Rome. (C. Coli.)
Skeleton Monte Corno, Ascoli,
(without Italy,
skull).
2 6. Marsala, Sicily.
[A. Hubert.)
6. Labyrinth, 200 ft.
Crete.
0. Thomas (p).
0. Thomas (p).
Lord Lilford (p).
Tomes Collection.
88. 1. 5. 46-47.
0. 7. 1. 4-5.
0. 7. 1. 32-34.
6. 4. 1. 8-9.
8. 8. 10. 1-9.
94. 7. 18. 10.
5. 8. 4. G.
5. 8. 4. 5.
11. 1. 2. 5-14.
7. 1. 1. 735.
(Type of Yespertilio ursinii Bonaparte.)
0. Thomas (p). 6. 8. 4. 24-25.
Miss D. Bate (c). 5. 12. 2. 9.
274
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276
CHIROPTERA
Family MOLOSSID^.
1865. Mulossi Peters, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 258.
1872. Molossidx Gill, Arrangement of the Families of Mammals, p. 17.
1878. Emhallonuridx (part ; Molossime, part, Molossi) Dobson, Catal.
Chiropt. Brit. Mus., p. 402.
1907. Molossidx Miller, Families and Genera of Bats, p. 241, June 29, 1907.
Geograjjlncal clistrihufion.- — Warmer parts of both hemispheres ;
in the Old World north to the Mediterranean region and
southern Asia.
Characters. — Essentially like the VespertHiotiidse except that
the secondary articulation of humerus with scapula is more
perfectly developed and the fibula is robust, adding appreciably
to strength of leg. Auditory bulla noticeably emarginate on inner
side. Tail projecting very consj^icuously beyond membrane, a
character by which the only European member of the group may
be immediately I'ecognized.
BemarJcH. — The family Molossidse is widely distributed in the
warmer parts of both hemispheres. Eleven genera are now
known, one of which is represented in the Mediterranean region
of Eurojae.
Genus NYCTINOMUS Geoftroy.
1818. Nyctinomus Geoffrey, Descr. de I'Egypte, ii, p. 114 ({egyptiacus).
1821. Nyctinoma Bowdich, Anal. Nat. Class. Mamm., p. 28 (Modification
of Nyciincmiiis).
1821. Nyctinomes Gray, London Med. Eepos., xv, p. 299, April 1, 1821
(Modification of Nyctinomus).
1822. Nyctinomia Fleming, Philos. of ZooL, ii, p. 178 (Modification of
Nyctinomus).
1842. Mops Lesson, Nouv. Tabl. Regno Auim., p. IS (Mops indicus Lesson
= Dysopcs mops F. Cuvier).
1878. Nyctinomus Dobson, Catal. Chiropt. Brit. Mus., p. 420 (part).
1902. Nyctinomops Miller, Proc. Acad. Nat. Sci. Philadelphia, p. 393,
September 12, 1902 [fcmorosaccxis).
1907. Nyctinomus Miller, Families and Genera of Bats, p. 251, June 29, 1907.
Type species. — Nyctinomus segypiiacus Geoffroy.
Geographical distribution. — Warmer portions of both hemi-
spheres, north to the southern United States and to the Mediter-
ranean coast of Euroj^e, east to the Pliilipjsines and Norfolk
Island.
Characters. — Dental formula: i 'J; or y.,* c '^, jmi j;-^, m :: ;' = 30
or 32 ; brain-case not unusually flattened, its dej^th at least half
its width ; bony palate with a small median anterior emargination
extending to behind level of roots of incisors.
Reynarhs. — As thus defined the genus Nyctinomus contains
* In the species occurring in Europe.
NYCTINOMUS
277
fibout forty species, two-thirds of wliich are peculiar to the Old
World, one of them occurring in the Mediterranean region of
Europe.
NYCTINOMUS TENIOTIS llalinesque.
1814. Ccphalotes teniotis Bafinesquo, Pr6c. des D6couv. Somiol., p. 12 (Sicily).
1825. Dino2)s ccstoni Savi, N. Giom. de' Letterati, Pisa, x, p. 235 (Pisa, Italy).
1810. Dysopcs savii Schinz, Europ. Fauna, I, p. 5 (Substitute for ccstoni).
1871. [Dysopcs cestonii] var. nifjrogrisciis Schneider, Neue Denkschr.
Schweiz. Gesellsch. Naturwiss., xxiv, p. 5 (articles separately
paged). Basel, Switzerland.
1877. Nyctinomus cesionii Dobson, Catal. Chiropt. Brit. Mus., p. 423.
1891. Nyctinomus ttenioiis Thomas, Proc. Zool. Soc., London, p. 182.
1897. [Dysopcs} viidas Schulze, Abh. Ges. Nat. iv, No. 10, p. 23 (Substitute
for cestoni). Not of Sundevall, 1842.
1910. Nyctmomus txniotis Trouessart, Paunc Mamm. d'Europe, p. 36.
Type locality. — Sicily.
Gcog7-aphical flistrihuiion. — Mediterranean region of Europe
and noi'thern Africa. Accidental ? at Basel, Switzerland.
Diagnosis. — Like the African Nyctinomus segyptiacus but
lai'ger, condylobasal length of skull about 23 mm. instead of
about 20 mm. ; lower incisor 3-3 instead of 2-2 ; small upper
premolar with crown area more than one-half instead of less
than one-fifth that of upper incisor. Distinguishable among
European bats by the generic characters ; forearm about 60 mm.
External characters. — Form heavy and robust, the legs short, the
feet large, the wings long and narrow, the membranes thick and
leathery, the ears very large, sub-orbicular, joined at their anterior
bases ; these characters in connection with the thick muscular tail,
projecting by at least one- third of its length be3^ond interfemoral
membrane, immediately distinguish the animal from all other
European bats. Muzzle projecting rather noticeably beyond
upper lip, oblicpiely truncate ; nostril pad well defined, wider
than high, the sub-circular nostrils opening forward and slightly
outward at its outer margin, its surface with very fine reticula-
tions and a few coarse wrinkles, glabrous except below, where it
is sprinkled with fine hairs, this hairy area continuous with the
l)rush-like fringe of i-ecurved blunt hairs along middle of upper
lip ; upper margin of pad broadly concave at middle, convex
laterally over the nostrils, its edge thickly set with small, laterally
compressed horny excrescences, about thirty-five in number,
the outermost lying a little above level of middle of nostril ; a
row of about ten similar excrescences crosses middle of pad
vertically. Upper lip very large and full, marked by manj^ deep
ol)li([ue wrinkles, its surface both abo^■e and below rather densely
hairy. A small wart on chin just behind level of symphysis.
Ear very large, sub-orbicular, the margin with no very decided
irregulaiities, though the anterior border usually shows some
slight flattening and the posterior border is faintly concave above
278
CHIROPTERA
and neai' base ; anterior basal margin of ear with a narrow
outward-folded hem, the exposed side of which is densely
pubescent except at extreme anterior margin, where it is glabrous
and set with about six terete wart-like projections, widely and
irregularly spaced ; jiosterior basal margin with a shorter, deeper,
inwardly-folded lobe just above concavity limiting posterior
base of antitragus ; keel well developed, not thickened at edge,
its height posteriorly ecjual to about one-third its length, the
margin of keel hairy, the line thus begun curving upward and
backward under upper margin of conch ; antitragus well defined,
its height slightly greater than that of keel, its length about
twice height, its anterior border with ridge-like continuation
forward to corner of mouth ; tragus squarish in outline, its
posterior border longest and with distinct angle below middle, its
anterior border shortest, its upper border almost horizontally
truncate ; antei'ior and upper margin of tragus fringed with long
loose hairs. Wings longer and nai'rower than in any other
European bat, the membrane inserted on side of tibia just above
ankle ; antebrachial membrane extending as a narrow fold along
forearm to base of thumb ; the fur of body extends on both
surfaces of wing to line joining middle of humerus with knee,
and on to extreme base of uropatagium ; otherwise the membranes
are essentially naked except for some fine dense pubescence on
upper side of propatagium. Thumb short and robust, with
moderately developed pad at distal end of metacarpal. Foot
broad and robust, more than half as long as tibia, the sole with
a low and rounded but evident pad at middle, four small elongated
pads in a row at bases of toes, and some smaller, less definite
callosities in space between this row and the large median pad.
Outer and inner toe thickened, their outer surfaces densely
covered with short stifiened hairs with recurved points ; sprinkled
among these shorter hairs and also at the ends of the other toes
are a few much longer bristles. Calcar about as long as tibia,
its point ill-defined ; no indication of keel or of terminal lobe.
Tail about half as long as head and body, robust and muscular,
the terminal third or half pi'ojecting beyond membrane.
Fur and colour. — The fur is everywhere dense and velvety
in texture, the hairs at middle of back about 7 mm. in length,
those on throat longer and looser. Colour a uniform light drab,
with faint darker shading in certain lights, the hairs pale ecru-
drab at extreme base. Ears and membranes in dry sjaecimens
blackish. Fringes on feet and hairy lines on ears, drab like
body.
SJcull. — The skull is large, but rather slender, about equal to
that of Myotis myotis in length. In general form it is distinguish-
able among those of the bats of Europe by the depressed brain-
case, the high, somewhat tubular rostro-interorbital portion, and
the conspicuously emarginate inner side of auditory liulhe. Dorsal
profile essentially straight from nares to lambda, though with a
NYCTINOMDS
279
slight convexity over anterior two-thirds of brain-case, and a
more aV)rupt though not very conspicuous swelling posteriorly,
the two convexities separated by a narrow concavity ; general
direction of dorsal profile more nearly horizontal than in any
other European bat, since the depth of occiput through condyles
is scarcely greater than that of rostrum through anterior portion
of tirst molar. Brain-case low and wide, its depth at middle
about half mastoid breadth, its surface smooth and evenly
rounded, with faintly indicated sagittal crest posteriorly and
median groove between latei-al swellings anteriorly ; lambdal
crest evident though not high ; outline of brain-case when
viewed from above a somewhat triangular ovate owing to the
rather squarely truncate posterior border and the strong contrast
between the wide mastoid region and narrow interorbital con-
striction ; floor of brain-case with median ridge and lateral
depressions posteriorly, the basisphenoid
with two shallow but rather well-delined
pits about as large as glenoid surface ;
auditory bulhe rather large, deeply emar-
ginate on inner side so that cochlea is
conspicuously exposed, the region in
front of middle of meatus reduced to a
narrow ring barely more than one-third
as wide as meatus, the extreme anterior
border with a flange-like inward-curved
projection. Interorbital region long, sub-
cylindrical, a little wider at lachrymal
level than posteriorly, but scarcely enough
so to impart a distinctly hour-glass general
form ; least interorbital breadth slightly
greater than breadth of tubular narial
region and slightly less than that across
roots of canines. Zygoma simple, not
bent upward, its margin with a barely indicated expansion behind
middle. Rostrum proper short and deep, not w^ell difierentiated
from interorbital region, the distance from orbit to front of pre-
maxillary about equal to depth at front of in} ; lachrymal ridge
short but well defined, the small, inconspicuous anteorbital fora-
men and minute lacluymal foramen opening forward under its
anterior edge, at level of middle of large premolar ; nares with
distinctly tubular latei'al margins separated from roots of canines
by evident grooves, the dorsal emargination extending about half
way to lachrymal level. Palate moderately wide, noticealjly
concave laterally, the anterior emargination small, about as deep
as wide, its posterior border scarcely extending behind level
of front of canine ; posterior border of palate double emargi-
nate with well developed median projection, the emarginations
extending forward to level of posterior margin .of m^ ; mesop-
terygoid- space large, its width anteriorly equal to that of
yycHnomus teniotU.
Nat. size.
280
CHIROPTERA
temporal fossa at sauie level, its length about twice width ;
hamulars small and inconspicuous. Mandible long and straight,
its axis scarcely bent upward posteriorly ; symphysis deep, scarcely
or not subtended by a concavity in lower border of ramus ;
posterior portion of mandible low, the depth through coronoid
process barely equal to distance from front of coronoid to back
of condyle ; angular process large, straight, directed backward,
outward and downward.
Teeth. — Relatively to size of skidl the teeth are rather large,
the lower molars in particular. The most obvious peculiarities
of the dentition as compared with that of other European bats are
the single, well developed incisor in each side of upper jaw, and
the presence oi a large hypocone, quite distinct from commissure
of jarotocone, in i«' and wi". Upper incisor simple, a little moi'o
than half as high as canine, the
shaft set at an angle so that the
points of the two teeth are much
nearer together than their bases ;
cingulum faintly indicated in front,
better developed and forming an
incipient cusp postero-externally ;
posterior surface of shaft somewhat
flattened ; space between incisor
and canine aljout equal to greatest
diameter of smaller tooth. Lower
incisors three on each side, much
imbricated and closely crowded in
narrow space at front of canines,
their cutting edge barely rising to
level ■ of canine cingulum. The
inner and middle tooth sub-equal,
rather deeplj' bifid, the outer
barely equal to outer lobe of the
others, its apex faintly notched.
Upper canine moderately large, its crown area somewhat more
than double that of upper incisor, the general outline of its
base an irregular triangle with its longest side directed
inward, and with the postero-external side slightly concave ;
cingulum narrow but well developed throughout, not tending to
develop small cusps ; lower canine with no special peculiarities,
its cingulum tending to form a slight anterior cusp. Small
upper premolar in the tooth-row, usually in contact with canine
and separated from large premolar by a narrow space, its crown
area somewhat less than that of upper incisor, its cingulum and
cusp well developed, the cusp about one-fourth as high as shaft
of canine, triangular in outline when viewed from the outside,
with well developed posterior cutting ridge ; lower premolars
similar in form but larger, their crown areas sub-equal, the shaft of
the second higher than that of first and about equal to larger cusps
FIG. 49.
Xyctiiiomus Icnlotis. Teetli X 5.
NYCTINOMDS
281
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JOJ CHIROPTERA
of lower molars ; lai'ge upper j^remolar with crown area about
two-thirds that of m}, its antero-internal cu.sp high and distinct.
First and second upper molars alike in sti'ucture, the first slightly
larger than second, the cusps all well developed and distinct,
showing no special peculiarities aside from the presence of a
conspicuous terete hypocone nearly as large and distinct as antero-
internal cusp of large premolar, and separated from commissure
of protocone by a deep notch ; third upper molar with crown
area not much less than that of second, except for the absence of
postero-internal heel and hypocone ; third commissure as long as
in the other teeth, but metacone smaller than paracone. Lower
molars with no special peculiarities, the cusps all well developed
and distinct.
Meaattroncnts. — Young adult male from Italy : head and
body, 87; tail, 56; free portion of tail, 20; tibia, 19; foot,
10' 4; forearm, GO* 4; third linger, 115; lifth linger, 65; ear
from meatus, 30' 4 ; width of ear, 32. Young adult female from
Greece : head and body, 84 ; tail, 57 ; free portion of tail, 27 ;
tibia, 20; foot, 11 "4; forearm, 61 ; third finger, 117; fifth
finger, 61 ; ear from meatus, 31 • 4 ; width of ear, 31. Two adult
females from Cintra, Portugal : forearm, 60 and 61 ; third finger,
110 and 115 ; fifth finger, 59 and 63. For cranial measurements
see Table, p. 281.
Specimens examined. — Six, from the followiug localities : —
Portugal : Cintra, 2.
Italy : No exact locality (probably from Pisa), 2. (B.M. and U.S.N.M.) ;
Sicily, 1 (U.S.N.M.).
Greece : No exact locality, 1.
2 9. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 6-7.
i. Italy (probably Pisa). Zoological Society (r). 10. 5. 24. 1.
(jSVui.)
?. Greece. Parreys. 46.0.15.121.
CAKNIVORA
283
Order CARNIVORA.
1827. Carnivora Gray, Griffith's Cuvior. Auim. KirigcL, v, p. 111.
Geoi/rnplrlcal dinfrihutinn. — Continents and larger islands of
the entire world, Australia,"' New Zealand, and the Antarctic
region excepted.
CJiaracters. — -Tei'restrial (rarely aquatic or semi-aquatic), non-
volant, placental mammals with rather high development of
brain, the cerebral hemispheres with distinct convolutions ; feet
unguiculate, never modified as fins or flippers ; dentition of a
modified tuberculosectorial tyjie, the posterior upper premolar
and antei'ior lower molar usually develojjed as special carnassial
or flesh-cutting teeth.
BcmarJcx. — The mammals of this order present much diversity
of form and structure, though less than in the case of the
Insectivora. Most of the living members of the group are
carnivorous in habits, and immediately recognizable among
placental mammals by the presence of a specially modified flesh-
tooth in each jaw. In cei'tain groups, however, as in the Ursidse
among the European representatives of the order, both habits
and dentition are of a more generalized type. The order contains
seven 7'ecent families, five of which occur in Europe.
KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES
OP CARNIVORA.
Larger cheek-teeth with crowns of a crushing type, the
cusps sub-equal, low, subterete, without noticeable
cutting edges ; upper carnassial 2-rooted, in front of
anteorbital foramen, its inner lobe posterior ; size
very large; form heavy ; feet plantigrade (Bears) ... LVsida;, p. 284.
Larger cheek-teeth with crown at least partly trenchant,
the outer cusps of one or more in each jaw narrow
and with well developed cutting edge, the inner
cusps reduced or absent ; upper carnassial 3-rooted,
behind anteorbital foramen, its inner lobe median
or anterior.
Cheek-teeth without crushing surfaces ; upper molar
scarcely larger than outer incisor ; claws com-
pletely retractile (Cats) Fclidnj, p. i55.
Cheek-teeth, at least the hindermost, with evident
crushing surface ; upper molar (or first when more
than one are present) much larger than outer
incisor ; claws partly or not retractile.
Tooth-row relatively long (more than half condylo-
basal length of skull) ; number of teeth in
European members of family 42 (Dogs) Canidx, p. 303.
* Represented in Australia by a species of Canis, probably introduced.
284
CARNIVOEA
Tooth-row relatively short (less than hal£ condylo-
basal length of skull) ; number of teeth in
European members of family not more than 40.
Auditory l)ulla divided into two chambers, the
boundary between which is marked externally
by an oblique constriction ; upper molars
usually (always in European genera) 2-2,
the crown of the first wider externally than
internally (Genets and Mongoose) Vivcrridn', p. HO.
Auditory bulla simple ; upper molars 1-1, the
crown wider internally than externally Miistclida' , p. 340.
Upper carnassial with evident crushing surface
on inner side, the crown triangular or
rhombic in outline ; upper molar large, the
length of its outer portion usually equal to
or greater than that of carnassial.
Skull normal, the rostrum longer than broad ;
external form not modified for aquatic
life, the toes long-clawed, not webbed,
the tail not conspicuously muscular ; fur
loose and coarse (Badgers) Mdlnas, p. 341.
Skull much flattened ; rostrum broader than
long ; external form modified for aquatic
life, the toes short-clawed, webbed, the
tail conspicuously muscular ; fur dense
and fine (Otters) Lulrinx, p. 354.
Upper carnassial without crushing surface on
inner side other than a small concave area
between small inner lobe and main cusp,
the crown not triangular or rhombic in
outline ; upper molar much reduced, the
length of its outer portion one-third to
one-half that of carnassial.
Dentition highly trenchant; small premolars
not opposite, at least one x^air capable of
shearing action ; upper carnassial with
posterior cusp narrow and trenchant ;
auditory bulla longer than broad; form
slender; feet digitigrade (Martens and
Weasels) Mufteliiuc, p. 3G4.
Dentition not highly trenchant ; small pre-
molars opposite, not capaljle of shearing
action, the points of all but pni^ and pni^
widely separated when jaws are closed ;
upper carnassial with posterior cusp
broad, almost flat-topped; auditory bulla
broader than long ; external form heavy ;
feet sub-plantigrade (Glutton) Guloniitie, p. 432.
Family URSIDiE.
1825. Ursid;i- Gray, Thomson's Annals of Philosophy, xxvi, p. 339,
November, 1825.
(jeot/rapliicnl distribution. — Northern hemisphere, south in the
Old World to the Atlas Mountains and the Malay Archipelago,
and in America to the Andes.
Characters. — Larger cheek-teeth of a strictly crushing ty23e,
the crowns wide and flattened, with large terete cusps, the last
DRsus 285
upper premolar and first lower molar scarcely differentiated as
carnassials, the former 2-rooted, its inner lobe at posterior border
of crown, its position so far anterior to level of anteorbital
foramen as not to he at point of greatest mechanical efficiency ;
auditory bulla flattened, without septum ; form heavy ; size
large ; feet strictly plantigrade ; digits, 5-5.
Memarlcs. — The family Ursidse, containing the bears, is at
jiresent represented by five or six genera, though the fossil
remains of others are known. The members of the group are so
easily recognizable by the peculiarities of the cheek-teeth that
they require no special comparisons with other carnivora. Two
genera occur in Europe.
KEY TO THE EUROPEAN GENERA OP UBSIDM.
Cheek-teeth relatively large ; incisors and canines not
specially enlarged and prehensive (Ordinary Bears) Ursus, ]). 285.
Cheek-teeth relatively small ; incisors and canines
enlarged and unusually prehensive in character
(Polar Bears) Thalarctos, p. 297.
Genus URSUS Linnwus.
1758. Ursus Linnaeus, Syst. Nat., i, 10th ed., p. 47 (arctos, by tau'touymy).
1857. Ursus Blasius, Saugethiere Deutschlands, p. 196.
1864. Euarctos Gray, Proc. Zool. Soc. London, p. 692 [americanns).
1864. Myrmarctos Gray, Proc. Zool. Soc. Loudon, p. 694 (eversmanni = arctos).
1898. Ursarctos Heude, Mdm. Hist. Nat. Emp. Chinois, iv, pt. i, p. 18
(yesoc7isis).
Tijpe species. — Ursus arctos Linnteus.
Geographical distribution. — Northern hemisphere from northern
limits of the great continental areas south to the Atlas Moun-
tains, the Himalayas and Mexico.
Characters. — Dental formula : i ^\ c J"*, pm '^-*, m P^ = 42 ;
inner upper incisor well developed, permanent ; first, second and
third premolars in both jaws small, single-rooted, readily deciduous,
especially jj?(i'-, j»/i._, and jjvHj ; molars large and robust, the length
of the two upjier teeth together eqiial to width of palate.
I{e7nar]cs. — The genus TJrsus as thus restricted is a very
homogeneous gi'oup practically confined to the north temperate
region. The species are at present so impei-fectly knowm that no
fair estimate can be made of their number. Recently about
thirty forms have been recognized, only one of which is definitely
known to occur in Europe.
URSUS ARCTOS Linnreus.
1758. [LVsifs] arctos Linnreus, Syst. Nat., i, 10th ed., p. 47 (Sweden).
1772. [JJrsus'] ursus Boddaert, Kortbegrip van het zamenstel der Natuur,
I, p. 4C (Renaming of arctos).
1788. [Ursus arctos'\ a nigcr Gmelin, Syst. Nat., i, 13th ed., p. 100
(Northern Europe).
286 CAENIVORA
1788. [Ursus ardos'] fi fiiscus Gmelin, Sj-st. Nat., i, 13th ed., p. 100 (Alps).
1778. [UrsHS arctos] y albus Gmelin, Syst. Nat., i, 13th od., p. 100
(Unknown ; based on the " ours blano terrestre " of Buff on).
1792. U'[rs;(s] arctos griscus Kerr, Anim. Kingd., p. 184 (Germany ; also in
northern North America).
1797. Ursjis arctos rufus Borkhauseu, Deutsche Fauna, i, p. 40 (Swiss and
Tirolean Alps).
1798. Ursus baclius Schrauk, Fauna Boica, I, p. 55 (Forests on the
Bohemian boundary).
1808. Ursus fuscus Ticdemann, Zoologie, i, p. 374 (Substitute for arctos).
1814. Ursus alpinus Fischer, Zoognosia, iii, p. 161 (Alps? Based on an
individual seen alive in Paris).
1820. Ursus arctos ma jar Nilsson, Skand. Fauna, i, p. 112 (Wooded portions
of southern Scandinavia).
1820. Ursus arctos minor Nilsson, Skand. Fauna, i, p. 123 (Northernmost
Scandinavia).
1827. [Ursus arctos] & brunncus Billberg, Synopsis FauniB Scandinavia?,
p. 15 (Northern Scandinavia).
1827. [Ursus arctos] y annulatus Billberg, Synopsis Faunre Scandinavia?,
p. 15 (Northern Scandinavia).
1827. [Ursus arctos] 5 argenteus Billberg, Synopsis Faunae Scandinavite,
p. 15 (Northern Scandinavia).
1827. [Ursus] otii/rmcphagus Billberg, Synopsis Faunae Scandinavise, p. IG
(Northern Scandinavia).
1828. [Ursus] formicarius Billberg, Synopsis Faunse Scandinavise, 2nd ed.,
p. 16 (Renaming of myrmephagus).
1829. U[rsus] pyrcnaicns Fischer, Synopsis Mamm., p. 142. Latinization
of "Ours des Pyrenees" F. Cuvier, Hist. Nat. des Mammif., v,
fasc. 44, 1824 (Asturias, Spain).
1829. U[rsus] norvcgicus Fischer, Synopsis Mamm., p. 142. Latinization
of "Ours de Norwege " F. Cuvier, Hist. Nat. des Mammif., ii,
fasc. 7, 1819 (Norway).
1836. ? ZJ;-s?(s/aZci5(C)- Reichenbach, Regn. Anim. Icon., i, p. 82 ("Pyrenees" ;
afterwards supposed to be an individual of " 17. ferox." See Natur-
gesch. des In- und Auslands, Raubsiiugeth., p. 299, 1852).
1842. Ursus pyrenxus F. Cuvier, Hist. Nat. des Mamm., Tabl. gen., p. 3
(Described in fasc. 44, 1824) (Asturias, Spain).
1847. ? Ursus curyrhinus Nilsson, Skand. Fauna, i, 2nd ed., p. 212 (Sweden ?
Type an individual raised in captivity).
1857. Ursus arctos Blasius, Siiugethiere Deutschlands, p. 196.
1801. Ursus arctos aureus Fitzinger, Wissensch.-pop. Naturgesch. dcr
Siiugeth., I, p. 372 (Norway).
1864. [Ursus arctos] var. 1. normalis Gray, Proc. Zool. Soc, London, p. 082
(Renaming of arctos).
1864. [Ursus arctos] sub-var. a. scandinavicus Gray, Proc. Zool. Soc,
London, p. 082 (Based on Nilsson, Ilium. Fig. Skand. Fauna,
pi. 23).
1864. ? [Ursus arctos] sub-var. c. rossicus Gray, Proc. Zool. Soc, London,
p. 682 {nomcn mulum).
1864. [Ursus arctos] sub-var. f. polonicus Gray, Proc. Zool. Soc, London,
p. 682' (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, pi. xxii,
fig. 3).
1864. [Ursus arctos] var. 2. grandis Gray, Proc. Zool. Soc, London, p. 684
("North of Europe " ; based on " a male purchased at Hull, living
iu the Zoological Gardens from 1852 to 1863").
uRsus 287
1804. [Urstts arctos] var. 4. stenorostris Gray, Proc. Zool. Soc, London,
p. G85 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, 2ad var.,
pi. xxii, fig. 4).
1864. Myrmarctos evcrsmanni Gray, Proc. Zool. Soc, London, p. C95
(Norway).
1910. Ursus arctos, U. arctos formicarius, U. arctos alpinus, and JJ. arctos
pyrenaicus Trouessart, Faune Mamm. d'Europe, pp. 67-08.
Type locality. — Sweden.
Geographical distribution. — Entire continent of Europe wher-
ever sufficiently extensive forests remain ; east into Asia ; west
formerly to Great Britain, where it became extinct about the
eleventh century ; not certainly known to have occurred in
Ireland.
Diagnosis. — Size moderate, condylobasal length of skull
ranging from about 260 to 350 mm. ; interorbital region notice-
ably elevated, the frontal profile strongly convex ; mesopterygoid
region not specially shortened and broadened, the width between
pterygoids decidedly less than half distance from hamular to
level of last molar ; colour brown or buffy, varying much in exact
shade, the legs usually darker than body, and feet darker than
legs.
External characters. — General form short and heavy, this
made more apparent by the long rather loose fur. Head
moderately pointed, rather broad posteriorly ; ear short, narrowly
rounded off above, nearly concealed in the fur, its tip not extend-
ing to eye when laid forward ; muzzle squarely truncate, its piad
naked, the upper border somewhat projecting backward, its lower
border separated from upper lip by a broad hairy area crossed
at middle by a nearly bare perpendicular line. Fore foot with
digits robust, inconspicuously graduated, the third and fourth
sub-equal and loiagest, the fifth and second sub-equal and slightly
shorter, the first with anterior edge of ball extending about to
middle of that of second, this interval greater than in the case
of the other digits ; claws strongly curved, blunt, without evident
cutting edges, their length at least twice that of those on hind
foot ; balls of digits large, pad-like, their surface, like that of
pads, coarsely rugose ; main pad wider than long, covering more
than half surface of palm, its outer border about twice as long as
inner, its porterior border slightly concave, its inner portion, at
base of thumb, marked off from rest of pad by a slight furrow ;
region between main pad and balls of digits densely furred ;
wrist pad about as large as ball of digits, near outer margin of
palm, its long diameter transverse ; region between wrist-pad and
main pad densely furred ; hairs along edge of palm standing out
stiffly, especially on outer side. Hind foot longer than fore foot,
the second and third digits sub-equal and longest, the first and
fourth slightly shorter, the fifth with anterior edge of ball at
middle of that of fourth ; pad like that of fore foot, but with a
broad backward extension passing along inner side nearly or
288
CARNIVORA
quite to heel ; region between pad and balls of toes, and at outer
side of backward extension densely furred ; fringe along edge of
foot conspicuous. Tail very short, concealed in the fur.
Colour. — The colour of body is usually a light brown or dull
buff, the head not essentially different, but feet and outer surface
of legs darker. Many individual difierences in colour have been
described, some of which are probably characteristic of geographical
races.
Skull. — Genera] form of skull rather robust, the rostrum
Via. 50.
i^r.^us arctos.
moderately long (distance from orbit to front of premaxillary
contained about 2^ times in condylobasal length), the brain-case
289
deep (depth to level of under side of poistglenoid process con-
siderably more than distance from tip of postorbital process to
middle of intei-parietal) but not unusually wide (mastoid breadtli
slightly exceeding depth to under side of postglenoid process).
Dorsal profile usually with an evident concavity in interorbital
region, but this character showing much variation ; highest point
at bregma or slightly further forward, the profile nearly straight
and sloping away at an angle of about 20° behind tliis j^oint to
slightly overhanging lambdal region ; ventral profile faintly and
Fig. 51.
Uistis arcton. x J.
evenly concave throughout. Brain-case broadly ovate in outline, its
greatest breadth about equal to distance from bregma to lambda,"
its depth at middle, exclusive of sagittal crest, slightly less than
greatest breadth ; lambdoid and sagittal crests well developed in
adults, tlie lambda noticeably projecting so that occiptal condyles
are scarcely visible when skull is viewed from above, and region
between crest and foramen magnum is deeply concave ; sagittal
crest dividing in region of bregma into two ridges, one of which
* Except in very old individuals in which the lambda is greatly
produced backward.
U
-i90 CAlRNn'ORA
runs to extremity of each jaostorbital process, the hinder margin
of which it forms ; occiput when viewed from behind moderately
broad, the depth from lambda to lower lip of foramen magnum
contained If to 1| times in mastoid breadth ; mastoid processes,
paroceipital processes, and condyles extending to about the same
level. Floor of brain case nearly flat, the basioccipital with
raised edges applied to inner surface of bullw, and in some
specimens marked by an evident concavity on each side of median
line ; auditory bulla flat, not rising above edge of basioccipital, the
greatest longitudinal diameter less than transverse diameter, the
URSUS 291
meatus distinctly tubular, usually longer than wide ; postglenoid
process heavy, rising to level of hamulars. Interorbital region
broad, the width across robust, triangular, postoi-bital processes
about equal to that of brain-case, the region immediately between
orbits always a little concave and sometimes conspicuously so,
that at base of each postorbital pi'ocess usually somewhat swollen.
Zygomata moderately expanded, the greatest zygomatic breadth
opposite anterior glenoid edge ; orbital process well developed,
liostrum equal to less than half condyloljasal length of skull, the
width across alveoli of canines equal to or less than depth at
front of orbit, the depth at front of nasal equal to about half
distance from orbit to front of premaxillary ; nares rather large,
their lateral margins slightly everted ; nasal bones elongate wedge-
shaped, squarely truncate anteriorly, their posterior extremity on
level with or extending slightly behind nasal branch of maxillary ;
anteorbital foramen over metacone of m^ or paracone of nr ;
palate nari-ow, its width between posterioi' molars contained about
3^ times in median length ; extension behind molars nearly
parallel-sided, its length equal to about three-quarters bi-eadth ;
mesopterygoid space 1 A- times to twice as long as wide, its anterior
border squarish or I'ounded, its lateral borders nearly parallel ;
hamulars small but distinct, slightly hooked outward. Man-
dible robust, the depth of ramus behind large premolar con-
tained about five times in length, the height of jjosterior portion
(measured to level of lower border) a little less than half length ;
coronoid process broad, its width at level of alveolus slightly
greater than height, the anterior border at first straight then
evenly convex to overhanging tip, the posterior border concave ;
angular process short, extending slightly if at all behind level of
articular process, its inner border nearly straight, its outer border
convex.
Teeth. — The teeth are moderately large relatively to size of
skull. Upper incisors forming a continuous row, separated at
each side from canine by a diastema about as wide as inner
incisor ; i^ and P sub-equal, the former slightly the smaller, the
anterior face smoothly rounded, a little more than half as wide
as high, the posterior face abruptly concave, with shelf-like
posterior extension, the cingulum slightly de^ eloped and forming
a rudimentary nodule on inner and outer side of /- and on outer
side of i^ near level, of middle of anterior surface ; P with crown
area nearly double that of r, and height nearly half that of
canine, its anterior surface smoothly rounded but with pronounced
nodule on inner side, its posterior sui-face gradually concave
and without shelf-like extension ; a thickened ridge along its inner
border and a low but somewhat trenchant ridge slightly outside
of middle. Lower incisors forming continuous row between
canines, their crown area increasing regularly from first to third,
their height approximately equal ; each has a high inner cusp and
a low outer tubercle best developed in ('3 ; posterior border slightly
u 2
292
CARNIVOEA
Fig. &3.
Wrsus arctos. Teetli nat. size.
URsus 293
concave, with inner, middle and outer ridges, the inner and middle
low and confluent in i\. Canines large and strong, oval or
slightly ovate in cross section, the longest diameter at level of
alveolus equal to or slightly greater than distance from alveolus
to median line of palate ; lower canine shorter and a little more
curved than upper ; a slightly de^■eloped posterior and antero-
internal longitudinal ridge, most evident in upper tooth ; no
cingulum. First and third upper premolars small, flat topped,
with slightly indicated cusp and posterior and antero-internal
ridge, the crown area slightly less than that of smallest incisor,
the first close to canine, the third close to large premolar ; third
near middle of space between first and second, much smaller than
the others and frequently deciduous, its crown indefinitely
rounded ; fourth upper premolar with crown area about half that
of first molar, its general outline triangular with apex directed
forward, the outer side longest, the posterior border shortest, the
contrast between them sometimes noticeable, in other instances
slight, the three cusps lying near respective angles, the anterior
highest (reaching level of main cusps of molars) and most robust,
the posterointernal and jiostero-external abruptly smaller, sub-
equal, the inner usually lower than the outer ; a small accessory
tubercle usually present at posterior base of postero-outer cusp ;
cingulum obsolete but usually visible along inner base of anterior
cusp and outer base of postero-external cusp ; first and second
lower premolars approximately like corresponding upper teeth in
both size, form and position ; third very early deciduous, usually
if not always absent in adult individuals ; fourth in contact with
first molar and with from one-third to nearly one-half its crown
area, the outline irregularly quadrilateral with well develoj^ed
antero-external cusp nearly as high as main cusps of molars, a
rudimentary antero-internal cingulum cusp, and a tuberculated
I'idge extending along outer side of crown from antero-external
cusp to posterior border ; occasionally a similar ridge is present
on inner side of crown, its anterior tubercle forming an evident
cusp at inner posterior base of main cusp. First upper molar
with crown much less than twice as long as broad, its outer side
bi-convex, its inner side evenly rounded, its two outer cusps
sub-equal in both height and diameter, the two inner cusps
decidedly lower than outer, and less well defined, owing to the
presence of a low, ridge-like tubercle between them ; both outer
and inner cusps when unworn have distinct though low anterior
and posterior trenchant ridge ; space between outer and inner
cusps occupied by a rather well defined longitudinal groove, the
surface of which is marked by irregular low ridges and furrows \
cingulum obsolete, but indicated in the regions between the
cusps ; second upper niolar nearly twice as long as broad, the
anterior two-thirds approximately like first molar, with the same
four cusps and intermediate longitudinal groove, the main axis
of which is, however, in axis of tooth-row instead of slightly
294
CARNIVORA
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296 CARNIVOEA
oblique to it ; postero-external cusp slightly smaller than antero-
external, and contrast between outer and inner cusps less marked
than in in^ ; posterior third of crown occupied by a flattened heel,
variable in form and size, but usually narrowing off, chiefly by
slanting inward of outer border, to about half anterior width of
tooth, and in some specimens bearing a low but evident third
inner tubercle ; surface of heel sculptured by irregular small
tubercles and furrows ; cingulum obsolete but usually evident
along anterior half of inner border. First lower molar about as
long as second but noticeably narrower, its crown showing more
traces of the jsrimitive trituberculate form than any of the other
teeth ; protoconid and hypoconid wide apart, separated by a deep
groove, the protoconid the highest cusp in the tooth, and with
evident antero-external commissure ; paraconid forming narrow
anterior extremity of crown and provided with a distinct com-
missure, similar to and joining that of protoconid ; metaconid
subterete, without commissure, near to and slightly behind inner
base of protoconid, a minute though evident accessory tubercle
just in front of it ; entoconid like hypoconid, at extreme posterior
edge of crown, the deep wide groove between it and metaconid
with small accessory tubercle at its lowest point. Second lower
molar about 1^ times as long as wide, its outline an irregular
parallelogram with rounded-off corners, the siirface of the crown
occupied principally by a flattened, irregularly sculptured crushing
surface, the cusps near border ; Ave cusps are usually well
developed : a rather large antero-internal and antero-external
opposite each other, and joined by a low transverse ridge ; a
small postero-external and two smaller postero-internal cusps ; a
small but evident accessory tubercle at anterior base of large
antero-internal cusp, this tubercle not infrequently dividing into
two. Third lower molar varying from rounded-triangular to
ovate in outline, its area about two-thirds that of riin, its surface
entirely flat excejit for a slightly raised rim which forms a
small antero-internal cusp.
Measurements. — Adult male from Sweden (mounted) : head
and body, 1900 ; tail, SO; hind foot, 195 ; ear, 90. For cranial
measurements see Table, p. 294.
Specimens examined. — Eight, from the following localities : —
Sweden : Kvickjock, Norbotten, 1 ; bo exact locality, 3 (B.M. and
U.S.N.M.).
Switzerland : Engadine, 1.
Adsteia-Hungaey : Near Hatszeg, Hunyad, 3.
Bcmarhs. — As regards the existence of geographical races of
the large European bear it is impossible to form any opinion on
the basis of the few sjiecimens seen. Ur.'ius nrctos is related to
the grizzly bear of North America, U. horribilis and its local
forms, but is readily distinguishable by the relatively greater
height of the frontal region and the consequently more abrupt
slope of posterior half of dorsal profile, a character suggesting
TIIALARCTOS
297
Umus richarfhnni ; forehead in most specimens rising tabriiptly
al)Ove level of rostrum so as to produce a noticeable concavity in
dorsal profile, but this character subject to marked variations,
the exact nature of which is not fully understood. The skulls
of the two species are of approximately the same size, those of
U. horrihilis perhaps averaging somewhat the larger. The teeth
of the two animals are also much alike.
9. Kvickjock, Norbotten, Stockholm Museum 90. 8. 1. 3.
Sweden. (e).
2 S. Sweden. (Lloyd.) Purchased (Stevens). 02. 3. 29. 7-8.
9 St. Engadine, Switzerland. H. Justen (p). 86. 1. 23. 1.
3 skeletons. Hatszeg, Transylvania, C. G. Danford (c). 78. 1. 16. 1-3.
Hungary.
Note on the Ursus formicarius of Bieler.
A peculiar small bear supposed to have been taken in the
Canton of Grisons, Switzerland, has been described by Professor
Bieler of the Lausanne Agricultural College as Ursus formicarius
Eversmann.* Through the author's kindness I have had the
opportunity to examine this skull. It is that of a rather young
individual, apparently a female, with basal suture open, but with
teeth showing slight indications of wear. In size it is smaller
than in a female of the same age or slightly younger from
Sweden (see Table of cranial measurements, p. 294), the inter-
orbital region is much depressed, so that the frontal profile is
nearly flat, and the mesopterygoid fossa is unusually broad and
short, the width between pterygoids equal to a little more than
half distance from hamular to level of last molar. The teeth on
the other hand are slightly larger than usual in females (see
Table, p. 295) ; inform they show no special peculiarities. Small
bears, presumably of this type, have been reported from Spain,
northern Italy, Russia and Scandinavia, and have formed
the basis of such names as Ursus formicarius, U. arctos minor
and Myrmarctos eversmanni. Until more is known of them, as
well as of the normal variation in ordinacy Ursus arctos^ their
status must remain in doubt.
Genus THALARCTOS Gray.
1825. Tlialarctos Gray, Ann. of Philosophy, N.S., x, p. 62, July, 1825.
1825. Thalassarctos Gray, Ann. of Philosophy, N.S., x, p. 339, November,
1825.
1896. TJialassiarcJius Kobclt, Boricht Senckenberg. naturforsch. Gesellsch.
Frankfurt am JIain, p. 93 (Substitute for Tlialarctos).
Type species. — Tlialarctos polaris Gray = Ursus viariiimns
Phipps.
* Compto-Rendu des Stances du Sixi^me Congrus Internationale do
Zoologie, Berne, 1904, p. 248. 1905.
298
CARNIVORA
G(jo(jraiiltical distribution. — North Polar region, south to
northernmost continental coasts.
Characters. — Similar to Ursus, but cheek-teeth much less
robust, the combined length of the two upjaer molars not equal
to width of palate, and canines and incisors enlarged and more
prehensive in general form.
Remarks. — The genus Thalarcios, though not very strongly
dift'erentiated from Ursus, is a well defined and perfectly natural
group.
THALARCIOS MARITIMUS Phipps.
1774. Ursics maritimns Phipps, Voyage toward North Polo, p. 185 (Spitz-
bcrgen).
1776. Ursus mariiius Pallas, Eeise durch verschiedeue Provinzen des
russischen Beichs, iii, p. 691 (Arctic Ocean).
1792. Ursus folaris Shaw, ^Museum Levorianum, i, p. 7 (P^enaming of
marinus).
1862. Thalarctos maritimus Gray, Catal. Bones Mamm. Brit. JIus., p. 105.
1908. ? Thalassarctos jenacnsis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Jena Island,
Spitzbergen).
1908. ? Thalassarctos spitzbergensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Seven Island,
Spitzbergen).
1910. Ursus [Thalassarctos) maritimus Trouessart, Faune Mamm. d'Europe,
p. 66.
Tijpe locality. — Spitzbergen.
Geographical distrihution. — Arctic Ocean, south on floating ice
occasionally to the northern coast of Norway. Details of
distribution unknown.
Diagnosis. — General characters as in the genus ; size very
lai'ge ; colour uniform whitish or buffy.
External characters. — Form lijnger and less heavy than in
Ursus arcios, the neck noticeably longer and head longer and
more pointed ; ear actually as well as relatively shorter ; fore
foot with palmar tubercles and balls of toes essentially as in
U. arctos but smaller ; pad on hind foot without backward
continuation along inner portion of sole ; claws much less
elongated than in U. arctos, not strongly curved, but with acute
jjoints and well developed cutting edges. Fur very dense, its
texture almost seal-like in the short summer coat.
Colour. — Entire animal a uniform whitish or buffy, the winter
pelage tending to be a creamy-white, the summer coat yellowish
buff.
Skull. — The skull is considerably larger than that of Ursus
arcios, with relatively longer brain-case, deeper, wider rostrum,
and less elevated frontal region ; lambdal region less produced
backward than in Ursus arctos, the condyles usually visible when
skull is viewed from above. Base of brain-case essentially as in
U. arctos, but portion at base of condyles more narrowed and
THALARC'TOS
299
elongate. Palate iioticea1)ly broadei' than in Uisus arctos, a
character made more conspicuous by the relative weakness of the
teeth. Mandible with no sj)ecial [leculiarities except that the
FIO. 54.
Thalnrclos maritimiis.
lower margin is nearly straight thi'oughout, the posterior concavity
being very slightl}' indicated, and lower border of angular process
only a little elevated above general outline.
Teeth. — While the general character of the dentition differs
300
CAKNIVOEA
notably from that of Ursus arctos in the reduction of the molars
and increased size and prehensiveness of the canines and incisors,
the details of the individual teeth present little that is specially
FIO. 55.
Thalarctos maritimus. x ;.
noteworthy. General form of upper incisors as in Ursus arctos,
but points of i^ and i'^ narrower and more hooked backward ; P
with cusp more slender and ridges nearly obsolete ; lower incisors
THALARCTOS
301
with lobes more sharply defined. Canines both above and below
essentially similar to those of Ursas arcfos, except for their greater
size. Owing to the greater width of palate the proportion of
FlO. 5C.
Thalarctos maritimus.
diameter of upper canine to palatal width is about as in the
smaller-toothed animal. Small premolars showing no special
peculiarities. Large upper premolar with relati\ely higher
302
CAP.NIVORA
Thalardos marUimux. Teetli iiat. size.
THALAKCTOS
303
anterioi' cusp than in Z7. nrctos, its inner side more flattened,
giving tlie tooth a more carnassial appearance ; postero-internal
cusp relatively less developed. Large lower premolar essentially
as in U. arctos but with somewhat more slender cusp. Molars
differing from those of Ursus arctos in their smoother, less sculp-
tured crushing surface, and slightly more trenchant cusps. Form
of viy not peculiar, though outer cusps are higher and narrower
and inner cusps relatively lower ; «!- with inner cusjjs obsolete
and heel relatively narrower and less developed. Anterior lower
molar with metaconid and its accessory tubercle reduced to a low
irregularly tuberculate ridge ; commissure of protoconid and
paraconid obsolete ; hypoconid and entoconid smaller and much
nearer together than in Urms arctos, though separated from
anterior cusps by a normally wide interval, in which, however,
there are no definitely formed accessory tubercles. Second lower
molar with the same elements as in U. arctos except for the
absence of all trace of an intermediary tubercle on inner side of
crown. Third lower molar with crown nearly flat, its margin
showing only the faintest trace of antero-internal and antero-
external elevations.
Measurements. — Adult male from Behi'ing Strait (mounted) :
head and body, 2670 ; tail, 90 ; hind foot, 370 ; ear, 80. For
cranial measurements see Table, p. 294.
Specijiiens examined. — Nine, from the following localities: — Spitzbergen,
1 (U.S.N.M.); Griffin Bay, Wellington Channel, 1; Melville Island, 1;
Arctic Ocean, 1 ; no history, 5.
[The Museum specimens appear all to have come from the
American side of the Atlantic]
Family CANID^.
1821. Canidx Gray, London Medical Repository, xv, p. 301, April 1, 1S21.
Oeographico I distrihution. — Essentially cosmopolitan ; in E urope
west to Ireland.
Characters. — Larger cheek-teeth of a combined trenchant and
crushing type, the last upper premolar and first lower molar
strongly differentiated as carnassials, the former 3-rooted, its
inner lobe in front of middle of crown, its position, somewhat
posterior to level of anteorbital foramen, at point of greatest
mechanical efficiency ; auditory bulla moderate!)' or considerably
inflated, without septum ; form rather light, the legs long ; size
moderate ; feet digitigrade ; toes, 5-4 or 4-4.
licinarlcs. — Notwithstanding its wide disti'ibution the family
Caniiiue is not rich in genera. About a dozen are now recognized,
three of which occur in Europe.
304
CAENIVOEA
KEY TO THE EUROPEAN GENERA OF CANID.E.
Interorbital region distinctly elevated ; postorbital processes
convex above ; pupil of eye round Ca«is, p. 304.
Interorbital region not elevated ; postorbital processes not
convex above ; pupil of eye elliptical.
Postorbital processes flat or very slightly concave above ;
forehead rising abruptly above level of rostrum ; ear
rounded Alopex, p. 318.
Postorbital processes distinctly concave above ; forehead
rising gradually above level of rostrum ; ear pointed VuIjm's, p. 325.
Genus CANIS Linnasus.
1758. Ganis Linnaeus, Syst. Nat., i, 10th ed., p. 38 (type by tautouymy
C. familiaris).
1810. Lupus Oken, Lenrb. d. Naturgesch., iii, pt. 2, p. 1039 [Canis lupus,
by tautouymy).
1837. Vulp)icanis Blainville, Ann. Sci. Nat., Paris, 2nd ser., Zool., viii,
p. 279, November, 1837 (Canis auretts Linngeus).
1839. Lyciscus H. Smith, Jardine'a Naturalists' Library, Mammals, ix,
p. 160 {Canis latrans Say).
1839. Thous H. Smith, Jardine's Naturalists' Library, Mammals, ix,
p. 193 (Canis antlms F. Cuvier).
1839. Sacalius H. Smith, Jardine's Naturalists' Library, Mammals, ix,
X3. 213 (Canis aureus Linnaeus).
1841. Oxygous Hodgson, Calcutta Journ. Nat. Hist., ii, p. 213 (Canis
aureus Linnaeus).
1857. Canis Blasius, Saugethiere Deutschlands, p. 177.
1868. Neocyon Gray, Proc. Zool. Soc, London, p. 506 (Canis latrans Say).
18G9. Dieba Gray, Catal. Cam. Pachyd. and Edentate Mamm. Brit. Mus.,
p. 180 (Canis anthus F. Cuvier).
Type species. — Canis famiUaris Liniifeus.
Geographical distribution. — Nearly as in the family, but
absent from the Malay Archipelago and South America ; in
Europe west within historic times to Great Britain, but now
restricted to the continent.
Characters. — Skull heavy and deep (depth of brain-case more
than one-third condylobasal length) ; interoi-bital region thickened
and elevated, the frontal sinuses i-ather large, the postorbital
processes thick, convex above, their edges rounded off ; dorsal
profile of forehead rising rather abruptly and noticeably above
level of rostrum ; dental formula : i ||, c j^J, ^jwt t^, in |^ = 42 ;
teeth heavy and large, the length of carnassial and upper molars
together contained about 21 times in palatal length ; canines
robust and not specially elongated, the point of ujjper tooth
extending scarcely beyond middle of mandibular ramus when
jaws are closed (fig. 65).
Bemarks. — Much uncertainty exists at present with regard
to the limits of the genus Canis. As here defined the group
includes the domestic dogs, the true wolves, the American prairie
CANIS
305
wolves, .-u.d the Old World jackals. Two species are known
%o:::;l^ :;::::''' - ^^ -^^-^^ ^« -p— ^^d ,y seve.ai
KEY TO THE EUROPEAN FORMS OP CANIS.
Condylobasal length of skull less than 200 mm • teeth
not so large as in the largest domestic dogs ('len"th
of upper carnassial 17 to 18 mm.); cingulum on
outer margm of m' broad and conspicuous (South-
eastern Europe ; Jackal) ^ r n„.
Condylobasal length of skull more " than Iw' mm"- ^- «"'^'««' P' 315.
teeth larger than in the largest domestic dogs'
(length of upper carnassial 25 to 27 mm.) • cingu-
lum on outer border of m' narrow, tending to bo
mcompleto at middle (Distribution general • true
Wolves) r 1
Size rather small "(exact' "dViension's'^unknowni '^' '^"^'
(Southern Spam n , , ■,
Size large, C. l. dettanus, ]). 615.
White of throat not extending uninterruptedly
WI,ih?nfn.^^?*''v^'''^"°'"'^°™ Europe) ... C.Llu,,„.,r, 313
White of throat extending uninterruptedly on to
cheek (Spain except extreme south) c. I. signatus, p. 314.
CANIS LUPUS Linnc-eus.
(Synonymy under subspecies.)
G«.i,r«y./caZ d/.<n-J«fion.-Originally throughout Europe from
SrfttwT r'in°'^,^"Tx.^^'''^' °«^^' exterminated in the
British Islands, Holland and Denmark
/09,f/"T'''-""^°°'ly^^^''''^ l^'^gth of skull more than 200 mm
(220 to 250 mm.) ; cheek-teeth larger than in the largest racS
o domestic dogs the upper carnassial 25 to 27 mm. In length
but structure not peculiar, the upper molars with narrow, incon!
spicuous cingulum on outer side (licr 61)
d^„fH'7,^;!/-^";™tr-^^e^«ral Form essentially as in doitestic
Sed if '? X'V ^^'' ^«derately long, erect, somewhat
Z h ' ^■^^"'^"^g '^^^""V" 'y' ''^'''^ ^^''^ ^«^^^^"'^^- Muzzle pad
completely bare. Fore foot with third and fourth digits s^ib-
e.]ual and longest, second and fifth shorter, the large Vd-like
tdtlvtHfol rl^^f^''" ?^"^^ "^ *^"'-^^ -"d fourth and the
fkof If ' hfi't-shaped main pad, the combined area of
u o digits greater than that of pad; thumb much shorter
than other digits, the nail smaller, but not peculiar in form its
extremity not reaching level of posterior boiler of ma n pa i
1 scarcely indicated, no pad at its base; wrist pad single, near
Hini w' ^y^'^^fTf^^h^t more than half that of bait of toes.
•Uion/ rf''"*'''"7 ^'^ ^? ^"^^' ^^"t ^^'^""-^ '-^"d posterior pad
Padsnndfir' '"^"f' 'l^'^^l^^ '''''''''^' .s"b-e.,ual throughout,
lads and balls narrowly edged with short hair
of legs yellowish brown or buff, darker along median region of
X
306
CARNIVORA
back, on posterior portion of head and outer surface of ears,
lighter and more incHned toward greyish at sides of shoulders
and between ears and eyes ; longer hairs of back and sides black-
tipped, producing an evident dark shading over middle of back.
I'IG. 5S.
Canis lujms. x i.
especially behind middle and at base of tail ; pencil narrowly
clear black, rest of tail essentially like back. Underparts and
inner surface of legs pale buft" or buffy white, not strongly
contrasted with sides, the chin and inturraiuia usually grizzled.
CANIS 307
frequently margined with blackish ; upper lip to muzzle pad and
including lower half of cheek dull whitish, usually not very
different from throat ; inner surface of ear light buff.
Skull . — In general aspect the skull diHers slightly if at all
I'lG. 59.
Canis liquis. x *.
from that of some of the larger races of domestic dogs, though
often attaining a greater size. The rostrum, however, appears
to bo relatively less rtibust than in dog skulls of approximately
the same length. Dorsal profile rising gradually from nares to
308
CAIINIVORA
just in front of orbit, then abruptly to a little in front of ])regma,
behind which it is nearly flat to strongly overhanging lambdal
region. Depth of brain-case through auditory bulla about 2h
Fig. CO.
Canis lupus.
times that of rostrum behind canine, and about equal to mastoid
breadth. Brain-case rather elongate ovate in outline when
viewed from above, its breadth above roots of zygomata about
l.", times that of rostrum over canines and apj^roximately equal
CANIS 309
to distance from bregma to most posterior point of occiput.
Posterior portion of occiput strongly concave when viewed from
the side, tlio condyles nearly hidden beneath the projecting
lambdal region. Floor of brain-case with no specially noteworthy
features, the auditory bullte sub-circular in outline, with short but
evident meatal tube, their surface evenly inflated (more so than
usual in domestic dogs) except for an evident flattening on antero-
external aspect. Sagittal and lambdoid crests well developed,
the former dividing just in front of bregma into two ridges
curving outward to form posterior border of postorbital processes.
Interorbital region modei'ately elevated, well defined, with distinct
longitudinal concavity between raised and thickened postorbital
processes. Zygomata widely spreading, the greatest zygomatic
breadth (at level of anterior glenoid edge) a little more than half
greatest length of skull ; orbital process well developed, the
orbit surrounded by bone through about four-fifths of its
circumference. Rostral breadth at canine about equal to depth
at front of carnassial ; premaxillary extending posteriorly to
about middle of nasal ; mnxillary extending back nearly to middle
of orbit, slightly exceeded by nasal ; anteorbital foramen about
9x5 mm. in diameter, over posterior root of third premolar.
Palate moderately wide, nearly flat, not extending posteriorly
beyond level of last molar, terminating in an obscure median
spine ; incisive foramina between canines, 11 to 17 mm. in length,
their combined breadth usually a little less ; mesopterygoid fossa
rather more than one-third as long as palate, considerably
narrower posterioi'ly than anteriorly. Mandible strong, but not
remarkably robust, the depth at posterior edge of carnassial
contained about six times in length ; symphysis rather long ;
coronoid process high, the depth of mandible through its middle
noticeably greater than distance from last molar to back of
condyle ; angular process heavy, nearly horizontal, distinctly
raised above level of under margin of ramus.
Teeth. — The teeth are relatively larger than in any of the
races of domestic dogs, though in form they show no tangible
features by which they may be distinguished. Upper incisors form-
ing a continuous, slightly convex row, the outer tooth separated
from canine by a distinct space ; size, when viewed from in front,
increasing regularly from first to third, but third abruptly much
larger than the others in cross-section and nearly half the size of
canine ; anterior surface of i^ and /- slightly more than half as
wide as high, smoothl}' rounded ofl", the cutting edge narrow but
not acute ; a small but distinct secondary lobe at each side of
fi'ont aspect, that of inner side a little below middle, that of
outer side about ecjually above ; posterior sui'face of crown con-
ca\e longitudinally though without backward-projecting basal
shelf ; a well developed median longitudinal rib, and a low but
noticeable cingulum, the latter terminating abruptly and forming
the lobes seen in front view ; outer incisor with no secondary
310
CARNIVORA
lobes, its general form intermediate between that of canine and
of inner incisors. Lower incisors forming continuous row between
canines, the three teeth essentially alike in form, but increasing
no. Gi.
Canis lupua. Tectli nat. size.
regularly in size from iirst to third though less conspicuously
than in the case of the upper incisors ; crowns (viewed from in
front) about twice as high as wide, distinctly bilobed, the outer
CANIS 311
lobe scarcely half as wide as inner ; on inner tooth the outer lobe
is nearly level with cutting edge, on second it lies slightly above
middle of crown, and on third slightly below middle of crown ;
posterior surface oblique, slightly concave, with noticeable longi-
tudinal furrow extending back from notch between lobes. Canines
large, usually 15 mm. or more in diameter at alveolus and about
twice as long, a size rarely if ever attained in domestic dogs,
their surface smooth except for a low antero-internal and
posterior-median logitudinal ridge, the upper teeth slightly longer
and less recurved than the lower. Premolars Jiioderately spaced
except that j-im-* is nearly or quite in contact with the
carnassial ; first, second and third teeth essentially alike in the
two jaws, those of the mandible, however, slightly the less robust ;
first premolar both above and below single-rooted, the crown
simple, that of pm^ subterete, that of j»wi nearly twice as long as
broad, the height in both slightly less than length, the crown area
approximately the same as tbat of corresponding inner incisor, the
small cusp a little in front of middle and with slightly developed
anterior and posterior ridge. Second and third premolar's two-
rooted, the crown about twice as long as wide, sub-elliptical in
outline, the inner margin sometimes (especially in 2J'»h ^'^^ jwu'-)
slightly concave, the long axis nearly parallel with sagittal plane
except in pm^, which is obliquely set ; main cusp a little in front
of middle of crown, its height distinctly more than half length of
crown, its anterior and posterior cutting ridge well developed, the
posterior bearing a distinct secondary cusp situated over middle
of posterior root and relatively larger in lower than in upper
teeth ; a slight shelf -like projection behind secondary cusp ; pm^
similar to jjJHg but considerably larger, its secondary cusp
better develoi^ed and succeeded by a small but evident postero-
basal cusp sprin.ging from the posterior edge of crown ; cingulum
of all the smaller premolars complete though low and incon-
spicuous. Upper carnassial large and robust, the length of crown
along middle slightly more than twice greatest breadth exclusive
of antero-internal lobe, the main axis of the tooth extending
evidently through middle of crown, so that the small, cuspless
inner lobe stands as an offset, slightly breaking the symmetry of
the outline ; main cusp slightly behind middle of crown, its
height more than half length of tooth, its axis slanting distinctly
backward, its anterior and outer surfaces evenly convex except
for the rudimentary longitudinal ridge on basal two-thirds of
fi-ont, its inner surface, together with that of posterior cusp,
flattened ; posterior cusp low and robust, obscured by its very
high nearly horizontal commissure which meets the somewhat
shorter but eciually trenchant conuiiissure of main cusp at an
angle of about 75" ; cingulum complete, though low and incon-
spicuous. Lower carnassial narrower than upper but with eiiually
high crown, the most elevated portion in front of middle instead
of behind it ; protoconid large and robust, resembling main cusp
312
CARNIVORA
of upper carnassial, but with well developed cutting edge both
in front and behind ; paraconid near middle of anterior portion
of crown ; its general form like posterior cusp of upper carnassial,
its connnissure bearing essentially the same relation to that of
protoconid as in the case of the two cusps of the upper tooth,
except that tlie relative lengths of the cutting edges is reversed ;
metaconid small but evident, at postero-internal base of proto-
conid ; liypoconid and entoconid low, occupying the posterior edge
of a well developed though relatively small heel (area of heel
scarcely more than one-third that of anterior portion of tooth)
separated from the cusjjs of the main triangle by a wide transverse
groove ; crown area of entoconid equal to about half that of
hypoconid, its cusp apjaroximately the same size as that of meta-
conid. Second lower molar essentially like heel of carnassial
but larger, its two anterior cusps corresponding in size and form
with liypoconid and entoconid of the large tooth, the posterior
edge of its crown with a small outer cusp resembling the antero-
inner, and sometimes with a slightly developed inner ridge or
rudimentary fourth cusp. Third lower molar single-rooted, the
crown subterete, about as large as that of first premolar, with low
central cusp and rudimentary longitudinal ridge. First upper
molar large, with high outer two-cusped sectorial portion and low
inner crushing portion, the two areas sliarjjly differentiated, the
antero-posterior diameter of the outer decidedly greater than that
of inner ; 23aracone and metacone conical, terete, with slightly
developed anterior and 2:)osterior cutting ridges, the area and
height of metacone about two-thirds those of paracone, the width
of base of which is at least equal to width of inner portion of
tooth ; protocone very low, with low but distinct anterior and
posterior commissures, each of which joins cingulum at base of
corresponding large outer cusps, and each of which bears an
intermediate cusp soon disappearing with wear, the posterior
intermediate cusp larger and more definite in form than anterior
cusp ; hypocone ridge-like, at postero-inner border of crown,
separated from protocone and its posterior commissure by a deep
groove. Second upper molar with about half the crown area of
first, its elements essentially the same, though so reduced that
tlie paracone is scarcely larger than protocime of large tooth,
intermediate cusps on commissures of protocone are barely
indicated, and hypocone is not distinguishable as a cusp distinct
from the cingulum. In both molars the cingulum on outer border
is narrow and inconspicuous relatively to the broad cusps ; in
region between paracone and metacone of m^ it is usually
obsolete (compare figs. 61 and 62).
Remarks. — The material availal^Ie for study has been so poor
tliat I have found it impossible to come to any conclusion with
regard to the existence of local forms of the European Wolf.
The following races have been distinguished by INlr. Caljrera.
There seems to be no good reason to doubt their validity.
CANIS 313
Tlu! only known characters by which the skull of Canifi lupus
can be distiuguishcd from that of the larger domestic dogs is the
greater average general size and the relatively larger teeth. In
a dog's skull with condylobasal length of 2,30 mm. the length of
upper and lower carnassials is respectively 21 "6 and 25 "0 mm.
In ten skulls with condylobasal length of more than 200 mm.
the average and extremes for these teeth are : upper, 20 • 5
(19-22) ; lower, 24 • (22 • 8-26 • 0).* In all the dog skulls which
I have examined, representing such different breeds as the pug,
fox-terrier, bloodhound, mastiff, ancient Egyptian, ancient
Peruvian, Eskimo (Greenland and Alaska) and American Indian,
the teeth are strictly of the wolf type, never showing any
approach to that of the jackal (fig. 62).
Canis lupus lupus Linnajus.
1758. [Cams'] lupus Linniseus, Syst. Nat., i, 10th ecL, p. 39 (Sweden).
1792. C[anis] lupus flavus Kerr, Anim. Kingd., p. 137 (France and
Germany).
1804. Canis lupus niger Hermann, Observ. Zool., p. 32. Not of Kerr,
1792 (Forest of Hagenau, Alsace, Germany).
1839. ? [Canis hqms'] var. canus do Selys-Longchamps, Etudes de Micro-
mamm., p. 144 (nomen nudum).
1839. ? [Canis lupus'] var. fulvus de S61ys-Longchamps, Etudes de Micro-
mamm., p. 144 (nomen nudum).
1841. Lupus oricntalis Wagner, Schreber's Saugthiere, Suppl., ii, p. 367
(Europe).
1857. Canis lupus Blasius, Saugethiere Deutschlands, p. 180.
1863. [Canis lupus] var. major Og6rien, Hist. Nat. du Jura, iii, p. 59
(Lower slopes of the Jura).
1863. [Canis lupus] var. viinor Og^rien, Hist. Nat. du Jura, in, p. 69
(Higher portions of the Jura).
1897. Canis lujnis minor Mojsisovics von Mojsvar, Thierlebeu der osterr.-
hung. Tiefebonen, p. 241 (Southern Hungary). Based on the
" Rohrwolf ," an animal supposed to be smaller and greyer than
true lupus.
1910. Canis lupus and C. lupus hjcaonf Trouessart, Faune ilamm.
d'Europe, p. 90.
Ti/pc locdlitij. — Sweden.
GccHjrupliiral distribution. — Northern and central Eui'upe,
exact limits of range unknown ; formex'ly west to Ireland.
Characters. — Size maximum for the species ; general colour
* Wingo (Danmarks Fauna, Pattedyr, p. 123, 1908) states that in the
skull of a dog from a prehistoric grave (Iron Age) in Denmark, the length
is 209, and that of the two carnassials 20 and 22-5 respectively, while in a
rather large modern "great Dane " the corresponding measurements arc
255, 22 and 28. This author (p. 124) regards the domestic dogs as derived
from Canis aureus.
t -Vpplicd to the wolf of the Pyrenees ; but Schreber's plate lxxxix,
the basis of the name, is a copy of BufEon's plate xli, representing an
animal brought alive to Paris from Canada.
314
CARNIVORA
not markedly tawny ; white of throat not extending to cheeks.
The few skulls examined agree with Asiatic specimens in having
the outer cusps of m^ moderately large, the paracone with trans-
verse diameter of base about equal to width of large flattened
jiortion of crown.
Measurements. — For cranial measurements see Table, j). 316.
Specimens examined. — Pour skulls, from the following localities : —
Sweden: No exact locality, 2 (U.S.N.M.).
Russia: No exact locality, 1 (U.S.N.M.).
Italy : Near SasscUo, Liguria, 1 (Genoa).
Canis lupus signatus Cabrera.
1907. Canis lupus signatus Cabrera, Bol. Real Soc. Espaii. Hist. Nat.,
Madrid, vii, p. 195.
1910. Canis htpus signatus Trouessart, Fauue Mamm. d'Europe, p. 91.
Ti/2)e localitjj. — Escorial, Madrid, Spain.
Geogrnjjhical distrihution. — Central Spain.
Characters. — Size and general appearance as in Canis Inpns
lupus ; colour a more tawny brown than in the northern animal,
particularly on muzzle ; white of throat extending unintei'iuptedly
to cheeks.*
Measurements. — Type (adult male), from Cabrera : head and
body, 1230 ; tail, 400 ; hind foot, 265 ; ear, 125. Young adult
male and female from Province of Burgos, Spain : head and
body, 1130 and 1180 ; tail, 350 and 380 ; hind foot, 225 and 220 ;
ear 120 and 115. For cranial measurements see Table, p. 316.
Specimens examined. — One from Seville, Spain, and two from Province
of Burgos, Spain.
liemarhs. — Tn dentition the Seville specimen differs from all
the other Old World wolves with which I have compared it in
the unusual development of the outer cusps of the upper molars.
The transverse diameter of paracone in m} conspicuously exceeds
width of the small inner 2:)ortion of tooth. Mr. Cabrera informs
me that the tyj^e shows much the same peculiarities. This
character is also present, though less pronounced, in the two
skulls from Burgos, which further differ from northern specimens
in the smaller size and more glol)ular form of the auditory
bulL-e.
i. Seville, Spain. (.1. Rjiiz.) Lord Lilford (p). 95. .3. 3. G.
6,9. Riocabado, Bm-gos. Hon. N. C. Roths- 11. 10. 5. 1-2.
(Rev. S. Gonzalez.) child (p).
* In the Burgos specimens the colour is not unusually tawny : back
and sides a coarse inixture of black, whitish, ochraceous-buff, and drab grey
(underfur), the black and whitish most conspicuous along back, the
ochraceous-buff ou legs and feet (clear and unmixed on latter) ; ear
ochraceous-rufous on outer side (darker and duller at tip), pallid
ochraceous-buff on inner surface ; throat and lower half of cheeks the same
pallid ochraceous-buff ; chin and interramia blackish.
CANIS . 315
Canis lupus deitanus Cabrera.
1907. Canis lupus deitanus Cabrera, Bol. Real Soc. Espafi. Hist Nat
Madrid, vii, p. 107. ''
1910. Canis lupus deitanus Troues.sart, Fauue Mamm. d'Europe, p. 91.
%>e Zorr</%.— Moratalla, Murcia, Spain.
Genf/mphical (h'.strihnt!on.~-l<^o\v known from the type locality
only. -^ •'
D!a<jnosis.~^miiUev than Canis hipm lupus and brighter in
colour, the general ai)pearance much as in C. aureus.
Measurements. — Unknown.
Specimens examined.~l have seen the two living examples in :\radrid
on which the form was based.
Iteniarh'i.^Tn general appearance the two Moratalla wolve.s
are strikingly different from Cam's lupm. Unfortunately their
ti-ue characters are not yet known.
CANIS AUREUS Linnteus.
1758. [Cams'] atireus Liouaeus, Syst. Nat., i, 10th ed., p. 40.
1835. Canis aureus var. moreotica I. Geoffroy, Exp6d.'sci. de Mov&e Zool
pi. I (Morea, Greece). ''
1841. C[anis] dalmatinus Wagner, Schreber's Saugthiere, Suppl ii n 383
(Dalmatia). '-'■ ' ' ^"
1841. C[anis-\ gr^'cus Wagner, Schreber's Saugthiere, Suppl ir p 383
(Peloponesus). ' '■'
1892. Canis aureus halcanicus Brusina, Glasnik Hrvatskoga Naravoslovno^a
Drustva, Zagreb, vii, p. 317 (Drava River, Croatia). "
Type locaUhj. — Province of Lar, Persia.
(ic(>[iraphical distribution.—Indin and westward through Asia
Min.)r to the Balkan Peninsula, north to Heves Comitat,
Hungary.
D/ag-no-s/s.— Smaller than Canis lupus (condylobasal len<^th of
skull less than 200 mm.) ; teeth not equal to those of the larger
domestic tlogs m size, the upper molars with wide, conspicuous
Icmgulum on outer .side (fig. 62).
_ CoZowr.— Upper parts buffy cinnamon, clouded by black hair
tips along dorsal region, nearly clear on sides, and becoming clear
bright cinnamon on outer surface of legs and ear and on area
around and behind base of ear ; muzzle more heavily washed
with black ; from middle of back to base of tail the cinnamon is
replaced by whitish, causing a rather noticeable contrast between
this region and the surrounding parts when fur is disarranged ■
tail essentially like l^ack, the basal half above greyish,°the
terminal half and underside bully much overlaid with black •
pencil blackish ; underparts and niner side of legs dull buffy
316
CARNIVOEA
o
1
Basal suture closed.
„ closed.
„ open.
„ closed.
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8. 10. 2. 50
6. 5. 1. 31
6. 5. 1. 33
6. 5. 1. 34
6. 5. 1. 35
3
C. lupus lupus.
Sweden .
Northern Sweden .
Russia .
Italy: near Sassello,|
Liguria . j
C. lupus signatus.
Spain : near Seville
Prov. Burgos
C. aureus.
Greece : Pirreus
Asia ]Minor ; Khotz
317
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318
CARNIVORA
Skull. — The skull is much smaller than that of Cam's lupus.
In form it differs slightly in the less elevated frontal region and
somewhat more inflated auditory bulla?.
Teeth. — While agreeing with those of Cain's lupus in general
form and in the position of the cusps, the teeth
are on the whole more trenchant in character, as
shown by the general tendency toward narrowness
of crown and prominence of ridges. This is
particularly noticeable in the upper molars, in
which the large cusps are relatively higher, more
slender, and less terete than in Canis lupus, their
cutting ridges much more developed ; transverse
diameter of metacone in m^ noticeably less than
width of inner portion of crown ; cingulum on
outer border of both upper molars wide and
cons2:)icuous in contrast with narrow cusps,
KiG. 62. showing no tendency to become obsolete in
Canis aureus, regioia between paracone and metacone. Lower
^''cheek-teeth"^ carnassial with metaconid actually as well as
Nat. size. relatively larger than in Canis lupus, and posterior
heel with area equal to nearly half that of anterior
portion of tooth, its cus2:)s strongly developed.
Measurements. — For cranial measurements see Table, p. 316.
Siiccimens examined^ — -One from Greece (Piraeus) ; numerous others
from Asia Minor and India.
Bcmarlcs. — The single specimen from Greece agrees sufficiently
with a series of five from Khotz, near Trebizond, Asia Minor,
to make it appear unwise, in the absence of more satisfactory
material, to use one of the Balkan names.
1. Pirfcus, Greece. (C. Mottaz.) Hon. N. C. Roths- 8. 10. 2. 49-50.
child (p).
Genus ALOPEX Kaup.
1829. Alopcx Kaup, Eutw.-Gesch. und Natiirl. Syst. Europ. Thicrw., i,
p. 83.
1857. Lcucocyon Gray, Proc. Zool. Soc, London, p. 512.
Tape species. — Canis lagopus Linnteus.
Gcograpliieal distribution. — Arctic region of l)oth Old and New
"Worlds ; in Europe south to southern Norway and Sweden.
Characters. — Skull intermediate in general form between that
of Canis and Vulpes ; occipital depth about one-third condylo-
basal length ; interorbital region more elevated than in Vulpes
owing to greater inflation of the frontal sinuses ; postorbital
processes thin, flat or slightly concave above, with liead-like,
overhanging edges ; dorsal profile of forehead rising abruptly
ALOPEX 319
above rostrum as in Conin ; teeth moderately heavy and large,
the length of cariiassial and upper molars together contained
about 2j times in palatal length ; canines and incisors inter-
mediate between those of Cams and Vulpes (see fig. 65) ; external
form fox-like, but ear short and rounded, not conspicuously
overtopping the surrounding fur.
liemdrks. — Although in most respects intermediate between
Canis and Vulpes the Arctic foxes form such a natural and
circumscribed group that it seems desirable to set them apart as
a distinct genus.* Half a dozen species have been described, two
of which come within the scope of the present work.
KEY TO THE EUROPEAN SPECIES OP ALOPEX.
Condylobasal length of skull about 130 in males,
124 in females (Scandinavia and Finland)... A. lagopus, p. 319.
Condylobasal length of skull about 120 in males,
114. in females (Spitzbergen) A. sjntzbcrgmcnsis, p. 324.
ALOPEX LAGOPUS Linnaius.
1758. [Canis] lagopus Linnaeus, Syst. Nat., i, 10th ed., p. 40 (Lapland).
1816. V[;ulpes] arctica Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1038
(Renaming of Canis lagopus).
1820. C[a7iis'] violpcs cxrulca Nilsson, Skand. Fauna, i, p. 88 (Lapland).
1827. ICanis lagopus] B argcntcus Billberg, Synopsis Faunae Scandinavia3,
p. 14 (Lapland).
1910. Vul2:ies lagopus Trouessart, Faune ]\Iamm. d'Europe, p. 96.
Ty^je locality. — Lapland.
Geo<jrapMcal distribtition. — Arctic portions of the mainland
of Europe and Asia ; in Europe south along the mountains of
Scandinavia to south-western Norway, and as an occasional
visitant as far as southern Sweden.
Dia(iuosis. — General characters as in the genus ; condylobasal
length of skull about 130 mm. in males, about 12 i mm. in
females.
External characters. — In general external charactei's Alopex
lagopus resembles Vulpes ruljX'S, though the muzzle is less
elongated, and the low, rounded ears (not extending to eye when
laid forward) impart a somewhat lui-fox-like appearance to the
head. Fur very dense, the underfur in summer about 12 mm.
deej) on back, nearly twice as deep and somewhat looser in
texture on sides (in winter longer throughout) ; longer hairs
rather sparse, not concealing underfur. Tail bushy, with
abuiulant underfur. Feet as in Cauis, but claws longer and
more .slender, and entire palm and sole covered with a dense
woolly growth of hair, 10-17 mm. deep in winter, shorter in
* See Collett, Norges Pattedyr, p. 275, 1911, for account of apparently
complete sterility of Arctic fox male with Red fox female, a fact which
indicates a fundamental physiological diflcrencc between the two animals.
320
CARNIVORA
suininer when it sometimes wears away sufficiently to expose
lialls of toes, and parts of pads.
Colour. — Summer pelage : ground colour of back, shoulders,
and outer side of legs drab, darkening to about prout-brown or
Fig. G3.
Alopcx laijoinia.
dark bister on feet, head, chin, interramial region and outer
surface of ears, the face thickly sprinkled with whitish hairs,
especially on cheeks and between eyes, the inteiramial region
tinged with slaty grey ; each of the longer hairs of back with
321
one cream-buff sub-terminal aiinulation, producing a noticeably-
speckled appearance throughout dark area ; flanks with a few
long, entirely butt' hairs ; sides of body and of neck light cream-
buff tinged with clay-colour, in striking contrast with dark areas,
the buff lateral area divided at shoulder by band about 60 mm.
wide where drab of back crosses to leg ; anteriorly the buff
lateral area extends to about level of ears where it abruptly gives
place to dark brown of head ; inner surface of ear light bufiy
grey ; underparts and inner surface of legs buffy greyish, slightly
contrasted with sides ; under surface of tail like sides of body,
upper surface essentially like back at base (where line of
demarcation is well defined), but becoming tinged with buffy
toward tip. Winter pelage : entirely white, tinged with yellowish
on throat, neck and face ; underfur and posterior surface of ear
light drabby grey. In the " blue " phase the entire animal is at
all seasons a bluish drab, usually washed with sepia on head and
feet, and sprinkled with pure white hairs on face, chin and throat.
Skull. — The skull is shorter and narrower than that of Vulpen
vulpes, but of nearly equal depth, a difference in form due in part
to the greater elevation of the interorbital region and in part to
the relatively greater depth of brain-case (depth equal to one-
third condylobasal length instead of noticeably less as in
V. milpes) ; muzzle less produced
than in the common fox and rela-
tively wider proximally ; zygomata
less abruptly spreading anteriorly,
so that the region of greatest
zygomatic breadth is noticeably at
glenoid level. In other respects
there is essential agreement with
the skull of V. vulpes. Anteorbital
foramen over space between 'pm^
and pm* ; auditory bulla relatively
as large as in the larger animal ;
depth of brain-case through bulla
equal to greatest breadth above
roots of zygomata ; postorbital pro-
cesses slightly less concave on
underside and somewhat more
flattened above ; mandible with
ramus relatively a little deeper
and more compressed than in the
common fox.
Teeth. — In general the teeth
closely resemble those of Vidpes
vulpes except for their slightly
smaller size. Incisors both above
and below with relatively wider crowns than in the common fox,
the cingulum of i^ and i? much more developed and forming a
Alopex lagopu.
322
CARNIVORA
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324 CARNIVORA
noticeable inner and outer secondary cusp (the former soon
wearing away in /') much as in Canis ; along posterior border of
crown the cingulum shows a strong tendency to develop irregular
tubercles, a condition not observed in Vulpes vulpes. Canines
and premolars with no special peculiarities, the form of the upper
carnassial quite as in V. vulpes. Lower carnassial with posterior
heel narrower than main portion of tooth but not otherwise
peculiar. First upper molar with metaconule relatively less
developed than in Vulpes vulpes.
Measurements. — For cranial measurements see Table, p. 322.
Specimens examined. — Ten, from the following localities : —
Norway: Roros, Trondhjem, 1; Tolgen, Hedemarken, 2; Dovre, 2;
Egersund, Stavanger, 1.
Sweden : No exact locality, 1 skull (U.S.N.M.).
Lapland : No exact locality, 3 skulls (B.M. and U.S.N.M.).
?. Roros, Trondhjem, Nor- Christiania Museum 95. 11. 14. 1.
way. (e).
skull. Egersund, Stavanger. K.H.Schaauning(c). 11. G. 3. 12.
2 9. Dovre. Christiania Museum 95. 11. 14. 2-3.
(E).
2 imm. Tolgen, Hedemarken. Christiania Museum 93. 3. 1. 1.
(e). 0. 5. 2. 1.
2 skuUs. Lapland. Wheelwright (c). 64.3.8.3-4.
ALOPEX SPITZBERGENENSIS Barrett-Hamilton and Bonhote.
1799. ? Canis fuliginosus Bechstein, Thomas Pennant's allgem. Uebersicht
d. vierfiiss. Thiere, i, p. 270 (Iceland).
1799. ? Canis grocnlandicus Bechstein, Thomas Pennant's allgem. Ueber-
sicht d. vierfiiss. Thiere, i, p. 270 (Greenland).
1898. Canis lagopus spitzhergcncnsis Barrett-Hamilton and Bonhote, Ann.
and Mag. Nat. Hist'., 7th ser., i, p. 287, April, 1898 (Spitzbergen).
T}'pe in British Museum.
1910. Vulpes lagopus spitzbcrgensis and ? V. lagopus fuliginosxis Trouessart,
Faime Mamm. d'Europe, p. 97.
Type locality. — Spitzbergen.
GeograpMcal distribution. — Spitzbergen ; also Iceland and
Greenland 1
Diagnosis. — Like Alopex lagopus but smaller, condylobasal
length of skull about 120 in males, about 114 in females.
Colour. — Type (summer pelage) : colour pattern well defined
and exactly as in A. lagopus, but dark areas wood-brown against
which the cream-bufi" annulations of longer hairs make no marked
contrast. Another skin, also in summer pelage, is a uniform
dark slaty drab throughout, the hind feet darker and more
brown ; .sides and underparts with a few long white hairs
(50 mm.) ; lijis with slight grizzling due to presence of short
white hairs.
Measurements. — For cranial and dental measurements see
Tables, pp. 322, 323.
VULPES
325
Specimens examined. — Six, all from Spitzbergen(B.M.and U.S.N.M.): —
6 skull. Spitzbergen. Stockholm Museum (e). 90. 8. 1. 2.
i, 9. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 2-3.
(96. 9. 2.3. 3. Type of species.)
$, 9 skulls. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 4-5.
Genus VULPES Oken.
1816. V[ulpcs] Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1033, in full on
p. 1034 and in index, p. 1268 (Vulpcs communis Oken = Canis
vulpcs Linnifius).
1821. Vulpes Bowdich, Anal. Nat. Classif. Mamm., p. 40 (Canis viilpes).
1857. Vulpes Blasius, Siiugethiere Deutschlauds, p. 178 (Sub-genus of
Canis).
Type species. — Canis vulpes Linnajus.
Geographical distrihution. — Northern portion of the northern
hemisphere from about the hmit of tree growth south to Morocco,
India and Mexico ; in Europe west to Ireland.
Characters. — Skull slender and low (depth of brain-case less
than one-third condylobasal length) ; interorbital region nearly
flat, the frontal sinuses scarcely inflated, the postorbital proces.ses
thin, slightly concave above, their edges overhanging and bead-
like ; dorsal profile of forehead rising very slightly and gradually
above level of rostrum ; dental formula as in Canis ; teeth
relatively light and small, the length of upper carnassial and
molars together contained about 2| to 3 times in palatal length,
the general character of cheek-teeth somewhat more trenchant
than in Catiis, the canines slender and elongated, the point of
upper tooth extending to about level of lower margin of mandi-
bular ramus when jaws are closed (fig. 65).
MemarJcs. — As thus restricted the genus Vulpes contains about
thirty-five forms, all jjeculiar to the northern hemisphere. Five
of these occur in Europe.
KEY TO THE EUROPEAN FORMS OP VULPES.
Size small, hind foot in adult male about 125 mm.,
condylobasal length of skull in both sexes less
than 130 mm. (Sardinia and Corsica) T'. ichnus,T, p. 336.
Size large, hind foot in adult male 135 to 165 mm.,
condylobasal length of skull in adult male 136 to
165 mm., in adult female 127 to 155 ram V. vulpcs, p. 326.
Teeth larger and more robust, the premolars tend-
ing to be slightly spaced or in contact, their
secondary cusps well developed (Scandinavia)... V. v. vulpes, p. 330.
Teeth smaller and less robust, the premolars tend-
ing to be widely spaced, their secondary cusps
usually obsolete or absent.
YeUowish and reddish tints bright ; posterior
half of back seldom much frosted with whitish ;
tail never clear grey (Central Europe) r. v. crucigem, p. 331.
Yellowish and reddish tints pale and dull ; pos-
terior half of back usually much frosted with
whitish ; tail often clear grey (Iberian Penin-
sula) V. V. yilacea, p. 333.
326
CARNIVORA
FtG. 65.
Incisors and canines from finnt, of fVon'-v (a), Alopex (i), and Vulpes (c).
VULPES VULPES Linnjeus.
(Synonymy under subspecies.)
Ty2je loralHii. — TTpsala, Sweden.
GeograpMcul distrihufion. — Europe from the Arctic coast to
the Mediterranean, and from Ireland eastward into Asia.
Diar/nosis. — Size large: hind foot, in adult male 135 to 165
mm.; condylobasal length of skull in adult male 135 to 165 mm.,
in adult female 127 to 155 mm.
External characters. — Form more slender and legs relatively
shorter than in Canis ; muzzle long and pointed ; ear high,
pointed, rising conspicuously above surrounding fur ; tail long,
VULPES 327
thick anfl bushy, with abundant underfur ; longer hairs of back
normally concealing the underfur ; feet as in Cauls, but with
the soles hairj' between the pads, the pads themselves sometimes
furred.
Colour. — General colour a yellowish brown brighter and more
inclined toward reddish along median dorsal region and on face,
duller and more yellowish or greyish on sides of body to shoulder
and on sides of neck to base of ear, the flanks and usually the
sides sprinkled with white hairs which may produce a decided
effect of frosting ; underparts very variable, ranging from
whitish to slaty black, rarely almost concolor with sides ; feet
dusky or blackish ; ear tawny or buff at base and on inner
surface, the terminal half of outer side black or very dark brown
in strong contrast with surrounding parts ; upper lip dull white.
Blackish and greyish variations not uncommon, especially at the
north.
Skull. — General form of skull slender and somewhat flattened,
with widely spreading, nearly parallel zygomata. Dorsal profile
almost flat from nares to slightly beyond midfUe of nasal bones,
then rising at a slight angle (10° or less) to or a little beyond
bregma, behind which it slopes away by an evenly convex curve
(distorted in old individuals by the development of the sagittal
crest) to lambda, which lies a little above level of middle of orbit ;
ventral profile essentially straight. Brain-case distinctly broader
than high, its outline ovate when viewed from above, the
lambdoid and sagittal crests well developed, the latter extending
forward about to bregma, where it di^'ides, sending a branch to
form posterior border of each postorbital process. Occiput
obliquely truncate, so that condyles are not visible from above,
but region between lambda and foramen magnum slightly if at
all concave. Floor of brain-case with no special peculiarities ;
auditory bulhe moderately and evenly inflated, slightly flattened
antero-externally, meatal tube short but distinctly indicated,
especially its hinder wall. Interorbital region flattened, with
median longitudinal groove, the postorbital processes prominent,
flattened, triangular, much shorter along posterior edge than
along antero-external edge, the margin slightly raised so that
the upper surface is some\\hat concave, the under surface so
abruptly concave that the process is much less thick than in
Alojjex and Cani.s. Rostrum moderately long (the distance from
orbit to gnathion about etjual to that from postorbital process to
lambda), rather abruptly narrowed proximally, so that the sides
are nearly parallel through a noticeable portion of their extent
(occasionally the sides diverge from region of greatest narrowing
to bases of canines) ; nasal slender, narrowing gradually back-
ward and extending nearly to level of middle of orbit ; nasal
branch of premaxillary extending to about middle of nasal and
usually not in contact with frontal ; posterior extremity of
maxillary extending slightly behind that of nasal ; anteorbital
328
CAENIVORA
foramen over posterior root of pm? or over space between pw/^
and ptu*. Palate relatively narrower than in the European
Fia. CG.
Viilpes vuljies. x i.
species of Alopex and Canis, its median posterior termination
about at level of middle of last mola.r ; mesopterygoid fossa
329
I
about twice as long as wide, the sides gradually converging
posteriorly ; haiiuilars slender, straight ; incisive foramina about
four times as long as wide, the median septum usually wider
than the foramen, the posterior margin a little in front of
posterior margin of alveolus of canine. Mandible slender, but
with no special peculiarities of form.
Teeth. — In general form and structure the teeth do not differ
very widely from those of Canis lupus. The canines are, however,
relatively much longer and more slender (fig. 65), the incisors are
weaker and narrower, and the premolars show a more decided
tendency to develop secondary
cusps. Upper incisors slender,
not closely crowded, their
crowns relatively narrower than
in either Canis lupus or Alopex
liujopus, the secondary cusps
obsolete, early disappearing
with wear, the cingulum barely
indicated ; lower incisors with
crowns less simplified than in
the upper teeth, the usual lon-
gitudinal groove present on
posterior surface, and i^ with
well developed outer basal lobe.
Upper canine slender, its dia-
meter at alveolus about 7 mm.,
its height when unworn usually
about three times as great ;
lower canine with diameter
contained about 2;V times in
height. Premolars with no
special peculiarities, their crowns
relatively narrower than in
Canis lupus, and secondary cusps
tending to be more developed,
the latter character varying in
different geographical forms.
Upper carnassial with inner
lobe better developed than in
Canis, and bearing a distinct terete cusp, its position a little
more forward and outward than in the dogs, so that it appears
to lie in or nearly in the main axis of the tooth. Lower carnassial
with 2)Osterior lieel essentially as broad as main portion of tooth,
the cu.sps es.sentially as in Canis, but general aspect of tooth more
trenchant. Ujiper molars differing from those of Canis in the
relatively smaller size of paracone and metacone, these cusps
apjiearing to stand in from border of crown, leaving a noticeable
cingulum beyond them ; paraconule obsolete. Second and third
lower molars with no marked peculiarities.
FIG. 67.
Vulpes vulpes. Teeth.
330
CARNIVORA
RcmarJiS. — With the material at hand it has been impossible
to reach wholly satisfactory conclusions with regard to the
number of local forms represented by the common foxes of various
parts of Europe. The existence of three such races, one in the
Scandinavian Peninsula, a second in Central Europe, and a third
in the Iberian Peninsula, seems well established ; but the status
of the forms inhabiting Italy and Greece is still in doubt.
VuLPES vtiLPKs VULPES Linuieus.
1758. [Canis] vulpes Liiiuffius, Syst. Nat., i, 10th ed., p. 40 (Sweden).
1758. [C'auis] alopex Liniiieus, Syst. Nat., i, 10th ed., p. 40 (Sweden).
1798. C[anis] vulpus Thunberg, Beskrifning pA Svenske Djur, Mamm., p. 7
(Variant of vulpcs).
1816. Vl^ulpes] vulgaris Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1034
(Renaming of vulpes).
1820. Canis nigro-argentcus Nilsson, Skand. Fauna, i, p. 91 (Lofoten
Islands, Norway).
1827. [Canis vulpus] y oiigrocaudatus Billberg, Synopsis Faunae Scandi-
navise, p. 12 (Uppland, Sweden).
1827. [Canis vul2ms] ( variegatus Billberg, Synopsis Faunas Scandinavise,
p. 1.3 (Uppland, Sweden).
1827. [Canis vulpus] i) Uncatus Billberg, Synopsis Faunae Scandinaviae,
p. 13 (Skane, Sweden).
1830. ? [Vulpes] communis Burnett, Quart. Jouru. Sei. Lit. Art, sxviii,
1829, p. 349 (Substitute for vulpes). Nomen nudum.
1898. Vulpes vulpes Thomas, The Zoologist, 4th ser., ii, p. 100, jMarch,
1898 (part).
1910. Vulpes vulpes Trouessart, Fauna Manun. d'Europe, p. 93 (part).
Type locality. — Upsala, Sweden.
Geographical distribution. — Scandinavian Peninsula.
Diagnosis. — Teeth larger and more robust than in the central
and southern races, the premolars tending to be slightly spaced
or in contact ; skull attaining maximum size for Eiu'opean
foxes.
Measurements. — For cranial and dental measurements see
Tables, pp. 334, 335.
Specimens examined. — Seven skulls from Sweden (for exact localities
see Table of cranial measurements), and eleven from Egersund, Stavanger,
Norway.
Remarks. — While I have seen no skins of this fox, the
distinctness of the race from those occurring in central and
southern Europe seems well established by the characters of
the teeth.
skull. Sweden. Wheelwright (c). 04. 3. 8. 2.
11 skulls. Egersund, Stavanger, K. H. Schaanning (c). 11. G. 3. 1-11.
Norway.
VULPES 331
VuLPEs VULPES CRUCiGERA Bechsteiii.
1789. [Canis] crucigera Bechsteio, Gemeinn. Naturgescb. Deutschlands,
I, p. 250 (Thiiringen, Germany).
1792. C[anis] Vulpes cilopex europaeus Kerr, Anim. Kingd., p. 142 (Bur-
gundy, France).
1797. Canis vulpes alba Borkhausen, Deutsche Fauna, i, p. 33 (Vogelsberg,
near Rudigshain, Hessen, Germany).
1797. Canis vulpes nigra Borkhausen, Deutsche Fauna, i, p. 33 (Hessen
and Thiiringen, Germany).
1801. Clanis] vl^idpes] lutea Bechsteiu, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. G28 (Thiiringen, Germany).
1801. C[a7iis] v[ulpes] cinerca Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 628 (Thiiringen, Germany)..
1832. Canis melanogaster Bonaparte, Iconogr. Fauna Ital., i, fasc. 1
(Neighbourhood of Rome, Italy).
1841. Vulpes hyponielas Wagner, Schreber's Saugthiere, Suppl., ii, p. 405
(Oberbayern, Germany).
1857. Canis vulpes Blasius, Siiugethiere Deutschlands, p. 191.
1861. ? V[2dpes] vulgaris meridionalis Fitzinger, Wisseusch.-pop. Natur-
gesch. der Saugeth., i, p. 194 (Dalmatia).
1910. Vulpes vulpes (part) and V. vulpes melanogaster Trouessart, Fauue
Mamm. d'Europe, pp. 93-94.
Type locality. — Thiiringen, Germany.
Geographical distribution. — Central and .southern Europe from
Ireland eastward and from the coast of the Baltic to the
Pyrenees, Italy and Greece.
Diagnosis. — Maximum size I'ather less than in V. v. vulpes,
and teeth distinctly smaller, the premolars rather widely spaced
and seldom if ever in contact ; general colour a bright yellowish
or reddish brown, the posterior half of back not conspicuously
frosted with whitish, and tail never clear greyish.
Colour. — In seventeen skins the general colour ranges from
nearly cinnamon-rufous to a light ochraceous-rufous, the sides
of neck and region immediately behind shoulder lighter than
median dorsal area (^in extreme in.stances clear buff with a
decided rufous tinge) ; posterior half of back with evident
white frosting in some specimens, scarcely any in others, but
this character never so pronounced as in average Spanish skins ;
underparts dull slaty overlaid with white, the slaty nearly always
predominating, except on throat, and not infrequently giving the
effect of an almost blackish tinge throughout, this apparently
not in the least dependent on regional or local climatic con-
ditions. In a flat skin from Cephalonia, Greece, the characteristic
slaty and white is confined to the throat and chin, all the rest
of the ventral region being a dull tawny-ochraceous like sides.
Two specimens from Tatoi, near Athens, taken in July, have
shed all the longer hairs of the back, leaving only the velvety
underfur. This is of the usual colour, a dull umber brown, in
one specimen with a slaty cast.
Sltill and teeth. — The skull and teeth do not attain so srreat
332
CARNIVORA
size as in true Vuljies ruJpes, and the small premolars are seldom
in contact, the spaces between them usually conspicuous.
Measurements. — Adult female from Cappagh House, Water-
ford, Ireland : head and body, 613 ; tail, 340 ; hind foot, 146 ;
ear from meatus, 90. Adult male and female from Tunbridge
Wells, Sussex, England : head and body, 690 and 630 ; tail,
343 and 370; hind foot, 160 and 141 ; ear from meatus, 95 and
82. Adult male from Ingelheim, Rheinhessen, Germany : head
and body, 578 ; tail, 440 ; hind foot, IGO ; ear from meatus, 98.
Adult male and female from Haute-Garonne, France : head and
body, 610 and 610; tail, 370 and 350 ; hind foot, 150 and 135 ;
ear from meatus, 94 and 89. Two adult males from Porlezza,
Como, Italy (Ghidini) : hind foot, 150 and 148-6. Adult male
from Borghetto S. Spirito, Italy : head and body, 745 ; tail, 380 ;
hind foot, 157. Adult male from Zinnigas, Siliqua, Sardinia
(measured from mounted specimen, Genoa) : head and body, 700 ;
tail, 300; hind foot, 144± ; ear from meatus, 83.
Specimens examined. — Fifty, from the following localities : —
Ireland : Kilmanock, Wexford, 2; Cappagh House, Waterford, 1.
Scotland : Ben Nie, Sutherland, 1 ; Inversanda, Ardgour, 1.
England : Northumberland, 1 ; Thame, Oxford, 2 ; Hassocks, Sussex, 1 ;
Tunbridge Wells, Sussex, 3 ; Ditchling, Sussex, 1 ; Mayfield, Sussex, 1
(Grant).
France : As-les-Thermes, Ari^ge, 3 ; Caterille, Haute-Garonne, 1 ; Pic
Sessire, Haute-Garonne, 1 ; St. Aventin, Haute-Garonne, 1.
Germany : Ingelheim, Rheinhessen, 1 ; Nuremberg, Bavaria, 1
(U.S.N.]\I.) ; Grossgraben, Silesia, 1 (Breslau) ; Rieseugiberge, Silesia, 1
(Breslau) ; southern Germany, 3.
Austria-Hungary : Trentino, 1 (Genoa) ; Gazza, Treutino, 1 (Genoa) ;
Vigolo Vattaro, Trentino, 1 (Genoa).
Switzerland: Geneva, 1 (Ghidini); Valais, 1 (Ghidini).
Italy : Porlezza, Como, 4 (Ghidini) ; Garbagna, Piedmont, 1 (Genoa) ;
Borghetto S. Spirito, 1 (Genoa) ; Vargo, Liguria, 1 (Genoa) ; Torriglia,
Liguria, 1 (Genoa) ; Cornigliano, Liguria, 2 (Genoa) ; near Genoa, 1 ;
Molasana, 1 (Genoa) ; Tuscany, 1 ; Pisa, 1.
Sardinia: Zinnigas, Siliqua, 1 (Genoa).
Greece : Cephalonia, 1 ; Tatoi, near Athens, 2.
Remarlcs. — The fox of Italy appears to be the same as that of
Central Europe, though further material from the southern
portion of the peninsula may show that it should be distinguished.
The three specimens from Greece are in such unsatisfactory
condition of pelage that their status cannot be determined with
any degree of certainty. If they represent a peculiar local race
this should probably take the name vteridioiiaUs Fitzinger.
9. Ben Nie, Sutherlandshire, E. R. Alston (p). 79. 9. 2.5. 80.
Scotland. {H. Brown.)
6. Inversanda, Ardgour. H. Leigh (c & p). 1. 16. 12. 1.
2. Kilmanock, Wexford, G. Barrett-Hamilton 6. 6. 20. 1.
Ireland. (c & p). 9. 12. 15. 4.
?. Cappagh House, Waterford. R. J. Ussher (c & p). 96. 12. 28. 1.
1. Northumberland, England. Rev. H. H. Slater 0.2.24.5.
(c & p).
vuLPES 333
2 6. Thame, Oxfordshire. Hon. N. C. Roths- 0. 10. 31. 1-2.
child (p).
2 (J. Ditchling, Sussex. Guy Dollman /9. 11. 3. 1.
(c & p). \10. 9. 13. 1.
3. Tunbridge Wells, Sussex. G. H. B. Grant (c). 1. 2. 1.5. 1-3.
2(5,9. Ax - las - Thermes, Arifege, V. Builles (c & p). 8.3.27.6-7.
Franco. 8. 3. 27. 14.
9. Caterille, Haute-Garonne, O. Thomas (p). 8. 7. 15. 2.
900 m. [A. Robert.)
9. PicSessiro,Haute-Garonno. 0. Thomas (p). 8. 7. 15. 3.
[A. Robert.)
6. St. Aventin, Haute-Ga- O. Thomas (p). 8. 7. 15. 4.
ronne, 900 m. (A. Robert.)
i. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 15.
Germany.
3 skulls. South Germany. Dr. A. Giinther (c). 59. 9. 6. 83, 88.
175. k.
S. Genoa, Liguria, Italy. MarquisG. Doria (p). 88. 12. 1. 3.
skull. Pisa. Zool. Soc. Mus. 68. 5. 4. 126.
lyg. S. Italy. (Prof. Savi.) Zool. Soc. Mus. 55.12.24.240.
9. Tuscany. Purchased (Dr. 45. 7. 22. 15.
RiippeU).
6, 9. Tatoi, Athens, Greece. Hon. N. C. Roths- 8. 10. 2. 22-23.
(C. Mottaz.) child (p).
VuLPES VULPES siLACEA Miller.
1907. Vulpcs vidpes silaceus Miller, Ann. and ^Mag. Nat. Hist., 7th ser.,
XX, p. 393, November, 1907. Type in British Museum.
1910. Vulpes vulpes silaceus Trouessart, Faune Mamm. d'Europe, p. 45.
Type locality. — Near Silos, Province of Buri^os, Spain.
Geographical distribution — Iberian Peninsula.
Characters. — Size about equal to that of V. v. vulpes, but
teeth noticeably less enlarged, the wider spacing of the premolars
especially evident ; general colour a dull buff without bright
yellowish or reddish tints, the posterior half of back conspicuously
frosted with whitish (except in abraded pelage) the tail often a
clear greyish.
Measurements. — Type (adult male) : head and body, 750 ;
tail, 370; hind foot, 150. Adult female from near the type
locality: head and body, 670; tail, 330; hind foot, 125. Old
male from Elche, Alicante : head and body, 770 ; tail, 480 ;
hind foot, 160; ear from meatus, 113.
Specimens examined. — Eighteen, from the following localities in
Spain : Olot, Gerona, 1 (probably not typical) ; vicinity of Silos, Burgos, 5 ;
near Burgos, 1 ; Arrechavaleta, Vitoria, 1 ; Torres del Alio, Coruna, 1 ;
Madrid, 1 ; near Seville, 3 ; Goto Dofiana, Huelva, 3 (B.M. and U.S.N.M.) ;
Elche, Alicante, 1 ; Barracas, Gastellon, 1.
Bemarks. — The Spanish fox is well characterized by its light
colour, large size and rather small, widely-spaced premolars as
compared with those of the Scandinavian form.
2 6. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 46-47.
(8. 8. 4. 46. Type of subspecies.)
(J, 2 9, 1. Burgos. Rev. Saturio Gon- 8.7.7.12-15.
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336
CARXn-ORA
9. Arrechavaleta, Vitoria. 0. Thomas (p). S. 2. 9. 49.
(y. Gon.'alez.)
6. Torres del Alio, Comua. Dr. Y. L. Seoane 94. 5. 29. 1.
(c & p).
6. :Madrid. M. de la Escalera (c). S. 9. 24. 7.
6, 9. SeviUe. {A. Ruiz.) Lord Lilford (p). 95. 9. 4. 7-8.
2 i. Goto Doiiana, Huelva. A. Chapman (c & p). 8. -3. 8. 4-5.
i. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 48.
i juT. Barracas, Castellon. O. Thomas (p). 8. 2. 9. 50.
(N. Gonzalez.)
VULPES ICHNUS.E Miller.
1907. Yulpes icliniisx Miller. Aun. and Mag. Xat. Hist., 7th ser., xx, p. 391,
November, 1907. Type in British Museum.
1910. Tidpcs vuJpcs ichniisx Trouessart, Faune Mamm. d'Europe, p. 94.
Type locality. — Sarrabus, Sardinia.
Geographical distribution. — Sardinia and Corsica.
Diagnosis. — Smaller than any of the races of Vidpes vulpes ;
both hind foot and condylobasal length of skull in adult male
less than 130 mm., ear from crown 60 to 70 mm.
Colour. — Face and head dark rufous becoming Hghter and
more dull on base of ears and on neck, and fading to ockraceous-
rufous on shoulders and back ; sides of neck, outer surface of
upper arm and region just behind axilla still lighter, a ta^^Tiy
butf, hairs of underfur on back drab grey at base, tawny clay
colour at tip ; longer hairs of head, sides, and back (behind
shoulders) much speckled by huffy white subtermmal areas about
5 mm. in length, the extreme tips reddish : feet and legs
ochraceous-rufous, slightly clouded with blackish and a little
speckled with bufiy white : tail like back above, the tawny
gradually fading out through a bufiy grey to the whitish buff"
tip, the longer hairs except in pencil black tipped (30-40 mm.) ;
underparts to front legs huffy wliitish tinged with hair-brown,
this nearly clear on middle of throat ; rest of underparts a
mixture of hair-brown and dull tawny, the latter predominating
along sides.
Measurements. — Type (adult male) : hind foot, 123 ; ear from
meatus, 70. Adult male and female from the type locality :
head and body, 640 and 590 ; tail, 280 and 290 : hind foot,
127 and 125 : ear from meatus, 74 and 71. Adult female from
Siliqua : head and body, 600 : tail, 350 ; hind foot, 127 : ear, 74.
Adult from near St. Florent, Corsica : hind foot, 123 : ear from
meatus, 73.
Specimens examined. — Eight, from the following localities : —
Coksica: Grotto Campu Consule, near St. Florent, N.W. Corsica, 1
(Major).
Saedesia : No exact locality, 1 (Turin) ; Lanusei, 1 ; Sarrabus, 3 (B.M.
and Genoa) ; Zinnigas Siliqua, 2.
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342
CARNIVORA
Geograplrical distribution. — Northern temperate portions of
Old World ; in Europe from central Scandinavia to the
Mediterranean, and from Ireland eastward.
Characters. — Skull narrow and high (depth of brain-case
much more than half mastoid breadth), the zygomatic arches not
widely expanded, and postorbital region not specially narrowed
(distance from point of greatest narrowing to zygoma less than
breadth of postorbital constriction) ; rostrum elongate, the
distance from orbit to gnathion about three-fourths length of
brain-case ; auditory bulla elevated along inner margin, but else-
where somewhat flattened, the meatal tube distinct ; paroccipital
process robust, standing out conspicuously behind bulla ; dental
formula : i ||, c t], jmi ti,* m ^^ = 38 ; teeth not specially enlarged
as compared with width of rostrum and palate (greatest trans
verse diameter of upper carnassial equal to a little more than
one-third distance between carnassials) ; three small premolars
(one upper and two lower) capable of trenchant action ; upper
carnassial with crown triangular in outline, the cutting portion
consisting of a single large antei'ior cusp with its slightly developed
anterior commissure and more distinct posterior commissure, the
crushing portion represented by the obliquely sloj^ing inner base
of this cusp and ridge ; upper molar rhombic in outline, its crown
area about three times that of carnassial, its greatest diameter
in axis of tooth- row, the four primitive cusps present ; lower
carnassial with the three anterior cusps well developed, sub-equal,
the posterior heel decidedly larger than anterior triangle ;
external form heavy, the head pointed, the ears short but plainly
visible, the body thick, the tail short, not muscular ; feet sub-
plantigrade, the toes with long fossorial claws and without webs ;
fur coarse and loose.
Reiiiarlcs. — The genus Meles contains about half a dozen
named forms, the status of several' of which is at present not
clearly understood. Two species occur in Europe.
KEY TO THE EUROPEAN FORMS OF MELES.
Maxillary tooth-row, exclusive of incisors, about
35 mm. ; auditory bullae rather strongly in-
flated, the inner Isorder not ridge-like (Crete) M. arcalus, p. 352.
Maxillary tooth-row, exclusive of incisors, about
40 mm. ; auditory bullae slightly inflated, the
inner border ridge-like (distribution general) M. mcles, p. 343.
General colour moderately jpale ; teeth averaging
' smaller, less frequently attaining maximum
size (Central and southern Europe) M. m. meles, p. 348.
General colour slightly paler ; teeth averaging
larger, and more frequently attaining maxi-
mum size (Iberian Peninsula) M. m. marianensh! , p. 352.
* In adults usually '-p-" owing to the early disappearance of the small
343
MELES MELES Linnjfius.
(Synonymy under subspecies.)
Type locality. — Upsala, Sweden.
Gi'cxjrapMcal distribution. — I^urope, west to Ireland, south to
the Mediterranean and north to central Scandinavia. Eastei'n
limits of range not known.
Diagnosis. — Size large (upper length of skull in adult more
than 120 mm.; maxillary tooth-row, exclusive of incisors, about
40 mm. or more ; hind foot about 90 to 110 mm.) ; auditory bulla
moderately inflated, the highest region close to inner margin and
forming an evident longitudinal ridge, the region between ridge
and meatus noticeably concave ; teeth large, the two lower molars
together 22 mm. or more in length ; postero-external border of
upper molar usually convex, though occasionally straight or
slightly concave.
External characters. — ^Pelage coarse and loose, practically
without underfur, the hairs at middle of back about 60 mm. in
length, those on sides longer ; underparts scantily haired, the
skin usually visible ; palm bare, usually with a slight bristly
pubescence at middle in area between tubercles ; a large
tubercular mass at base of digits, convex in front, concave behind,
wider on outer border than on inner border, showing no tendency
to trilobation ; a roundish tubercle about half as large at postero-
external border of palm, separated from anterior mass by a wide
space ; a small, ill-delined pad at base of thumb ; sole densely
haired from heel to a little beyond middle, then completely naked ;
plantar tubercles essentially like those on palm except that
small pad at base of hallux is absent, and the two large masses
tend to coalesce, owing to absence of the wide intervening space ;
surface of pads on both palms and sole finely rugose, this
especially noticeable in dried specimens ; muzzle pad entirely
naked, but separated from upper lip by a narrow hairy band.
Mammte : a 2-2, * 1-1 = 6.
Colour. — Back and sides a coarse grizzle of black and buffy
white, the black usually predominating on back, the lighter
colour on sides. Throat, median ventral area and all four legs
and feet black or blackish ; face, chin and entire neck (except
underside) clear whitish except for a broad dark brown or black
band beginning on each side about 15 mm. behind nostril pad
and extending back, including eye and ear, to middle of neck,
where it fades in.sensibly into colour of back ; width of white
median area on face usually greater than that of the dark lateral
stripe, and about ecjual to that of the light area between lateral
stripe and dark ventral area : ear black, its anterior border
white in strong contrast ; eye usually a little below middle of
dark baud ; tail like back at base, soon fading to soiled white.
Skull. — Except for the greatly (.leveloped sagittal crest, the
344 CARNIVORA
height of which in old individuals is equal to nearly half inter-
orbital breadth, the skull of Meles meles is rather smooth and
without marked angularity. Depth of brain-case (without crest)
and depth through posterior portion of orbit nearly equal, and
about two-thirds mastoid breadth ; depth at front of nasal about
two-thirds that at orbit. Dorsal profile (without crest) nearly
flat from middle of brain-case to interorbital region, convex over
posterior half of brain-case ; rostrum sloping downward at angle
of about 30° with surface of brain-case. When crest is fully
FIG. 08.
Meles melex.
developed it reduces this angle to 20° or less. It is slightly
convex anteriorly, rather abruptly convex posteriorly, the hinder-
most portion slightly overhanging. Postorlntal piocesses short
but well developed, sharply outlining the small orbit (greatest
diameter of orbit slightly less than half that of interorbital
region) which, though widely open posteriorly, is more than half
encircled with bone. Rostrum moderately long, the distance
from gnathion to front of zygoma about equal to width of palate
including molars, and continued about 3,'; times in condylobasal
length of skull. Zygomata widel}' spreading posteriorly, narrow
MELES 346
anteriorly, the widest portion opposite front of glenoid fossa, the
arch Aery slightly bowed upward. Anteorbital foramina rather
large, sub-circular in outline, their greatest diameter about half
that of orbit. Ventral prolile of skull slightly concave through-
out. Palate moderately wide, the distance between molars
contained about 2 J times in distance from gnathion to level of
posterior edge of molar. Incisive foramina small, at level of
space between canine and P, elongate pyriform in outline, the
minute median foramen a little behind middle. Posterior exten-
sion of palate about equal to distance between molars, and slightly
more than half that from molar to hamidar. Mesopterygoid
Fig. 69.
Meles meles. x i.
space short and wide, its length only a little more than distance
between tips of hamulars. Auditory bullte slightly inflated,
irregularly triangular in outline, the meatus lying in angle formed
by the large, forward-projecting mastoid process. The surface is
irregular, with evident ridge near inner margin along crest of
most highly inflated region, this ridge often terminating anteriorly
in a bluntly pointed projection. Length of flattened portion
extending inward from meatus about equal to transverse diameter
of inflated portion of bulla, but the two regions not sharply
defined ; least distance between bulla> about \}, times diameter of
inflated portion ; paroccipital process short, triangular (not ridge-
like), its extreme base applied to posterior border of bulla.
346
CARNIVORA
Mandible heavily built, the I'amus nearly straight, the lower
border with the usual upward curve posteriorly, its depth at
middle about one-third that through coronoid process ; angular
riQ. 70.
Meles meles. x J.
process short, thick, and ill-defined, lying close to base of articular
process ; coronoid process squarely truncate above, its height
above articular process about equal to width at articular level.
Teeth. — The teeth are moderately large relatively to size of
skull, the incisors and canines rather short, the crown area of the
upper molar fully double that of any of the other maxillary teeth,
this last peculiarity unique among the European Mitstelidse. Upper
incisors robust, the crowns somewhat higher than wide when
viewed from in front, the teeth closely crowded in a slightly
convex row, the longitudinal diameter of crown a little greater
than transverse diameter ; i^ and r sub-equal, the latter slightly
the larger, their anterior surface convex with two faint longi-
tudinal grooves soon disappearing with wear, the cutting edge
entire, the posterior surface concave with narrow but distinct
heel ; P with crown area about double that of P and nearly half
347
that of canine, its crown higher than in the other incisors,
owing to lower insertion of root, but extremity of cutting edge
not extending beyond level of smaller teeth ; laterally the cutting
edge is continued down outer-posterior margin of shaft where it
functions against lower canine. Lower incisors not so large as
upper, forming a continuous, nearly straight row between
cauiiaes, the shafts straight, sloping obliquely forward, the root
of 1.2 implanted behind the others ; cutting edge irregularly
2-lobed, that of i^ longer than the others. Canines rather short
and weak, the shaft simple, that of upper tooth nearly straight,
that of lower tooth strongly curved backward, the enamel surface
of both essentially smooth, though that of lower canine shows
indications of a rudimentary cingulum in front. Anterior
premolar both above and below a minute or spicular tooth closely
crowded between canine and
second premolar, pm^ early
deciduous, pvii usually more
persistent. Second premolars
similar to each other, the crown
area about equal to that of P,
the outline irregularly oval when
viewed from above, the cusp
about as high as long, its apex
sHghtly in front of middle of
crown ; rootof each tooth single,
that of pm.T, showing a tendency
to become divided longitudi-
nally. Other small premolars
(pm^, pvi^ and j^^'^i) alike in form,
the crowns laterally compressed,
triangular when viewed from
the side, with apex slightly in
front of middle, jj)«^ and pm^
nearly alike in size, prii^ some-
what longer ; cingulum very slightly developed, not forming
secondary cusps. Upper carnassial triangular in outline, the
posterointernal border longest ; outer, trenchant portion of teeth
consisting of a main anterior cusp resembling ^jjh^ in size and form,
joined by a nearly horizontal commissure to a moderately high
posterior cusp ; cingulum low but bearing three distinct secondary
cusps, one at anterior base of main cusp, one (sometimes obsolete)
near middle of antero-internal border, and the third and largest
at middle of posterointernal border. Lower carnassial with
anterior triangle well developed, the three cusps sub-equal in
height, the metaconid subterete, the protoconid more compres.sed
than paraconid, its commissure slightly longer than that of the
anterior cusp ; crushing portion of crown longer and wider than
anterior triangle, its inner portion occupied by a large basin-like
concavity, its border with two large cusps on outer edge (of
Fig. 71.
Meles meles. Teeth.
348
CARNIVORA
which the anterior is the larger), a large cusp similar to postero-
external cusp at middle of inner margin ; posterior margin
occupied by three or four minute cusps soon disappearing with
wear, and a similar minute cusp sometimes in angle behind
protoconid and nietaconid. Second lower molar flat, terete, its
area about half that of crushing portion of carnassial ; middle of
crown with basin-like concavity ; margin with small but evident
antero-external and postero-external cusps, and a smaller
elevation at middle of inner edge. Upper molar rhomboidal
in outline, the inner and outer margins parallel, the former
nearly li times as long as latter; outer border occupied by two
rather large, subterete, conical cusps, the anterior (pai'acone)
slightly larger than posterior (metacone), so that greatest trans-
verse diameter of crown is slightly behind anterior border ;
remainder of crown a shallow basin-like concavity with rugose
surface and raised, irregularly nodulate margin, the middle of
concavity crossed by a ridge, convex internally, extending from
anterior base of paracone to posterior base of metacone and
bearing from three to five small cusps soon obscured by wear,
the three anterior separated from two posterior by a deep angle ;
the anterior group probably representing the protocone, the
posterior group the hypocone.
liemarJcs. — Two moderately well differentiated local races are
represented by the material that I have examined, one confined
to the Iberian Peninsula, the other general in distribution.
British specimens do not differ appreciably from Swedish
examples of the typical form.
Meles meles meles Linnseus.
1758. [TJrsus'] meles Linnaeus, Syst. Nat., i, 10th ed., ]}. 48 (Sweden).
1785. [MelesJ taxiis Boddaert, Eleuchus Animalium, i, p. 80 (Europe).
1789. [Urstts meles] /3 alba Gmelin, Syst. Nat., i, 13th ed., p. 102.
1789. [Ursus meles] y maculata Gmelin, Syst. Nat., i, 13th ed., p. 102.
1808. Taxus vulgaris Tiedemann, Zoologie, i, p. 376 (Renaming of Ursus
meles).
1816. Meles europn'us Desmarest, Nouv. Diet. d'Hist. Nat., iii, p. 465
(Renaming of Ursus meles).
1822. Taxus meles F. Cuvier, Hist. Nat. des ^Nlamm., iii, fasc. 36, January,
1822.
1827. [Meles] communis Billberg, Synopsis Faunae Scandinavise, p. 16
(Renaming of Ursus meles).
1827. [Meles conwiunis] & caninus Billberg, Synopsis Faunae Scandiuavise,
p. 17 (Scandinavia).
1857. Meles taxus Blasius, Saugethiere Deutschlands, p. 204.
1894. Meles meles Dahl, Die Heimat, iv, p. 125, June, 1894.
1899. M{eles] m{eles] typicus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., iv, p. 384, November, 1899.
MELES 349
1906. Meles meles britannicus Satunin, Mittoilungen des Kaukasi.schen
Museums, ii, p. 115 (Based on the cranial measurements of
xf r'4 specimens recorded by Barrett-Hamilton, Ann. and Mac.
iQin ,.f*'-H>stv Tthser., IV p.384). Co-types in British Museum.
1910. Meles meles and M. meles britannicus Trouessart, Faune Mamm
d'Europe, p. 69.
Type locality. — Upsala, Sweden.
Geographical distribution.—Centvul and southern Europe from
southern Sweden to the Pyrenees and Italy, and from Ireland
eastward ; eastern limit of range not known.
Characters.~S>knll and teeth moderately large, the teeth
rarely attaining maximum size for the species ; colour moderately
light, the sides not often conspicuously whitish.
3Ieasuremcnts.— Adult male and female from Woodpark
Ga way, Ireland : head and body, 686 and 618; tail, 153 and
loO; hmd foot, 108 and 102; ear from meatus, 48 and 45
Adult female from Boucoune, Gers, France : head and body 670 •
tail, 170 ; hind foot, 100 ; ear from meatus, 45. Adult female
from Ingelheim, Rheinhessen, Germany : head and body 672 •
tail, 180; hind foot, 104; ear from meatus, .50. For cranial
measurements see Table, p. 350.
Specimens cxnmi7ied.-Thirty-&ve, from the following localities •-
Ireland : Woodpark, Galway, 2. '
England: Burnley, Lancashire, 1; Ross, Hereford, 1; Kentchurch
Herefordshire, 4; Tetworth, Oxfordshire, 1 ; Ibstone, Buckingham hire 1-
NoRw;v''T;'= B'?^°»' Devonshire; Cornwall 1; noexactloca ty,4:
^orway: Holme, Mandal, 7 ; Egersund, Stavanger 2
bWEDEN : Middle Sweden, no exact locality, 1 (Stockholm) ■ TAkern
Phaxce : Poret de Bouconne, Gers, 1
Bavarlafr"'' Ingelheim, Rheinhessen, 1; Strass, near Burgheim,
Italy : Liguria, 1 (Genoa).
6, 9. Woodpark, Galway, Ire- R. p. Hibbert (p) 95 6 ^ i
^^''^- , 95. 12. 3. 1.
o „„ , -- , .\(^o-typesoiM.7n.britannicus,^a.t\xmn.)
V. Burnley, Lancashire, Hon. N. C. Roths- 10 31 4
England. child (p). ■ ■ • •
i. Ross, Herefordshire. Hon. N. C. Roths- 0. 10. 31. 3.
o o ■ i T^ , child (p).
■»■, 3 ]uv. St. Kentchurch, Hereford- W. E de Winton
shire. (c & p).
i. Tetworth, Oxfordshire. Hon. H. Parker
6. Ibstone, Buckingham- A. H. Cocks (c & p) 4 1 25 1
shire. ^ ' • • . .
i. Horsham, Sussex. E. C. Hawes (c & p) 7 4 13 1
w it'- ^ ^ictou, Devonshire. J. C.Stagdon(c& p). 97.2 23 1
Sr^ S°T'^11- LordLilford(P). ^ 89.10.28.1.
skull. Sweden. Purchased (Wheel- 64. 3. 8. 1.
If 00 Wright).
4 <5, i 9. Holme, Mandal, Norway. R. J. Cuninghame 8. 8 9 7-12
(c & p) '
2 skulls. Egersund, Stavanger. K. H. Schaanuing /ll. 6. 3. 13
(c)' (11.16.23.2.
350
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352 - CARNIVORA
?. Foret de Bouconne, Gers, O. Thomas (p). 8. 7. 15. 5.
260 m. France. (A.
Robert.)
9. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 16.
Germauv.
Meles meles maeianensis Graells.
1897. [Meles taxus'] var. marianensis Graells, Mem. Real Acad. Cien.,
Madrid, xvii, p. 170 (Central Spain).
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7tli ser., iv, p. 384, November, 1899 (Seville, Spain). Type
in British Museum.
1910. Meles meles mecliterraneus Trouessart, Faune Mamm. d'Europe, p. 70.
Tyjje locality. — Central Spain.
Geographical distribution. — Iberian Peninsula.
Diagnosis. — Skull and teeth large, the teeth frequently attain-
ing maximum size for species ; colour light, the sides often
conspicuously whitish, especially in region bordering dark ventral
area.
Measurements. — For cranial and dental measurements see
Table, p. 351.
Specimens examined. — Seven, from the following localities in Spain ;
Arrechavaleta, Vitoria, 1 ; Quintanar de la Sierra, Burgos, 2 ; near
Seville, 2 ; Goto Doiiana, Huelva, 2.
Benmrks. — Although not very strikingly differentiated, the
large-toothed, pallid, Iberian badger seems worthy of recognition
as a geographical race.
6. Arrechavaleta, Vitoria, N. Gonzalez (c). 8.7.7.18.
Spain.
6, 9. Quintanar, Burgos. Rev. Saturio Gou- 8. 7. 7. 19-20.
c?, ?. Seville. (A. Ruiz.) Lord Lilford (p). 95. 3. 3. 7-8.-
(95. 3. 3. 7. Type oi mediterraneus Barrett-Hamilton.)
<?, 9. Goto DoSana, Huelva. A. Chapman (p). 8. 3. 8. 6-7.
MELES ARCALUS Miller.
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and ^Nlag. Nat.
Hist., 7th ser., iv, p. 131, November, 1899 (part).
1906. Meles meles mediterraiieus Bate, Proc. Zool. Soc, London, 1905, ii,
p. 318, April 5, 1906.
1907. Meles arcalus Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 394,
November, 1907. Type in British Museum.
1910. Meles arcalus Trouessart, Faune Mamm. d'Europe, p. 70.
Tj/jje locality. — Lassethe Plain, Crete.
Geographical distribution. — Island of Crete.
Characters.— Size small (upper length of skull in adult less than
110 mm. ; maxillary tooth-row, exclusive of incisors, about 35 mm. ;
353
DENTAL MEASUREMENTS OE MELES MELES AND M. ARC ALUS.
Locality.
Number.
Sex.
Upper molar.
Lower molar
(first).
1 M. meles meles.
Sweden : no exact locality
64. 3. 8. 1
9
13-0 X 11-4
15-4 X 7
Wegeliu ....
Stockholm
6 ?
15-0 X 13-0
16-2 X 7
8
Skane ....
i '!
15-2 X 13-0
18-0 X 8
2
Ulriksdal ....
,,
<5juv.
13-4 X 11-6
15-8 X 7
S
Sm&land ....
V ?
14-2 X 11-8
16-2 X 7
4
Orebro ....
i iuv.
14-2 X -il-S.
15-8 X 7
Takern ....
6 ■>
13-G X 11-2
15-6 X 7
2
Norway : Holme, Mandal
152622
6
li-6 X 10-4
14-0 X 6
4
»» »»
8. 8. 9. 9
6
12-8 X 11-8
14-G X 7
8. 8. 9. 10
6
12-6 X 11-2
14-6 X 7
M '*
8. 8. 9. 11
9
11-2 X 10-8
14-0 X 6
8
8. 8. 9. 12
9
12-2 X 10-4
14-0 X 7
Egersund, Stavanger
11. G. 3. 13
15-0 X 11-4
14-8 X 7
England : Ross, Hereford
0. 10. 31. 3
6
13-2 X 11-4
16-0 X 8
Cornwall
98. 10. 28. 1
9?
14-0 X 12-0
16-6 X 7
8
no exact locality
2116
<5?
13-4 X 12-0
16-4 X 8
., ,, , .
21]f
9?
14-0 X 12-4
16-2 X 8-2
„ . .
—
14-2 X 12-2
13-G X 11-8
—
Burnley, Lancashire
0. 10. 31. 4
9
14-4 X 12-2
16-2 X 8-2
Cornwall
—
15-0 X 12-2
18-0 X 8-2
Ireland : Woodpark, Co. Galway
95. 12. 3. 1
15-6 X 12-8
17-0 X 7-8
,, ....
95. 6. 2. 1
14-8 X 12-2
16-0 X 7-8
Denmark: Zealand.
Andersen
6
14-0 X 13-2
17-0 X 7-8
Copenhagen*
6
15-0 X 12-6
18-5 X 8-2
,,
6
15-0 X 13-7
18-3 X 8-3
,, .
,,
6
15-5 X 13-6
18-0 X 9-0
,, .
9
14-6 X 12-5
17-5 X 7-7
various localities, i ^laxi- 1
13 specimens : \ mum j
16-3 X 13-6
_
) Mini-»
ImumJ
13-0 X 12-0
various localities, (Maxi-i
17 specimens : \ mum 1
18-7 X 8-5
1 Mini- 1
" ^mumf
15-0 X 7-0
[Franco: Bouconne, Gers.
8. 7. 15. 5
9
14-0 X 12-0
lG-2 X 7-2
Germany: Ingelheim, Rheinhessen .
8. 11. 2. 16
9
14-8 X 12-8
l'^-2 X 7-8
Italy : Borghctto S. Spirito, Liguria
33 Genoa
9
13-G X 110
15-6 X 7-2
M. meles marianensis.
Spain : Quintanar, Burgos
8. 7. 7. 19
6
15-2 X 12-6
17-4 X 8-G
; Seville ....
95. 3. 3. 7t
6
16-0 X 12-0
16-6 X 7-8
,,
95. 3. 3. 8
9
14-6 X 12-6
16-8 X 8-2
Goto Dofiana, Huelva .
8. 3. 8. G
6
16-4 X 13-2
17-8 X 8-0
....
8. 3. 8. 7
9
14-8 X 12-0
17-6 X 8-2
1 M. arcalus.
|Orete
5. 12. 2. 17t
9 juv.
13-6 X 11-2
15-4 X 7-2
5. 12. 2. 38
ad.
13-8 X 11-G
• -Measured l.y H. Wlnge.
f Type of iiiediten-aiieiii' Banett-Hamilton.
t Type.
354
CARNIVORA
hind foot about 25 mm.) ; auditory bulla strongly inflated, the
highest region near middle of bulla proper (exclusive of meatal-
tube) and so broadly rounded as not to form a longitudinal
ridge, the region between highest portion and meatus not notice-
ably concave ; teeth smaller than in Meles meles (lower molars
together about 20 mm. in length), with smaller cusps tending to
be more strongly developed ; metacone of m^ relatively larger
than M. meles, so that breadth (jf crown through this cusp is as
great as or greater than that through paracone ; colour as in
Meles meles marianeiisis or slightly ^laler.
Measurements. — For cranial and dental measurements see
Tables, pp. 351, 353.
Specimens examined. — Four, all from Crete.
>5, ?. Lassethe, 2,820 ft. Crete. Miss D. Bate (c). 5.12.2.16-17.
(5. 12. 2. 17. Type of species.)
skull. Katharo, Crete. Miss D. Bate (c). 5. 12. 2. 38.
?. Crete. H. 0. Jones, R.N. (c). 99.6.13.1.
Sub-Family LUTEINS.
1857. Lutiin;v Baird, Mamm. North Amer., p. 148.
Geographical distribution. — Nearly cosmopolitan ; absent only
in the Antarctic and high Arctic regions, Madagascar, Australia
and the Pacific Islands.
Characters. — Teeth of the same general type as in the
Melinse ; skull much flattened, and rostrum so shortened that its
length is less than its width ; external form long and slender,
the fur very dense, the legs unusually short ; toes webbed, the
claws short or absent ; tail long and highly muscular.
BemarJcs. — The family Lutrinse., the members of which appear
to be essentially badgers modified for semi-aquatic life, contains
four genera, one of which occurs throughout Europe from
Ireland eastward.
Genus LUTRA Brisson.
1762. Ltitra Brisson, Eegnum Animale in Classis ix distrib., 2nd ed., p. 13
{Lutra Brisson = Mustcla Intra Linnseus).
1780. Liitra Briinnich, Zoologiae Fundamenta, p. 34. First use of name
by an author following the Linntean system (Mustcla liiira).
1806. Lutris Dumeril, Zoologie Analytique, p. 12 (Modification of Lutra).
1817. Lutrix Rafinesque, Analyse de la Nature, p. 59 (Substitute for Lutra).
1843. Lontra Gray, Ann. and Mag. Nat. Hist., xi, p. 118, February, 1843
{Lutra canadensis Schreber).
1843. Latax Gray, Ann. and Mag. Nat. Hist., xi, p. 119, February, 1843
{Lutra lataxina F. Cuvier). Not Latax Glager, 1827.
1843. Lataxina Gray, List Spec. Mamm. Brit. jNIus., p. 70 {L. mollis Gray
= Lutra lataxina F. Cuvier).
1857. Lutra Blasius, Saugethiere Deutscblands, p. 236.
1865. Barangia Gray, Proc. Zool. Soc. Loudon, p. 123 {B. sumairana Gray
= Lutra barang F. Cuvier).
I
LUTRA 355
1865. Lutrogale Gray, Proc. Zool. Soc. London, p. 127 (Lutra monticola
Hodgson).
1865. Nutria Gray, Proc. Zool. Soc. Loudon, p. 128 {Lutra felina Molina).
1867. Lutronectes Gray, Proc. Zool. Soc. London, p. 180 {L. whiteleyi
Gray = L. lutra Linnseus?).
Type species. — Lutra * Brisson = Mudela lutra Linnaeus.
Geographical distribntion. — Same as that of the sub-family.
Characters. — Skull broad and low (depth of brain-case only
about half mastoid breadth), the entire dorsal profile nearly
straight, the zygomatic arches so widely expanded and post-
orbital region so much narrowed that distance from point of
greatest narrowing to zygoma is greater than postorbital con-
striction ; rostrum so shortened that posterior border of narial
opening approaches level of anterior zygomatic root, and distance
from orbit to gnathion is much less than half length of brain-
case ; auditory bulla flattened, with conspicuous meatal tube ;
paroccipital process low and ridge-like but distinct from bulla ;
dental formula : i ||, c ^ , pm ^ , m ^ = 36 ; teeth large as com-
pared with width of rostrum and palate (greatest transverse
diameter of upper carnassial equal to half distance between
carnassials) ; all of the small premolars opposed and capable of
trenchant action with those of opposite jaw ; upper carnassial
with crown triangular in outline, the outer side occupied by a
moderately high cutting edge formed by two cusps and a con-
necting ridge, the inner side (about half the total crown area)
by a flat crushing surface ; upper molar rhombic in outline, its
crown area about equal to that of carnassial (usually somewhat
smaller), its greatest diameter transvei'so to axis of tooth-row,
the four primitive cusps present ; lower carnassial with three
anterior cUsps well developed, sub-equal, the posterior heel
slightly larger than anterior triangle ; external form highly
modified for aquatic life, the body long and of approximately the
same width as neck and head, the tail long, very muscular, broad
at base, tapering distally, the legs short, feet broad, toes webbed,
short-clawed, the fur soft, dense and impervious to water.
Remarlcs. — Although more widely distributed than any other
living genus of land mammals, Eptesicus and Mi/otis perhaps
excepted, Lutra is not rich in species. A dozen or fifteen forms
are currently recognized, only one of which occurs in Europe.
LUTRA LUTRA Linnaius.
1758. [Mustela] Intra LinniEUS, Syst. Nat., i, 10th ed., p. 45 (Sweden).
1777. [Lutra'] vulgaris Erxleben, Syst. Regui Anim.,i, p. 448 (Renaming of
lutra).
1792. M[ustela] Lutra piscatoria Kerr, Anim. Kingd., p. 172 (Renaming of
lutra).
* Not " Lutra lutra" (see footnote under Meles, p. 841).
•2 A 2
356
CAHNIVOIJA
1816. ? Lutra fluviaiilis Leach, Syst. Catal. Spec. Inclig. Mamni. and
Birds Brit. Mus. , p. G (nomen nudum : '' River Otter ").
1827. [Lutra vulgaris] (8 marinus Billberg, Synopsis Faunas Scandinavia,
p. 28 (Coasts of Scandinavia).
1830. ?[Luti-a] fluviatilis Burnett, Quart. Journ. Sci. Lit. Art. xxviii,
1829, p. 349 (Substitute for lutra), nomen nudum.
1834. \_Lutra] nudipes Melchior, Den Danskc Stats og Norges Pattedyr,
p. 50 (Coast of northern Norway).
1834. Lutra roensis Ogilby, Proc. Zool. Soc. London, p. Ill (Roe Mills,
near Newton Lemavaddy, Londonderry, Ireland). Type in British
Museum.
1857. Lut7'a vulgaris Blasius, Siiugethiere Deutschlands, p. 237.
1884. ILzitra'] lutra Lataste, Actes Soc. Linn, de Bordeaux, xxxviii,p. 34.
1885. [Lutra] angustifrons Lataste, Actes Soc. Linn, de Bordeaux, xxxix,
p. 168, August, 1885 (Bone, Algeria). Perhaps in part only :
specimen from Liguria, Italy, referred to this form on p. 239,
September, 1885. Type in Lataste collection.
1910. Lutra lutra Trouessart, Paune Mamm. d'Europe, p. 86.
Type hiraUty. — Upsala, Sweden.
Geugrapliical distribution. — Europe and northern Africa, east-
ward into Asia ; limits of range not known. In Europe west to
Ireland and north to the Arctic coast.
Diagnosis. — Size medium (head and body in adult male about
700 mm., condylobasal length of skull, 105 to 123); tail about
three-quarters as long as head and body ; naked muzzle pad with
upper border strongly convex at middle ; skull much flattened,
the depth of brain-case not conspicuously more than lialf mastoid
breadth ; interorbital region narrow, its least width less than
distance from front of zygoma to anterior extremity of pre-
maxillary ; teeth not specially enlarged, the greatest diameter of
upper carnassial not greater than width of palate between
carnassials.
External characters. — General form long and slender, the
limbs very short, the feet liroad, with conspicuously webbed toes,
the head short flat and ill-defined from neck, the ears incon-
spicuous, the tail long, broad at base, tapering toward tip,
covered with the same short waterproof fur as body. Head
rounded and flattened, not well defined externally from the
muscular neck ; ear rounded, densely haired on both surfaces,
scarcely rising above level of fur, tlie antitragal lobe valve-like;
a second and third valve-like lobe above and behind meatus ;
muzzle short and wide, the nostril pad entirely naked, its
surface reticulate, its upper margin strongly convex at middle,
the lower slightly so, its lower border separated from mouth by
the densely haired upper lip, the width of which at middle is
about equal to height of pad ; whiskers stiff and bristly, the
longest extending about to ear when laid back. Legs short, feet
broad and rounded, with shoi't toes joined by a naked membrane
extending to base of terminal phalanges ; claws short but strong,
non-retractile, those on fore-feet best developed (about 8 mm. in
LUTRA 357
length) ; psilin bai-e, a Large, hejirt-shaped, obscurely ti'ilobed
tubercular mass behind base of digits, and a round posterior
tubercle separated from the larger pad by a deep groove ; sole
with a bare area and heart-shaped pad essentially as on pahu,
but with posterior tubercle represented by an ill-defined prolonga-
tion of the main pad ; posterior portion of sole densely furred.
Fur very dense and waterpi'oof, alike in texture throughout the
body and tail, the hairs of underfur 10 to 15 mm. in length, the
longer overlying hairs, which almost completely conceal the
underfur, about 25 mm. in length. Mamma? : 6* (probably i 3-3).
Colour. — Winter pelage : upper parts, legs, feet and tail a
rich dark brown (about the prout-brown of Ridgway or some-
FIG. 72.
Lutra Intra. Nat. size.
what darker), with a drabby cast more evident in some lights
than in others, the hairs with a conspicuous metallic gloss ;
underfur light grey, the extreme tips of its hairs changing
abruptly to prout-brown ; on underparts the drab becomes more
coiLspicuous as well as paler, usually assuming a tinge of cream-
l)ufi", the throat and cheeks fading to buffy white ; interramial
region and upper lip with irregulai- white mottlings, the hairs of
which are white to base ; whiskers and claws light horn-colour.
The exact colour is subject to considerable v;iriation, but the
material examined is not sufficient to show whether such
differences as occur are correlated with locality or season.
* Southwell, Field, rii, p. 1043, December 19, 1908.
358
CARNIVORA
Sometimes the brown is darker and richer than usual, or the
drab may be especially pronounced. Occasionally the long hairs
of the back ai'e a light dull buff, imparting to the animal a
peculiar faded appearance.
Skull. — General form of skull broad and flattened, more so
than in any other European carnivore, the depth of brain-case at
middle scarcely more than half mastoid width ; the brain-case exces-
sively narrowed anteriorly, Inroad postei'iorly and with greatly
Fig. 73.
Lufra Intra. Kat. size.
deveKiped lambduid crest, ])Ut low though evident sagittal crest ;
widely spreading zygomata forming a marked contrast with narrow
postorbital region ; r'ostrum short, deep and robust. Dorsal profile
nearly flat throughout, though region in front of middle of post-
orbital constriction slopes gradually downward. Owing to depth
of rostrum and shallowness of brain-case the dorsal and ventral
profiles are approximately parallel. Postorbital processes short
but evident, though not forming any considerable portion of border
of rather large, somewhat upturned orbit, the greatest diameter
LUTRA 359
of which nearly e(]uals width of flattened interorbital region.
Rostrum short, broad and deep, the distance from front of
zygoma to gnathion barelj^ equal to greatest breadth across
canines, about equal to depth at front of orbit, and contained
about 4^ times in condylobasal length of skull. Zygoma
widely spreading throughout, the arch heavy, not much bowed
upward, the anterior root conspicuously perforated by the large
anteorbital foramen the upper margin with low but evident
yiG. 74.
Lutra lutra. Nat. size.
angular projection marking posterior border of orbit. Palate
rather narrow, the distance between molars contained about
three times in that from gnathion to level of posterior border of
molar : incisive foramina moderate, about half as wide as long,
lying entirely between canines ; posterior extension of palate
about ecjual to distance between molars and about one-third that
from molar to hamular ; mesopterygoid space narrow anteriorly,
widei- posteriorly, its length about double the width between
haiiuilars. Auditory bulla small, flattened, though a little
3(i0
CARNIVORA
iiiHated along inner margin ; outline triangular-flask shaped, the
tubular portion of meatus not well defined from rest of bulla ;
least distance between bullse about equal to greatest diameter of
bulla including meatus. Paroccipital process low and ridge-
like, widely removed from inflated portion of bulla. Mandible
very robust, the ramus slightly bowed outward posteriorly, its
lower margin essentially straight except for a slight upward
curve behind level of tooth-row, the depth of ramus at middle
about one-third that through coronoid process ; angular process
very short ; coronoid process narrowly rounded oS" above, its height
above articular process about equal to width at articular level.
Teeth. — The teeth are large and strong, with well developed
cusps and commissures and relatively small crushing ai-eas, the
small premolars of upper and lower mandible titting closely
between each other when jaws are shut ; crown area of upper
molar about equal to that of car-
nassial, sometimes less. Upper
incisors forming a nearly straight
transverse row, the anterior faces
of the smaller teeth exactly in line,
those of the two larger teeth slightly
more posterior ; i^ and i~ small,
sub-equal, the shafts compressed,
the cutting edges rounded, simple ;
P higher than the others, somewhat
resembling the lower canine in form,
its apex curved outward and back-
ward beyond level of smaller teeth,
its posteroexternal surface with two
concavities separated by a trenchant
ridge ; lower incisors subterete, the
crown of v, obscurely and unsym-
metrically bilobed, that of the others
simple ; cross section of i-^ about one-third that of iy Canines
robust though not greatly elongated ; shaft simple, v\ithout
evident cingulum, that of upper tooth slightly curved backward,
that of lower tooth abruptly recurved, its axis set obliquely
outward. Anterior premolar both above and below well
developed and functional, the point of the upper tooth lying
internal to that of lower when jaws are closed ; pm^ small, single
rooted, its crown area about equal to that /^, its form es.sentially
like that of succeeding tooth, though with cusp relatively less
developed ; pm" and ptiv^ successively larger, two-rooted, the
crown area of imi^ fully three-fourths that of canine, each tooth
with a well-developed cusp lying in front of middle of crown,
and a distinct anterior and posterior cutting ridge ; cingulum
moderately developed ; outline of crown of pm'^ elliptical, that of
p)m^ ^with inner margin bulging inward behind middle, though
without developing a deflnite inner lobe ; pm., and pm^ much like
Flc. 75.
IjUtra Intra. Teeth.
LUTRA 361
pin- in l)oth size and form, the three teeth cutting a<j;ainst each
other when jaws are closed ; pm^ larger than the other small
premolars, its cusp l<3ss anterior in position, its crown decidedly
in wider posteriorly than anteriorly. Upper carnassial triangular
general outline, though this is somewhat obscured by the broadly
rounded inner portion and the somewhat projecting postero-
external lobe ; outer and posterior borders sub-equal and longest ;
trenchant portion of tooth with high anterior cusp connected
with well developed posterior cusp by a high, abruptly angled
commissure; crushing portion nearly as wide as long, its area
about as great as that of trenchant portion, its inner border
noticeably raised at middle though without secondary cusps ;
cingulum moderately developed along outer border of crown, and
forming a distinct though small antero-basal cusp. Lower
carnassial with anterior triangle well developed, the protoconid
and paiaconid sub-equal, the metaconid somewhat smaller ;
crushing portit)n of crown wider than antei'ior triangle though
not so long, the areas of the two portions of the tooth
approximately equal ; hypoconid evident though not high, the
outer surface of its base in line with that of protoconid, the
inner surface continuous with the concave though scarcely
basin-shaped main portion of crushing area ; cingulum narrow
but evident throughout, not specially developed in region
bordering outer base of hypoconid. Second lower molar flat,
subterete, slightly wider than long, the inner and outer margins
each with a small cusp somewhat behind middle, the cusps joined
by a low transverse ridge. Upper molar about equal to carnassial
in crown area, but length of its outer border decidedly less than
that of preceding tooth ; crown slightly constricted near middle,
the outer portion bilobed, the anterior lobe bearing a low paracone
and a broad outer jjrojection representing the parastvle, the
posterior lobe bearing a low but robust metacone ; iinier portion
of crown with a large protocone and low hypocone, the former
sending forward a conspicuous commissure to base of paracone,
the latter connected with cingulum that extends around base of
protocone.
Measitrements. — Adult male from Warwickshire, England :
head and body, 712 ; tail, 495 ; hind foot, 134 ; ear, 28. Adult
female from Csehtelek, Hungary : head and body, 640 ; tail, 380 ;
hind foot, 115; ear, 22. For cranial and dental measurements
see Tables, pp. 362, 363.
Specimens examined. — Thirty-three, from the following localities: —
Ireland : Londonderry, 1 (t}'pe of roensis) ; Ahascragh, Co. Galway, 1.
England : Pembrokeshire, 1 ; River Stour, Dorsetshire, 1 ; Rugby,
Warwickshire, 1 ; Norfolk, 1 ; no exact locality, 1.
Norway: Egersund, Stavanger, 5.
Sweden : Jockmock, Lappmark, 1 (Stockholm) ; Gnesta, Soderman-
land, 1 (Stockholm) : Skane, 1 (Stockholm) ; no exact locality, 2.
France : St. Gilles, Gard, 1 (Lataste) ; Etupcs, Doubs, 1 (IMotta/).
Germany : Southern Germany, 2.
362
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364
CARNIVOKA
Austria-Hungary : Karlsbad, Bohemia, 1 ; Csehtelek, Bihar, Hun-
gary. 1-
Italy: Torriglia, Liguria, 1 (Lataste).
Spain : Castrillo de la Eeina, Burgos, 3 (B.M. and U.S.N.M.) ; Galicia, 1 ;
Seville, 5.
Seiuarlcx. — Although the specimens show some rather note-
worthy variations in cohnir, and in cranial and dental measure-
ments, the matei'ial examined is not suthcient to indicate the
existence of geographical races in Europe ; while the north
African animal to which the name augustifrons has been applied
appears to be not separable from true Intra.
1.
Newton Lemavady,
Londonderry, Ireland.
{Miss A. Moody.)
6 st. Ahascragh, Galway.
3 juv. st. Pembrokeshire, Wales.
6. Rugby, Warwickshire,
England.
? St. R. Stour, Dorset.
1. England.
4 (i, 1 9, Egersund, Stavanger,
skulls. Norway.
ad., juv. South Germany.
6. Csehtelek, Bihar, Hun-
gary.
1. Karlsbad, Bohemia.
S, 9. Castrillo de la Reina,
Burgos, Spain.
1 juv. Galicia.
skull. Seville.
9, juv. Seville. {Dr. A. Ruiz.)
skull. Seville.
Zoological Society's 67. 12. 14. 4.
Museum. (Type of L. roensis
Ogilby).
Purchased (Row- 5. 7. 10. 1.
land Ward).
Purchased (Row- 6. 5. 21. 1-4.
land Ward).
Dr.T. S. Townsend 9. 12. 16. 1.
(c & p).
J. C. Mansel Pley- 98. 6. 13. 1.
dell (c & p).
K. H. Schaanning
(c).
Dr. A. Giiuther (c).
Hon. Mrs. N. C.
Rothschild (p).
Lord O. Russell
(c & p).
Rev. Saturio Gon-
zalez (c).
Dr. V. L. Seoaue
(c & p).
Lord Lilford (p).
Lord Lilford (i').
Col. L. H. Irby
(c & p).
99. A.
11.6.3.14-18.
59. 9. 6. 62-68.
10. 9. 14. 1.
57. 2. 14. 1.
8. 7. 7. 16-17.
94. 6. 18. 1.
74. 10. 7. 2.
76. 3. 4. 1.
95.9.4.10-11.
2. 11. 16. 1.
Sub-Family MUSTELINiE.
1835. Mustelimr Swainsou, Nat. Hist, and Classif. Quadr., p. vii.
(jrfO(jr<iphiral (Jidrlhntivn. — Northern hemisphere, south into
northern Africa, the Malay Archipelago, and northern and
western South America ; in Europe west to Ireland.
Character!^. — Dentition highly trenchant, upper carnassial
without crushing surface other than a small concave ai'ea between
inner lobe and main cusp, the crown much longer than broad,
not triangular or rhombic in outline, the posterior cusp compressed,
trenchant, barely half as high as main cusp ; upper molar much
reduced, the length of its outer portion one-third to one-half that
of carnassial ; small premolars alternating when jaws are closed,
at least one pair capable of shearing action ; skull varying in
MARTES 365
form but never greatly flattened and never very robust, the
rostrum always at least as long as broad ; external form slender,
the legs usually rather short ; feet digitigrade ; toes partly
webbed ; tail varying in length, slender or bushy, never con-
spicuously muscular.
Reinarhx. — As here understood the sub-family Mustelinse
contains the three genera, Maries, Miisfcln and VormeJa, all of
w Inch occui' in Europe.
Genus MARTES Pinel.
1792. Murtes Pinol, Actes Soc. d'Hist. Nat., Paris, i, p. 55 (M. domestica
Pinel = M. foina Erxleben).
1820. Martcs Nilsson, Skand. Fauna, i, p. 38 (M. foina and M. sylvatica =
martes).
1829. Zibellina Kaup, Entw.-Gescli. u. Natiirl. Syst. Europ. Tliierw., i,
p. 31 (M. zibellina).
1857. Mustela Blasius, Siiugethiere Deutsohlands, p. 211.
1911. Martes Thomas, Proc. Zool. Soc, London, p. 139, March, 1911.
Type species. — Mustela martes Linnaius.
Geographical distrihution. — Isorthern hemisphere from the
limits of tree growth south to the Mediterranean, the Malay
Archipelago, and the central United States ; in Europe west to
Ireland.
Characters. — Skull narrow, moderately high (depth of brain-
ca.se much more than half mastoid breadth), the dorsal profile
moderately curved, the zygomatic arches not specially wide-
spreading, and postorbital region not unusually narrowed (distance
between region of greatest narrowing and zygoma normally less
than breadth of postorbital constriction) ; rostrum narrow and
somewhat elongate, its width noticeably less than that of inter-
orbital region, the distance from anterior rim of orbit to gnathion
exceeding width of rostrum Ijetween anteorbital foramina; auditory
bulla3 moderately inflated, the meatal tube evident though .short,
the longitudinal diameter of bulla greatest ; paroccipital process
small, slightly projecting, partly distinct from bulla ; dental
formula : / — , c — , pm t*, in -* = 38 ; cutting edges of five small
;i-;r i-i' -^ 4-4' '2-2 ' o o
premolars (2 upper and 3 lower) capable of trenchant action ;
upper carnassial long and narrow, not triangular in outline and
without crushing surface, the small inner lobe standing as an
ofl'set to anterointernal extremity of crown, the sectorial portion
consisting of a high anterior and low posterior cusp with some-
what concave connecting ridge ; upper molar pyriform or pan-
durate in outline, its long axis nearly perpendicular to that of
tooth-row, its crown mainly fiat, but with a small paracone, still
smaller, sometimes obsolete metacone, and crescentic ridge-like
protocone ; lower carnassial wider posteriorly than anteriorly,
the anterior triangle much distorted, the metaconid reduced to
3G6
CARNIVORA
a well detined postero-internal process on base of protoconid, the
posterior crushing heel slightly more than half as large as
trenchant jiortion of tooth ; external form slender, somewhat
catlike or squirrel-like, the head moderately elongated, the
muzzle pointed, the ears high and conspicuous, the tail long,
bushy ; feet digitigrade, the moderately long claws partly
retractile ; fur long, dense and soft.
Hemnrks. — The genus Martes contains about ten species, some
of which are represented by numerous geographical races. Three
are known from westei'n and southern Europe.
KEY TO THE EUROPEAN FORMS OF MABTES.
Third upper premolar with crown strongly
convex on inner side, slightly concave on
outer side ; width of inner lobe of upper
carnassial nearly equal to that of tren-
chant portion of cro-wn ; greatest diameter
of upper molar about equal to length of
outer border of carnassial ; fur usually
finer and softer, and throat-patch more
yellow (Pine Martens) M. martes, p. 366.
Throat-patch cream-buff, general colour
darker (Central and northern Europe)... ,V. m. martes, p. 372.
Throat-patch buff-yellow, general colour
lighter (Mediterranean region) M. m. latinorum, p. 373.
Third upper premolar with crown evenly bicon-
vex ; width of inner lobe of upper car-
nassial barely half that of trenchant
portion of crown ; greatest diameter of
upper molar noticeably less than length
of outer border of carnassial ; fur usually
coarser and less soft, and throat-patch more
whitish (Beech Martens).
Condylobasal length of skull in adult male,
76 to 79 mm. ; pale throat patch always
much encroached on by brown of sur-
rounding parts, occasionally obliterated
(Crete) ! M. bunites, p. 380.
Condylobasal length of skull in adult male,
7'J to 84 mm. ; pale throat-patch seldom
much encroached on by brown of sur-
rounding parts and never obliterated
(Central and southern Continental
Europe) M. foina, p. 374.
General hue of upper parts drab iV. /. foina, p. 375.
General hue of upper ]5arts wood-brown .. M.f. mediterranea, p. 380.
MARTES MARTES Linna>US.
(Sj'uonymy under subspecies.)
Geographical distribution. — Entire wooded region of Europe,
from Ireland eastward into Asia, and from the Mediterranean
coast and islands northward to the limits of tree growth.
Diagnosis. — Third upper jjremolar with outline of crown
MARTES 357
concave on outer side, irregularly convex on inner side, the rec^ion
ot greatest width behind middle ; upper carnassial with inner
projection robust, its diameter in line of tooth-row nearly or
quite equal to greatest width of trenchant portion of tooth behind
middle of crown ; upper molar large, its greatest diameter about
equal to outer length of carnassial, the metacone more than half
as large as paracone, the outer border of the crown usually
notched; general colour yellowish brovvn, the throat patch
Fig. 76.
Martes martes. Nat. size.
always distinctly tinged with yellow ; underfur moderately long
External charactcrs.—Fovm slender and graceful, suggesting
that of both cat and squirrel ; legs moderately long ; tail about half
head and body, densely furred and bushv, the hairs at tip about
one-third as long as vertebr*. Head pointed ; eyes of medium
size ; ear rising distinctly above fur, its height from crown about
^5 mm. the outline rounded but forming a slightly indicated
point above, the entire surface densely furred : muzzle pad
368
CARNIVORA
well defined, completely naked, communicating with upper lip
by narrow median line. Feet densely furred throughout in
winter, the pads bare in summer ; palm with deeply divided,
trilobed, heart-shaped tubercular mass at bases of median digits,
a small round pad at base of thumb, and a larger round pad at
outer posterior border near wrist ; sole with similar trilobed
mass, but elements more distinct, almost forming three tubercles :
small pad at base of hallux like that at base of thumb ; no
posterior pad. Fur very dense and soft, the hairs of undei'fur
about l'5 mm. long at middle of back in winter, shorter in
summer, the longer hairs reaching about 40 mm. Mamma? : 4.
Colour. — The colour varies considerably with season and
climate, but is usually characterized by rich yellowish brown and
blackish tints, the legs, tail and underparts darker than the
back, the head noticeably lighter than the body, the throat and
fore part of chest yellowish, and interramia, chin and muzzle
dusky. Claws light horn-colour.
Skull. — As compared with that of the European species of
Mustela the skull of Marfcs marten is characterized by a general
Fig.
Martes inartex
smoothness, lightness and lack of angularity. Dorsal profile
strongly convex posteriorly, flattish behind orbits, the rostrum
falling away at an angle of about 20' ; a slight concavity usually
present near middle of nasals. Brain-case elongate-ovate in
outline when viewed from above, nearly as deep as wide, the
MARTES 369
sagittal crest moderately developed in old individuals and slightly
overhanging occiput. Floor of brain-case with no special
features ; a slightly developed median ridge. Auditory bulla?
moderately inflated, the surface smooth, the general outline
broadly flask-shaped (longitudinal diameter greate.^t), the meatal
tube well defined, i-ather wider than long ; space between buUie
about equal to diameter of bulla without tubular portion.
Postorbital region rather abruptly constricted, much as in
Mustda, its width about half that of brain-case. Interorbital
I'IG. 7S.
Marten martes. Nat. size.
region wider than rostrum, the discrepancy in breadth greater
than in Mustda crminen. Postorbital processes well developed
though short. Rostrum relatively longer than in 3Iustda
erminca or M. ptitoriits, the distance from orbit to gnathion about
equal to width between outer margins of anteorbital foramina,
the width across canines much less than distance from foramen
to gnathion. Orifice of anteorbital foramen over anterior root of
carnassial, and under anterior border of rather large orbit.
Zygamata rather widely but gradually spreading, strongly bowed
2 b
370 CARNIYOKA
upward behind middle. Palate rather narrow, its width
between molars equal to about 1^ times greatest diameter of
molar, incisive foramina small, ovate, slightly oblique, at level of
front of canine, the minute median foramen slightly behind
middle ; posterior extension of palate broader than long, reaching
about half way from level of molar to hamular ; mesopterygoid
space about a quarter longer than broad, the hamulars short,
sHghtly everted.
Teeth. — In proportion to size of skull the teeth are larger
than in the members of the other genera of MusteHme occurring
in Europe ; transverse diameter of upper molar much more than
half width of palate between molars. Upper incisors forming a
straight row separated at each side from canine by a space about
equal to transverse diameter of i^ and P together ; crowns strongly
compressed, the antero-posterior diameter fully twice transverse
diameter, the anterior face convex, the posterior concave with
faintly indicated cingulum ; ?'- slightly
larger than ?'^ i^ abruptly much larger
than the other two teeth together, its
crown more than one-third as high as
that of canine and with the posterior
concavity extending to outer basal
25ortion. Mandibular incisors smaller
and lower than the upper teeth, their
crowns projecting obliquely forward,
the root of r.^ implanted in jaw behind
level of the other two ; crowns ob-
scurely bitid, the outer lobe smaller
than the inner ; crown area of i^
pj(, -g about one-third that of the com-
Martcs martes. Teeth. Nat. size, pressed 'i.^ ov the subterete /,. Canines
with no special peculiarities ; diameter
of the upper tooth along alveolus equal to one-half width of
jialate between canines, that of lower somewhat more ; shaft
of upper tooth with posterior longitudinal ridge, and an antero-
internal ridge which near base curves backward across inner
side of shaft nearly to base of posterior ridge ; cingulum barely
indicated ; shaft of lower canine directed a little forward at
base, a little backward beyond middle, the anterior profile
strongly convex, the posterior equally concave ; surface of
enamel, e.specially on basal half of tooth, much roughened by
longitudinal wrinkles. Anterior premolar both above and below
single-rooted, small (crown area about equal to that of i^ and i'-'
together), subterete, the height of the single ill-developed cu.sp
scarcely ecjual to diameter of crown in p7)i^, much less in pm.i.
Other premolars, except upper carnassial, two-rooted, compressed,
the outline of crown triangular when viewed from the side, with
apex slightly in front of middle of tooth. Crown area of pw*''^
about three times that oi^mi^ ; outline of crown flattened-elliptical
MAllTES 371
or slightly concavo-convex, tlio concavity, when present, on outer
side; cusp simple, its height a little less than length of crown
along hase, its posterior surface with slightly developed longi-
tudinal ridge. Crown of j)Hi'^ somewhat longer than that of piii^,
but its area much greater owing to widening of strongly convex
inner side ; cusp essentially as in pni^ though somewhat higher.
Second lower premolar about equal in size to jyin-, but axis of
shaft more anterior in position ; pm.-^ essentially like pm^, but
cusp with faintly indicated anterior longitudinal ridge, and
posterior ritlge with a slight nodule or rudimentary basal cusp
(sometimes absent) ; pm^ noticeably larger than pm-^ and with a
well developed secondary cusp at middle of posterior border of
main cusp. Upper carnassial ( pm*^) long and narrow, the width
of crown just behind internal lobe less than half length along
outer border, the internal lobe robust, its diameter in line of
tooth-row nearly or quite equal to width of trenchant portion of
crown, its axis nearly perpendicular to main axis of tooth, its cusp
well developed, about as high as that of pm^, and separated from
main cusi? by deep concavity ; main cusp robust, its height
contained about 1§ times in length of crown, its axis slanting
a little backward, its anterior border with well developed
longitudinal ridge ; posterior cusp about half as high as main
cusp, its outer surface sloping obliquely to well developed
cingulum, the two cusps connected by a high, sharply trenchant
and obtusely angled commissure. Lower carnassial with crown
nearly 2^ times as long as broad, the sectorial portion of the
tooth consisting of a much distorted triangle, the paraconid
forming anterior extremity of tooth, its anterior border nearly
perpendicular, its posterior commissure meeting anterior commis-
sure of the higher protoconid at an abrupt angle, the two together
acting in opposition to the angled commissure of upper carnassial ;
metaconid reduced to a subterete postero-internal process on
base of protoconid, its area scarcely one-sixth that of larger
cusp ; crushing portion of crown slightly broader than trenchant
portion, its outline sub-circular with slightly raised edge, this
edge forming a noticeable though low posteroexternal cusp
separated from base of protoconid by a shallow but distinctly
angled notch. Second lower molar about equal to heel of
carnassial in size and essentially like it in form, the crown flat,
with faintly indicated outer ridge and low postero-internal cusp.
Upper molar large, its crown area at least equal to that of
carnassial, its greatest diameter about equal to outer length
of carnassial (see measurements, p. 378), the diameter of inner
portion of crown usually much greater than that of outer
portion, the median constriction well marked ; outer portion of
crown with two small cusps, probably the ]iaraoone and metacone,
tlie latter more than half as large as former, the outer margin of
tooth often though not invariably notched between them ; inner
portion of crown with a slightly curved, ridge-like antero-
2 B 2
372
CARNIVORA
external cusp (usually notched or partly divided into two), the
surface elsewhere finely wrinkled ; in some specimens a minute
tubercle, perhaps representing the hypocone, occurs near posterior
border of crown in region of constriction between outer and
inner portions ; cingulum low but evident, especially around
inner margin of crown.
Hemarks. — The well - known pine marten is so strongly
characterized as to require no special comparisons with any other
European species except Maries foina (see account of latter).
Two geographical races are currently recognized, though their
status is by no means clear.
Martes martes martk.s Linn;eus.
1758. [Mtistcla'] martes Linnaeus, Syst. Nat., i, lUth ed., p. 46 (Sweden).
1816. Miustda] sylvestris Oken. Lchrb. d. Naturgescli., iii, pt. 2, p. 1029
(Renaming of martes).
1827. M[artcs] vulgaris Griffith, Cuvicr's Anim. Kiugd., v, p. 123
(Renaming of martes).
1847. Martes sylvatica Nilsson, Skand. Fauna, i, 2nd ed., p. 41 (Renaming
of martes).
1857. Mustela martes Blasius, Siiugetliiere Deutschlands, p. 213.
I'JIO. Mustela martes Trouessart, Faune Manim. d'Europe, p. 72.
1911. [Maries'] martes Thomas, Proc. Zool. Soc. London, p. 139, March,
1911.
Tijpe locality. — Vicinity of Upsala, Sweden.
Oeogrn-pliical distribution. — Europe north of the Mediterranean
region, from Ireland eastward into Asia.
Diagnosis. — Throat patch cream-bufF or slightly more yellow.
Culoar. — Upper parts a rich dark brown, usually rather near
the bister of Ridgway, the tips of the longer hairs blackish ;
underfur light grey, the tips of the hairs tinged with drab or
with wood-brown ; face essentially like back or not so dark, the
muzzle and chin usually not darker than forehead ; ear edged
with bufly drab ; tail very dark brown (nearly the seal-brown of
Ridgway), its underfur raw-umber or somewliat darker ; feet
and lower portion of legs blackish ; throat-patch varying
considerably, but as a rule rather pale, nearly cream-buif or
slightly more yellow.
Measurements. — For cranial and dental measurements see
Tables, pp. 376, 378.
Specimens examined. — Twenty, from the following localities : —
Ireland : Kenmare, Kerry, 1 ; Go. Kerry, no exact locality, 1.
Scotland : No exact locality, 1.
England: Keswick, Cmnberland, 1; Cockerniouth, Cumberland, 1;
Pontrilas, Herefordshire, 1 ; Ludford Park, Herefordshire, 1 ; no exact
locality, 3 (B.M. and U.S.N.M.).
Norway : Egersund, Stavanger, 3.
Sweden : No exact locality, 3 (U.S.N.M.).
Denmark : No exact locality, 1 (U.S.N.M.).
Germany : Southern Germany, 2.
Austria-Hungary : Sohemia, 1.
MARTES '"^TS
9. Kenmaro, Kerry, Ire- E. Dodson (c). 94. 3. 27. 1.
land.
1. Kerrv. J. M. Gurney (p). 72. 1. 10. 1.
1. Scotland. (McLcaij.) Hargitt Collectiou. 8G. 9. 9. 1.
St. Keswick, Cumberland, W. R. Wilson (c). 96. 11. 6. 1.
England.
6. Cockermoutli, Cum- W. R. Wilson (c). 97. 1. 15. 1.
berland.
5. Pontrilas, Hereford- Mrs. St. John A. 85. G. 10. 1.
shire. Matthews (p).
6. Ludford Park, Here- H. J. Bailey (c & p). 0.2.23.1.
fordshire.
2. Wales (1828). Earl Cawdor (p). Gl. c. d.
2(5,19, Egersund, Stavanger, K. H. Schaanning 11.6.3.19-21.
skulls. Norway. (c).
2 skulls. Southern Germany. Dr. A. Giinther (c). 188i, 1299f.
6. Bohemia. Lord 0. Russell 50. 12. 23. 1.
(c & p).
Martes MARTES LATiNORUM Barrett-Hamilton.
1904. Mitstela martes latinorum Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xiii, p. 889, May, 1904 (Nurri Mountains, Sardinia).
1910. Mustela martes latinorum Trouessart, Faune Mamm. d'Europe,
p. 72.
Tyjye locallfi/. — Nurri Mountains, Sardinia.
Geograi)liical distrihntkm. — Mediterranean region (Italy, Sar-
dinia, Balearic Lslands).
Diagnonig. — Throat j^atch buft-yellow ; general colour lighter
than in true martes.
Colour. — Upper parts a rather light brown, the longer hairs
between the raw-umber and mars-brown of Ridgway, the
general effect darker in certain lights, e.specially on posterior
half of body ; underfur a pale bluish grey, essentially as in true
martes, the tijas of the hairs ranging from pale wood-brown to
clear butf; face Isabella-colour streaked with whitish hairs, the
muzzle and lips between mars-brown and j^i'out-brown ; ear
edged with bufly drab ; tail like back at base, darker at tip ;
feet and legs washed with dark brown ; throat-patch varying
considerably, but as a rule approaching buff-yellow.
Measurements. — Adult male from Porlezza, Como, Italy :
liead and body, 470; tail, 235; hind foot, 94:'6; ear from
meatus, 42. Adult female from San Cristobal, Minorca, Balearic
Islands : head and body, 430 ; tail, 230 ; hind foot, 87 ; ear
from meatus, 45. For cranial and dental measurements see
Tables, pp. 376, 378.
Specimens examined. — Nineteen, from the following localities : —
Italy : Porlezza, Como, 11 (Ghidini) ; near Rome, 4 ; Milan, 1.
Sakdinia : Nurri Jlountaius, 1 (type).
Spain : Maucor, Majorca, Balearic Islands, 1 ; San Cristobal, Minorca, 1.
Itemarlcs. — The status of this form is very unsatisfactory. So
fai- as can be judged from flu- few specimens examined there is
374
an average difference, however, between the Mediterranean
animal and that occurring north of the Alps. Two of the
specimens from Porlezza are darker than the others, in this
respect agreeing with the northern animal.
1. Italy. (C. Coli.)
1. Milan.
1. iSTurri Mountains, Sar-
dinia.
9. JIancor, ilajorca, Bale-
aric Islands. (Riutoii.)
9. San Cristobal, Minorca.
G. Barrett-Hamilton
E. Caveudish Taylor
(P)-
E. N. Buxton (p).
{Type of subspecies.)
0. Thomas (p). 1. 3. G. 1
11. 1. 2. 29.
5. 6. 6. 8.
95. 4. 16. 1.
0. Thomas & R. I. 0. 7. 1. 43.
Pocock (c & p).
MARTES FOINA Erxlehen.
(Synonymy under subspecies.)
Geograpliical distrihuiion. — Central and southern Continental
Europe, from the Atlantic coast eastward, and from the
Mediterranean to the Baltic.
Dkt(jn<isls. — Third upper premolar with outline of crown
nearly biconvex, the outer side occasionally
flattened, the gieatest transverse diameter
at middle ; upper carnassial with inner
lobe slender, its diameter in line of tooth-
row equal to only about half greatest
width of trenchant poi'tion of tooth behind
middle of crown ; upper molar not so large
as in M. martes, its greatest diameter less
than outer length of carnassial, the meta-
cone less than half as large as paracone ;
external form as in M. marten, but fur
of less fine quality ; colour usually more
greyish or drab than in 31. martes and seldom with the rich
brown tints of the related species, the throat-jjatch never strongly
tinged with yellow. Mammte : i.
Skull. — The skull resembles that of Marion martes, but may
usually be distinguished by its greater breadth and less depth.
Brain-case noticeably wider than high, the general outline when
viewed from behind nearly as in Mimtela erminea and distinctly
less elevated than in Martes inartes. Interorbital region and
rostrum wider than in Martes martes, and concavity ()f dorsal
profile in nasal region much more pronounced. Anteorbital
foramen usually smaller than in the related species, though in
the same position. In other respects the skulls of the two
animals show no tangible diflerences.
Teeth. — Except as already pointed out the teeth agree with
those of Martes martes.
KiG. so.
Lfirgei' cheek-teeth of Marte-i
7nartes (re) and M.foina {h).
JS'at. size.
MAETES 375
BemnrJiS. — Though readily distinguishable from Martes marles
by tlio character of the skull and teeth iff. foina is sometimes
difficult to recognize by external peculiarities alone. Usually
the colour has a slaty or drab cast that is highly characteristic,
and the <|uality of the fur is inferior to that of the pine marten ;
but I have seen tanned skins of animals killed in winter which
were impossible to identify with any degree of certainty.
Two geographical races have. been described.
Martes foina foina Erxleben.
1777. {Mustela] foina Erxleben, Syst. Regni Anim., i, p. 458 (Germany).
1792. Martes domestica Pinel, Actes Soc. d'Hist. Nat., Paris, i, p. 55 (France).
1801. M[;ustela] f[oina] alba Bechsteiu, Gemeinu. Naturgesch. Deutsch-
lauds, I, 2nd ed., p. 759 (Thiiringen, Germany).
1857. Mustcla foina Blasius, Siiugethiere Deutschlands, p. 217.
18G9. Mustcla martes var. fagorum Fatio, Famie Vert. Suisse, i, p. 318
(Sweden) : name wrongly attributed to Linnaeus, Syst. Nat., i,
12th ed., p. ()7.
1910. Mustcla foina Trouessart, Faune Mamm. d'Europe, p. 72.
Type locality. — Germany.
Geographical distrihution. — Range of the species except
southern Spain.
Diagnosis. — Longer hairs of back tipped with sepia, the
general hue of upper parts drab.
Colour. — Underfur very light grey (about grey No. 10 of
Ridgway) with an evident buffy cast on distal third of hairs;
long hairs ranging from wood-brown to mars-brown, becoming
darker at tips. As the long hairs nowhere conceal the underfur
the general effect is a light drab resulting from the blending of
the two colours, very uniform thi'oughout dorsal surface, though
usually a little paler on neck and darker on posterior portion of
back. Tail essentially like body, but darkening toward tip, the
pencil often blackish. Legs and feet washed with dark brown.
Ear huSy drab externally, creamy white along rim, the inner
surface greyish white. Muzzle, lips and chin bi'occoli-brown.
Throat-patch buffy white, darkening to cream-buff' along outer
edges.
Measurements. — Adult male from near St. Gallen, Switzer-
land : head and bodj?^, 453 ; tail, 260 ; hind foot, 85 ; ear from
meatus, 3-1. For cranial measurements see Table, p. 377.
Specimens examined. — Twenty-three, from the following localities : —
France : ]\Iauonville, Mourthe-et-Moselle, 2.
Germany: Ingelheini, Rheinhessen, 2; Nuremberg, Bavaria, 1
(U.S.N.M.) ; southern Germany, 7 (skulls).
Switzerland : Geneva, 1 (Mottaz) ; Vallee-de-Joux, Vaud, 1 (Mottaz) ;
Thavngeu, SchaShauseu, 3 (B.M. and U.S.N.M.) ; Aarburg, Aarau, 1
(U.S.N.M.); Oberrich, St. Gallon, 1.
Spain : Silos, Burgos, 1 ; Donasantos, Burgos, 1 (U.S.N.!M.).
Italy : Near Rome, 1.
376
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380
CARNIVORA
Greece : Mt. Vuno, Cophalonia, 1 (colour normal ; size apparently
rather small).
?, juv. IManonviUe, Meiirthe-et- Lord Lilford (p). 95. 9. 5. 1-2.
Moselle, France.
6. Ingelbeim, Eheinhessen, C. Hilgert (c). 8. 11. 2. 17.
Germany.
6. Ingelheim, Eheinhessen. G. Barrett-Hamilton 11. 1. 2. 102.
7 skulls. South Germany. Dr. A. Giinther (c). 1299, a-g.
2 S juv. Schaffhausen, Switzerland. 0. Thomas (p). 2. 8. 4. 22-23.
(E. H. Zollikofcr.)
6. Oberrich, St. Gallen. 0. Thomas (p). 2. 8. 4. 21.
{E. H. Zollikofer.)
S. Silos, Burgos, Spain. Rev. Saturio Gonza- 8. 7. 7. 21.
lez (c).
1. Rome, Italy. B. Cavendish Taylor 5. 5. C. 7.
1. Mt. Vuno, Cephalonia, J. I. S. Whitakcr (p). 8.10.1.9.
Greece. (C. Mottaz.)
Martes foina mediterranea Barrett-Hamilton.
1898. Mnsfcla mediterranea Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., i, p. 442, June, 1898.
1910. Mustela foina mediterranea Trouessart, Faune Mamm. d'Europe,
p. 73.
Type locality. — Sierra de Jerez, Cadiz, Spain.
Geographical clistribution. — Southern Spain.
Characters. — Longer hair.s of back tipped with a light
yellowish brown, the general hue of upper parts lighter, yellower
and less drab than in true/o/)(«.
Measurements. — For cranial and dental measurements see
Tables, pp. 377, 379.
Specimens examined. — Two, the type, from the province of Cadiz, and
a second specimen (mummy) from the Sierra Nevada.
BemarTcs. — If the two individuals of Martes foina mediterranea
are not abnormal in colour the race is rather well characterized.
I have seen only one skin of true foina that ajaproaclies them, a
specimen from Rome.
1 juv. Sierra de Jerez, Cadiz, Spain. A Chapman (c & p). 98. 3. 18. 1.
(Type of subspecies.)
MARTES BUNITES Bate.
1899. Mustela foina leucolaclin:ia Barrett-Hamilton, Ann. and IMag. Nat.
Hist., 7th ser., iv, p. 383 (Not of Blauford).
190G. Mustela foina bunites Bate, Proc. Zool. Soc. London, 1905, ii, p. 318,
April 5, 190G.
1910. Mustela foina bunites Trouessart, Faune ^Nlamm. d'Europe, p. 73.
Tyjje locality. — Kontopalo, Kania, Crete.
(reograjMcal (listrihution. — Crete.
Diagnosis. — In general like Martes foina, but size not so
MUSTELA 381
large (condylobasal length of skull in adult male not reaching
80 inm.), rostral portion of skull not so broad, and pale throat-
patch so much encroached on by brown of surrounding jiarts,
that it is occasionally absent.
Colour. — The colour is lighter and more yellowish (less drab)
than in Maries foina folna, as the long hairs continue a clear
wood-brown to tips, producing a general effect more nearly a
light isabella-colour than drab. The difference is, ho\\ever, not
very great. Tail and feet essentially as in M. f. foina, though
not so dark. Throat-patch greatly reduced, absent in one of the
se^■en skins,* and in its most extreme development represented
by a mere horseshoe-shaped mark, one extremity of which lies in
front of each fore leg, while the anterior portion broadens out
to form an irregular patch covering entire throat immediately
behind interrainial region.
Slodl and teeth. — The skull and teeth resemble those of
Martrs foina except for their uniformly smaller size and the
relatively somewhat less breadth of the rostrum.
Meamrrmcnts. — External measurements of tyjae (young-adult
male) : head and body, 403 ; tail, 255 ; hind foot, 79 ; ear from
meatus, 39. For cranial and dental measurements see Tables,
l^p. 377, 379.
Specimens examined. — Seven, all from Crete.
Miss D. Bate (c). 5. 12. 2. 18-20.
Miss D. Bate (c). 5. 12. 2. 21.
Miss D. Bate (c). 5. 12. 2. 22.
(Type of species.)
Miss D. Bate (c). 5. 12. 2. 37.
H. 0. Jones, K.N. (c). 99. 6. 13. 2-3.
Genus MUSTELA Linnreus.
(Sj-nonymy under sub-genera.)
Type species. — Mustela erminea Linnaeus.
Geoijrapliiral distribution. — Northern hemisphere fi'om the
Arctic coast south in the Old World to northern Africa and the
Malay Archipelago, and in America to the Andes ; in Europe
east to Ireland.
Characters. — 8kull in general resemljling that of Martes, but
rostrum so shortened that distance from orbit to gnathion is less
than width of rostrum between anteorbital foramina ; auditory
bulla without meatal tube, its outline variable but never Hask-
shaped ; paroccipital process small and flattened, closely applied to
posterior martrin of bulla ; dental formula : i 1"-, c — , mm —
* .\ small tuft of white hair persists behind each angle of mouth, and
another in front of each fore leg, the largest of .these about 15 mm. in
diameter.
6,2.
1.
i.
Kanea, Crete
Katharo.
Kontopalo.
skull.
2.
Crete.
Crete.
382 CAENIVORA
m ^1 = 34 ; upper carnassial as in Maries, its posterior (;usp low
but well developed, the height of its main cusp about half outer
border of crown ; upper molar between pyriform and pandurate in
outline, the constriction evident though not deep, the main axis of
the crown nearly perpendicular to sagittal line ; lower carnassial
without metaconid, the posterior lieel crossed by a longitudinal
trenchant ridge ; other teeth essentially as in Maries ; external
form slender, the muzzle obtuse, the ears low and rounded, the
legs short, the tail variable in length and in quality of hair, but
never so bushy as in Martes.
Bemarks. — The genus Mustela, the most widely distributed of
the strictly terrestrial Mustelinse, is also the richest in species.
These fall into three main groups, extreme members of which
might be regarded as generically distinct. The existence of
species with intermediate characters makes it impossible to
define these groups by anything more than their average
differences.* They are therefore here treated as sub-genera. All
three are represented in Europe.
KEY TO THE EUROPEAN FORMS OP MUSTELA.
Brain-case broad, the mastoid width decidedly
greater than distance between basion and
palation ; auditory bullae triangular in outline
(Sub-genus Pntorius, Polecats) M. putorms, p. 419.
Underfur usually buffy grey (Central Europe) M. p. putoHus^ p. 423.
Underfur usually yellowish (Southern Spain) il.p- aureolus^ p. 425.
Brain-case narrow, the mastoid width less than
or about equal to distance between basion and
palation ; auditory buUfe not triangular in
outline.
Rostrum flattened above; inner margins of
auditory bulte strongly divergent pos-
teriorly ; tail bushy ; habits semi-aquatic
(Sub-genus Lutreola, 'Minkii) M. lutnola, p. 415.
Rostrum convex above; inner margins of auditory
bullffi nearly parallel ; tail not bushy ; habits
terrestrial (Sub-genus J\lustda, Stoats and
Weasels).
Tail with conspicuous black terminal area in-
cluding more than pencil ; skull with ros-
trum usually not so wide as interorbital
region; condylobasal length of skull never
less than 43 -4 mm. in adult male, or than
35 '4 mm. in adult female; hind foot
usually more than 43 mm. in males and
34 mm. in females (Stoats).
Brown of sides noticeably encroaching on
yellowish or whitish of underparts, the
line of demarcation irregular ; upper lip
always brown M. Jiibcniica, ]). 398.
* At least so far as Lidrcola and true Mustela are concerned. Putorms
appears to be sharply circumscribed, so that it might readily be regarded
as a distinct genus (see remarks on p. 419).
MDSTELA 383
Brown of sides not encroaching on yellowish
or whitish of undcrparts, the line of
demarcation straight ; upper lip wholly
or partly white.. m. crminca, p. 385.
Kegion extending from front of rostrum to
and including postorbital processes
relatively broad and short ; size below
the maximum for the European races ;
condylobasal length of skull in adult
males, 43-4 to 48-6 mm., mandible in
adult males, 23 to 26-4 mm. (Scandi-
navin Peninsula, except extreme south
of Sweden) .,/. ,. ,,„„■ gg^
Kegion extondmg from front of rostrum
to and including postorbital processes
relatively narrow and long ; size maxi-
mum for the European races ; condy-
lobasal length of skull in adult males,
47 to 52-5 mm., mandible in adult
males, 25 to 30 mm.
Zygomatic arches broadly spreading
(ratio of zygomatic breadth to con-
dylobasal length about 59) ; posterior
extension of palate unusually short
and broad (Islands of Islay and Jura
Scotland) ,V. ,. ,,-,,-„^ 3,j7_
Zygomatic arches moderately spreading
(ratio of Z3'gomatic breadth to condy-
lobasal length about 54); posterior
extension of palate not unusually
short and broad.
White winter coat normally assumed ;
condylobasal length of skull in
adult male seldom over 50 mm. ;
in adult female seldom over
45 mm.; upper carnassial not
enlarged and thickened (Conti-
nental Europe from southern
,,r,--?^'^'^®" *° *^^ -"^'PS ^^^ Pyrenees) M. c. «jun, p 3S'J
White winter coat normally not
assumed ; condylobasal length of
skull in adult male often over
50 mm. ; in adult female often over
45 mm. ; upper carnassial enlarged
m •, •., , ^"tl tJiickened (Great Britain) il. c .tabilU^n Z2Q
iail without conspicuous black terminal area,
though a tuft of black or blackish hairs
may bo present in pencil ; skull with ros-
trum usually wider than interorlntal
region ; condylobasal length of skull, except
m specimens from the Mediterranean
region, often less than 40 mm. in males, and
34 mm. in females; hind foot -(except in
M. africana) usually less than 41 mm. in
males and 25 mm. in females (Weasels).
Size about as in the smaller stoats, condy-
lobasal length of skull in adult male,
48 to 50 mm., mandible in adult male
about 28 mm., its depth at front of car-
nassial, 4 • 5 to 5 mm. ; tail nearly one-half
as long as head and body (afiicanus
group).
384 CARNIVORA
Brown of sides noticeably encroaching on
yellowish or whitish of underparts,
the line of demarcation irregular
(Malta, Azores) M. afiicana, 412.
Brown of sides not encroaching on yellowish
or whitish of underparts, the line of
demarcation straight (Crete) .1/. fjalintliias, p. 414.
Size less than in the stoats, condylobasal
length of skull in adult male, 37 to
47 mm. ; mandible in adult male not
more than 24 mm., its depth at front
of carnassial less than 4-5 mm. (true
Weasels) M. nivalis, p 401.
Winter pelage often white ; size small, hind
foot of males, 29 to 34 mm. (Northern
and central Europe) M. a. nirallf, p. 402.
Winter pelage very rarely white ; size
medium or large, hind foot of males
usually 32 to 41 mm. (Mediterranean
region).
Skull of adult male not infrequently
44 to 4G'6 mm. in condylobasal
length ; hind foot of adult male 34 to
41 mm. ; colour rather dark (Central
Mediterranean region) M. n. boccamda, p. 405.
Skull of adult male seldom if ever exceed-
ing 44 mm. in condylobasal length ;
hind foot of adult male 30 to 37
mm. ; colour rather light (Iberian
Peninsula) M. n. iberka^ p. 407.
8ub-genus MUSTELA Linnanis.
1758. Mustcla Linnteus, Syst. Nat., i, 10th ed., p. 45 (Type by tautonymy
M. erminea LinniBUs).
1829. Ardocjalc Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,
p. 30 [erminea).
1829. Idis Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i, p. 35
(vulgaris = nivalis). Not of Schinz, 1824-1828.
1841. Gale Wagner, Schreber's Saugthiere, Suppl., ii, p. 234 {uiilgaris =
nivalis).
1857. Fcctorius Blasius, Saugethiere Deutschlands, p. 219 (part).
18G5. Ncogale Gray, Proc. Zool. Soc, London, p. 114 {brasiliensis, aureo-
vcntris and xanthogenys).
1871. Mustclina, Bogdanow, Tpvj. Oiiuu Eciei-TB. Kasaiici;. .ViiiiBppc. I, Mem. i,
p. 167.
1899. Enmustcla Acloque, Faune de France, MammifcTcs, p. G2 {vulgaris
and enninca).
1911. Mitstda Thomas, Proc. Zool. Soc. London, p. 138, April, 1911.
Tijpe species. — 3Instcla eniiiiica Linn;eus.
Gcograpli'tcal (listrihutum. — Northern liemisphere, south in the
Old World to northern Africa, and in the New World to the
Andes ; in Europe west to Ireland.
Charartcrs. — Form very slender ; tail not bushy ; fur not
modified for aquatic life ; skull without noticeably projecting
mastoid processes ; auditory bulhe not triangular, the inner and
outer borders nearly parallel ; posterior boi'der of jmr^ shearing
MUSTELA 385
against anterior border of jj)«3 ; other small premolars not
capable of trenchant action.
RrmarJcs. — The sub-genus Mustela is the most widely-
distributed group in the genus. It is also the richest in species
and local races. About seventy of these are now recognized,
eleven of which occur in Europe.
MUSTELA ERMINEA Linnseus.
(Synonymy under subspecies.)
GeograpTiical disfrihufion. — Europe from the Arctic coa.st
to the Pyrenees and Alps, and from Great Britain eastward
into Asia.
Diagnosis. — Skull with narrow brain-case and slightly pro-
jecting, scarcely angular mastoid region, the mastoid breadth
less than distance from basion to palation ; rostral width across
canines usually less than interorbital width ; auditory bulhe much
longer than broad, nearly parallel sided, rounded posteriorly,
truncate anteriorly ; form very slender, the tail slender and not
bushy ; colour brown above, whitish below, the line of demarca-
tion straight ; a white winter pelage assumed in colder parts of
range ; tail always with black tip including more than terminal
pencil ; upper lip always at least partly white ; ear usually with
a whitish rim.
External characters. — General form long and slender, the
legs short, the body cylindrical, the neck long and nearly as
thick as body, tail considerably longer than outstretched hind
leg. Ear short but appearing distinctly above fur of head, its
outline evenly rounded, both outer and inner surface densely
clothed with short hair ; muzzle rather broad and short, the
nostril pad sharply defined, entirely naked, separated from upper
lip by a narrow hairy area. Palm and sole completely furred in
winter, the tubercles bare in summer ; on both palm and sole
there is a trilobed, heart-shaped tubercular mass at base of
median digits and a small round pad at base of inner digit ; on
palm there is an additional posterior rounded pad near wrist.
Fur dense and soft, the longest hairs on back about 10 mm.
in winter, less in summer ; tail rather closely haired, the pencil
full, usually wider than basal portion of tail, its longest hairs
about half as long as vertebra3 ; fur turning white in winter
except in warmer parts of range. MammsB : i 4 — 4 = 8.
Colour. — Upper parts and outer surface of legs yellowish
brown, usually a little darker along middle of back and on head,
the underfur and bases of the longer hairs much paler ; under-
parts, inner surface of legs and upper surface of feet whitish or
yellowish in strong contrast, the line of demarcation between
2 c
386
CAKNIYORA
the two colours straight and definite, extending along middle of
sides of body and neck and passing a little below ear and eye to
lower edge of muzzle pad, normally leaving upper lip white ; ear
brown, the rim whitish ; no dark spot behind angle of mouth ;
tail with terminal tifth and entire pencil black, elsewhere
concolor with back except for an ill-defined yellowish or whitish
median area which occasionally extends along under surface from
base toward or to black tip. In white winter pelage the entire
animal is white or whitish (often tinged with yellow), with the
Fig. 81.
Muatda erminea. Nat. size.
excej)tion of the black area of the tail, which remains as in the
dark coat.
Skull. — In general appearance the skull resembles that of
MtisfcJa jmtorins (p. 421). This is especially true of the dorsal
profile, the form of the palate, rostrum and interorbital region, in
all of which the differences between the two animals are very slight.
Breadth of rostrum over canines rarely equal to least interorbital
width. Position of auteorbital foramen as in M. 2)i(tor{us, but
orifice relatively larger, the width of plate sej^arating it from
MUSTELA 387
orbit distinctly less than that of foramen. Postorbital process
short, not evidently directed backward. Zygomata more widely
spreading than in M. putorins, strongly and evenly bowed when
viewed from the side, the orbital process barely indicated and
posterior widening absent. Owing to shortness of postorbital
process and virtual absence of orbital process of zygoma, the
orbit is less margined with bone than in M.puturius. Brain-case
longer and narrower than in Mustela putorius, the outline when
viewed from above elongate ovate not distorted by the presence
of conspicuously projecting mastoid regions ; sagittal crest slight,
even iii old individuals ; lambdoid crest moderately developed,
scarcely overhanging, the condyles usually visible from above.
Floor of brain-case nearly flat, though with the usual median
ridge and lateral and posterior depressions present. Auditory
bulla? moderately inflated throughout, about three-quarters as
wide as long, nearly parallel sided, squarely truncate in front,
rounded behind, the meatus not tubular ; anterior extremity of
bulla not in contact with haraular, and separated from foramen
ovale by a broad nearly tiat area, the length of which is
about one-half distance between foramina ; inner extremity of
glenoid surface marked oft" by a wide, shallow notch. Postoi;bital
region abruptly constricted, short, without indication of the
neck-like elongation which forms so conspicuous a feature of
the skull of M. putorius. Mandible with lower border slightly
convex, its posterior fourth somewhat elevated above general
outline ; angular process small but better developed than in
M. putorius.
Teeth. — Except for their smaller size the teeth are essentially
like those of Mustela putorius. Enamel of lower canine not
rugose. Second upper premolar and third lower premolar more
compressed than in the larger animal, and with base of cusp
shorter, so that there is a slight but evident concavity or
flattening at both anterior and posterior base. Upper carnassial
with inner lobe projecting more forward than in M. putorius, so
that its anterior border extends nearly to level of that of outer
margin of tooth. Upper molar and posterior lower molar as in
Mustela putorius, but rii^ with, metacone more reduced, and m.,
with crown area distinctly smaller than that of heel of carnassial.
Mustela ermine a erminea Linnaeus.
1758. [Mustela] erminea Linnajus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1792. M^ustela] erminea hyhcrna Kerr, Anim. Kingd., p. 181 (name applied
to the northern true ermine).
1816. ]S[\jistcla] hcrminea Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1026
(Renaming of erminea).
1827. [Mustela. erminea'] macnlata Billberg, Synopsis Faunae Scandinavise,
p. 8 (Scandinavia).
1857. Fcetorius erminea Elasius, Siiugethiero Deutschlands, p. 228 (part).
2 c 2
388
CARNIVORA
1877. Putortus erniinea Goues, Fur-bearing Animals, p. 109 (part).
1910. Putorius (Ictis) ermincus Trouessart, Faune Mamm. d'Europe, p. 78
(part).
Ti/2)e locaUty. — Upsala, Sweden.
Geogra^jliical distribution. — Scandinavian Peninsula, except
extreme south of Sweden ; eastern limits of range unknown.
Characters. — Region extending from front of rostrum to
and including postorbital processes relatively broad and short;
size below maximum for the European races : condylobasal length
of skull in adult male, 43 "4 to 48 • 6 mm. ; mandible in adult
male, "23 to 26*4 mm. ; basal portion of tail (in summer pelage)
usually lighter below than above.
Measurements. — Hind foot in each of two males from the
vicinity of Christiania, Norway, 43 mm. ; in a female from the
same region, 35 mm. Male and female from Nordre Fron, Lower
Gudbrandsdal : hind foot, 43 and 34 mm. Two males from
Lappmark, Sweden : hind foot, 44 and 46 mm. Adult female
from Upsala, Sweden : hind foot, 36 mm. Adult male from
Stockholm, Sweden : hind foot, 44 "6. For cranial measurements
see Table, p. 392.*
Specimens examined. — Twenty-five, from the following localities : —
Sweden: Karesuando, Lappmark, 1 (U.S.N.M.) ; Wilhelmina, Lapp-
mark, 1 (U.S.N.M.) ; Jemtland, 1 (U.S.N.M.) ; Upsala, 1 (U.S.N.M.) ;
Stockholm, 2 (U.S.N.M. and Stockholm).
NoEWAY : Aker, near Christiania, 2 ; Brekkebygden, Trondhjem, 1 ;
Nordre Fron, lower Gudbrandsdal, 2; Marestuen, Fillefjeld, Bergen, 1,
Egersund, Stavanger, 13.
HemarTcs. — The Scandinavian form of Mustela erminea is
distinguishable from the other European races by its slightly
smaller size and by the broader rostro-frontal region of the skull,
characters which appear to be fairly constant. Of the other
races it perhaps most closely resembles M. erminea ricinae from
the islands of Islay and Jura, west Scotland. The white winter
coat is normally assumed throughout the range of this form.
c5, 6 juv. Aker, Christiania, Nor- Christiania Museum 93. 3. 1. 2, 4.
way. (e).
5. Brekkebygden, Trond- Christiania Museum 93. 3. 1. 3.
hjem. (e).
6, ?. Nordre Fron, Gudbrands- Christiania Museum 93. 3. 1. 5-6.
dal. (e).
?. Marestuen, Fillefjeld, E. K. Alston (p). 79. 9. 25. 21.
2,.50O ft.
13skulls. Egersund, Stavanger. K. H. Schaanning (c). 11.6.3.22-34
* In eight skuUs of males from. Egersund, Stavanger, Norway (all
sutures closed, teeth slightly or not worn), received after the Tables were in
type, the averages and extremes are as follows : condylobasal length, 44-2
(43 -4-45 -2); zygomatic breadth, 24-2 (;23-6-25-4) ; mandible, 23-8 (23-0-
24 '8). Two females from the same locality: condylobasal length, 35'4
and 38-0; zygomatic breadth, 18-6 and 20-0; mandible, 18-2 and 19-2.
389
MUSTELA ERMINEA iESTIVA Kerr.
1792. Mlustela] crminca .rstiva Kerr, Anim. Kingd., p. 181 (Germany;
based ou Schreber's pi. 137a).
1820. Mustela erminea major Nilsson, Skand. Fauna, i, p. 34 (Carlskrone,
Blekingc, Sweden).
1857. Fatorius erminea Blasius, Siiugethiere Deutscblands, p. 228 (part).
1910. PiitoriiiH (Ictis) ermineiis Trouessart, Faune ]\Iamm. d'Europe, p. 78
(part).
Type locality. — Germany.
Gcoijraphical distrihniion. — Continental Eui-ope from southern
Sweden to the Alps and Pyrenees.
Characters. — Region extending from front of rostrum to and
including postorbital processes relatively narrow and long ; size
larger than in M. erminea erminea ; condylobasal length of skull
in adult male, 47 to 51 mm. ; mandible in adult male, 25 to
29 mm.
Measurements. — Two young adult males from Pic du Midi,
Hautes- Pyrenees, France : head and body, 235 and 240 ; tail,
82 and 85 ; hind foot, 44 and 44 ; ear from meatus, 17 and 17.
Three males from Strass, near Burgheim, Bavaria : head and
body, 265, 270 and 280 ; tail, 90, 100 and 102 ; hind foot, 44,
47 and 48. Two females from the same locality : head and body,
240 and 240 ; tail, 80 and 90 ; hind foot, 38 and 40. Average
and extremes of eight adult males from the vicinity of
St. Gallen, Switzerland: head and body, 271 (251-292); tail,
104 (94-111); hind foot, 47 (45-50). Three adult females
from the same region : head and body, 226 (218-242) ; tail, 89
(85-98); hind foot, 38-8 (37-6-41).
Specimens examined. — Seventy-nine, from tbe following localities : —
Denmark : No exact locality, 2 (skulls, Copenhagen) ; Amager, 1
(skull, Copenhagen) ; Faaborg, Fyn, 4 (skulls, Copenhagen) ; Vestervig,
Jutland, 1 (skull, Copenhagen).
Holland: Texel Island, 1.
France : Pic du Midi, Hautes-Pyr6n6es, 2 ; Manonville, Meurthe-et-
Moselle, 1.
Germany: Neustadt, Wied, 1 ; Ingelheim, Bheinhessen, 8 ; near Magde-
burg, 2; Bleiche, Saxony, 5; Moritzburg, Saxony, 1 (U.S.N.M.) ; Rudol-
stadt, Thiiringen, 3 ; INIarxheim, Bavaria, 4 ; Strass, near Burgheim,
Bavaria, 5 ; no exact locality, south Germany, G (skulls).
Austria-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 1.
Switzerland : Frete-de-Sailles, Vaud, 1 (U.S.N.M.) ; Frutigen, Bern, 1
(U.S.N.M.); Rorbas, Bern, 1 (U.S.N.M.); Ziirich, 1 (U.S.N.M.); St.
Gallen, 4 (U.S.N.JI.) ; Degersheim, St. Gallen, 5 (U.S.N.M.) ; Rorschach,
St. Gallen, 1 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 ; Untervatz, Grisons, 3
(B.M. and U.S.N.M.) ; Priittigau, Grisons, 1 (U.S.N.M.) : Vals, Grisons, 1
(U.S.N.M.); Obersaxen, Grisons, 1 (U.S.N.M.); Pontresina, Grisons, 1
(U.S.N.M.) ; Scanfs, Grisons, 1 (U.S.N.M.) ; Poschiavo, Grisons, 1 (U.S.N.M.);
Ems, Grisons, 2; Osogua, Ticiuo, 1 (U.S.N.M.).
Hemarks. — The ermine of central Europe differs from that of
Scandinavia in sligiitly greater size and in the narrower rostro-
frontal region of skull. The white winter pelage is normally
assumed throughout the ranse of the form.
390 CARNIVOEA
As shown by both external and cranial measurements the two
specimens from the Pic du Midi are unusually small. It is
possible that they represent a peculiar Pyrenean race.
5. Texel Id., Holland. J. L. Bonhote (c & 8. 10. 26. 1.
P).
2 S. Pic du Midi, Hautes Pyre- O. Thomas (p). 8. 9. 1. 52-53.
n6es, France. (A. Bobert.)
6. Manonville, Meurthe - et - Lord Lilford (p). 11.1.1.139
Moselle. (Lomont.)
?. Untervatz,Grisous,Switzer- 0. Thomas (p). 4. 4. 5. 33.
land. {E.H.ZoUikofcr.)
6,9. Ems, Grisons. (£. Jf. ;?oZiJ- 0. Thomas (p). 2.8.4.25-26.
kofer.)
i. Neustadt, Wied, Germany. Lord Lilford (p). 11. 1. 1. 138.
(Schneider.)
5 <?, 3 9. Ingelheim, Eheinhessen. C. Hilgert (c). 8. 11. 2. 22-28.
2. Magdeburg, Prussia. Dr. W. Wolterstorff 0. 2. 8. 6-7.
(P)-
4 i. Bleicbe, Saxony. [H. Tor- Lord Lilford (p). 11. 1. 1. 140-
mtz^ 143.
6. Marxbeim, Bavaria. Lord Lilford (p). 11.1.1.145.
{Wolterstorff.)
$. Strass, Burgbeim. {Kor- Lord Lilford (p). 11. 1. 1. 144.
hitz.)
6. South Germany. Dr. A. Giintber (c). 59.9.6.64-69.
1. Csallokoz - Somorja, Press- Budapest Museum 94.3.1.1.
burg, Austria-Hungary. (e).
MusTELA ERMiNEA STABiLis Barrett-Hamiltou.
1904. Putorius ermineiis stabilis Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., sin, p. 394, May, 1904.
1910. Putorius {Ictis) ermineus stabilis Trouessart, Faune Mamm. d'Europe,
p. 79.
Ti/jje locality. — Blandford, Dorset, England.
Geographical distribution. — Mainland of Great Britain.
Diagnosis. — Size slightly greater than in 31. erminea Kstiva :
condylobasal length of skull in adult male, 49 to 52 '4 mm.;
mandible in adult male, 27 to 30 mm. ; teeth usually larger than
in the continental races, a peculiarity especially noticeable in the
upper carnassial ; colour averaging slightly darker above and
less strongly yellow below than in the continental forms (though
in extreme instances the underparta are between buff and straw-
yellow, quite as in the brightest continental specimens) ; tail
usually without distinct lighter area below.
Measurements. — ■ External measurements of type (adult
female) : head and body, 244 ; tail, 105 ; hind foot, 43 ; ear from
meatus, 22. Another adult female from the type locality : head
and body, 245 ; tail, 102; hind foot, 43; ear from meatus, 21.
Two adult males from Banstead, Surrey : head and body, 270
and 278 ; tail, 119 and 119 ; hind foot, 47 and 49. Two adult
males from Wales : head and body, 274 and 280; tail, 120 and
MUSTELA 391
120; hind foot, 48 and 48-6. Adult male from Farr, Daviot,
Inverness : head and body, 254; tail, 110; hind foot, 48 "5.
Specimens exanmicd. — Seventy-four, from the following localities : —
Scotland : Thurso, Caithness, 1 ; Suthorlaudshire, 1 ; Cromarty, 4
Annadale, Skye, 2 ; Farr, Daviot, Inverness, 1 (Wilson) ; Stockbriggs
Lanark, 9; Wyseley, Dumfries, 2; Ecclefechan, Dumfries, 1 (U.S.N.M.).
Wales: North Wales, no exact locality, 1; Nannerch, Flintshire, 2
Vaynol, Carnarvonshire, 2 ; Usk, Monmouthshire, 2 ; St. Brides, Pembroke
shire, 1 ; St. Fagan's, Cardiff.
England: Riding-Mill-ou-Tyne, Northumberland, 1; Westmoreland, 1
Doncaster, Yorkshire, 2 ; Leeds, Yorkshire, 1 (U.S.N.JI.) ; Gainsborough
Lincolnshire, 1 ; Derbyshire, 1 ; Sandringham, Norfolk, :3 ; Lowestoft
Suffolk, 1 ; Bury St. Edmunds, Suffolk, 1 ; Friswell, Cambridgeshire, 8
Rugby, Warwickshire, 1 ; Graftonbury, Hereford, 3 ; Triug, Hertford, 2
Pelden, Hertfordshire, 1; Banstead, Surrey, 2; Laseley Park, Guildford
Surrey, 1 ; Buckland, Somerset, 2 ; Blandford, Dorset, .3 ; Eversley
Hampshire, 1; Whitechurch, Hampshire, 1; Selborne, Hampshire, 1
Staplehurst, Kent, 1 ; Horsham, Sussex, 2 ; Mayfield, Sussex, 3.
Bemarks. — In the British stoat the change to the white
winter coat does not take place so regularly and completely as in
the continental forms. This is particularly true in central and
southern England. Apart from this character, of problematical
value, the animal is distinguishable by its heavy teeth. A few
small though apparently well developed skulls from the north
of Scotland indicate the possible existence there of a local form
somewhat resembling true erminea.
9. Thurso, Caithness, Scot- W. R. Sherrin (p). 9. 1. 9. 1.
land (Mrs. J. Edis).
3 9. Cromarty. W.R.Ogilvie-Graut 11.1.3.175-177.
(c & p).
<5, 9. Annadale, Skye. Dr. Hastings (c&p). 11.1.3.395-396
<^. Sutherlandshire. W. Paterson (c&p). 8.11.26.1.
3 6, St. Stockbriggs, Lanarkshire. B. R. Alston (c & p). 79. 9. 25. 16, IS,
20.
3 6, 9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 14, 15,
17, 19.
9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 84.
(5,9. Naunorch, Flintshire, A.Richardson 11.1.3.192-193.
Wales. (c&p).
2 9. Vaynol, Carnarvonshire. J. E. Harting (p). 11. 1. 3.194-195.
2 6. Usk, Monmouthshire. J. S. Phillips (c & p). 87. 8. 6. 1-2.
St. Brides, Pembroke- Hon. C. Edwardes 90. 12. 5. 1.
1. St. Pagans, Cardiff. J. Cording (c). 90. 6. 19. 1.
st. Derbyshire, England. Index iluseum. 94. 4. 8. 1.
St. Sandringham, Norfolk. H.M. King Edward 96. 4. 13. 1.
VII. (p).
2 St. Sandringham, Norfolk. H.M. King Edward 96. 11. 24. 1-2.
VII. (p).
9 St. Lowestoft, Suffolk. F. S. Worthington 98. 2. 15. 1.
(c & p).
St. Bury St. Edmunds, Suf- T. Harcourt-Powell 87. 2. 28. 1.
folk. (c & p).
3<J,4 9. Friswell, Cambridgeshire. W. Farren (c). 4. 5. 3. 1-8.
i. Rugby, Warwickshire. E. E. Austen (c&p). 11.1.3.181.
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MUSTELA 397
6,2 9. Graftonbury, Hereford- W. E. de Winton (p). 11.1.3.178-180.
shire.
2 9. Tring, Hertfordshire. Hon. N. C. Boths- 11.1.3.182-183.
child (p).
6. Felden, Hertfordshire. 11. 1. 3. 184.
1. Nr. Hounslow, Middle- W. Phillips (p). 60. 1. 23. 2.
sex.
9. Guildford, Surrey. G.Dalgleish (c &p). 4.4.3.1.
6 St. Horsham, Sussex. Sir E. G. Loder, 6. 7. 14. 1.
Bart, (c & p).
6,9. Mayfield, Sussex. C. H.B. Grant (c&p). 11.1.3.186-187.
6. Selborne, Hampshire. Dr. B. Bowdler 2. 11. 29. 1.
Sharpe (c & p).
9. Whitchurch, Hampshire. W. E. de Winton 11. 1. 3. 185.
(c & p).
6, 9. Buckland, Somerset. B. Hooper (c & p). 11. 1. 3. 190-191.
9. Blandford, Dorset. J. C. ManseU Pley- 98. 5. 13. 2.
dell (c & p). (Type of subspecies.)
2 9. Blandford, Dorset. W. T. Blanford 11.1.3.188-189.
(c & p).
MuSTELA EEMINEA EICIN^ Miller.
1904. Puforius erminea Barrett-Hamilton, Aim. Scottish Nat. Hist., v,
p. 203, October, 1904.
1907. Putorius erminea ricime Miller, Ann. and Mag. Nat. Hist., 7th ser.,
XX, p. 395, November, 1907.
1910. Putorius ermineus richix Trouessart, Faune Mamm. d'Europe, p. 80.
Type locality. — Islay House, Islay Island, Scotland.
Geographical distribution. — Islands of Islay and Jura, off coast
of south-west Scotland.
Characters. — Size a little less than in M. erminea stahilis and
about equal to that of M. e. aestiva : condylobasal length of skull
in adult male, 47 to 50 mm. ; mandible in adult male, 27 to
29 mm. ; zygomatic arches unusually wide-spreading ; posterior
extension of palate relatively wider than in any of the other
known forms.
Measurements. — External measurements of type (adult male) :
head and body, 254 ; tail, 105 ; hind foot, 43 ; ear from meatus,
22. Two other males from the type locality : head and bodv,
270 and 270 ; tail, 111 and 114 ; hind foot, 47 and 46. Two
adult females from the type locality : head and body, 23 1 and
234 ; tail, 95 and 105 ; hind foot, 38 and 39. Two adult males
from Jura : head and body, 220 and 225; tail, 120 and 125;
hind foot, 45 and 46 "5.
Specimens exatnined. — Thirteen, six from Jura and seven from Islay
(B.!M. and Cambridge).
Itemnr1c.<>. — While readily distinguishable from the large true
atubilis of England, the Islay and Jura stoat will probably prove
to be more nearly related to the smaller form occurring on the
398
CARNIVORA
mainland of Scotland, when the status of the Scotch animal is
more clearly understood.
1. Islay, Scotland. H. Morrison (c & p). 7. 10. 9. 1.
(Type of subspecies.)
6, 2 ?. Islay, Scotland. H. Morrison (c & p). 11. 1. 3. 196-198.
MUSTELA HIBERNICA Thomas and Barrett-Hamilton.
1895. Putorius Jdbcrnicus Thomas and Barrett-Hamilton, The Zoologist,
.3rd ser., xix, p. 12.5, April, 1895 (Ireland).
1895. Putorius hibernicus Thomas, The Zoologist, 3rd ser., xix, p. 226,
June, 1895 (Isle of Man).
1900. [Mustdrc] hihernica Schulze, Zeitschr. fiir Naturwissensch., lxxiii,
p. 221.
1910. Putorius (Ictis) hibernicus Trouessart, Fauna Mamm. d'Europe, p. 80.
Ty2)e locality. — Enniskillen, Co. Fermanagh, Ireland.
Oeogra])hiccd distribution. — Ireland and the Isle of ^lan.
Diagnosis. — ^Like Mustela erminea but slightly smaller (hind
foot in adult male rarely attaining 50 mm. ; condylobasal length
of skull in adult male, 44 to 50 mm.) ; ear and upper lip without
whitish edging ; brown of sides of body usually encroaching on
yellowish or whitish of underparts, frequently so much so as
to extend completely across throat and middle of belly, the
line of demarcation, except in rare instances, very irregular.
External characters. — The general external characters, includ-
ing the proportion of tail to head and body, and the long,
full pencil, are as in Mustela erminea ; manmipe usually more
numerous : a2 — 2, « 3 — 3 or 4 — 4 = 10 or 12.
Colour. — Except in the rare instances in which a partial or
complete white coat is assumed in winter, the actual colour is
very constant throughout the year, and such variation as there
is appears to be due chiefly to the greater or less abundance of
the long hairs on back. These hairs are a dark glossy brown
very nearly the bistre of Ridgway. The hairs of the dense
underfur are smoke-grey at base, changing to a yellowish broccoli-
brown distally. From the varjang coralnnations of these elements
diflerent shades of bi'own result which range from a yellowish
bistre toward isabella-colour, the bistre usually more pure along
median dorsal region and on head, the isabella-colour on sides,
legs and tail. Upper lij) and entire ear concolor with dark
area. No dark spot behind angle of mouth. Feet usually dark
but frequently varied with whitish, this colour appearing at tips
of toes and spreading upwards. Tail without evident lighter
area on under side. Underparts varying from a creamy white
to a pale straw-yellow. The line of demarcation between the
dark and light areas is very variable in both form and position.
In the extreme development of the light area the line of demarca-
tion extends along sides in the same position as in 3lusicla
erminea, except that it passes somewhat further below ear and
eye and ends at or just above angle of mouth. In the extreme
399
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400
CARNn'ORA
extension of the dark area the brown forms a band 20 mm. wide
across throat just in front of fore legs, and spreads over whole
median region of belly, so that the white is confined to three
patches, one extending from chin to throat band, another occupy-
ing chest between front legs and spreading on inner side of
forearm, and the third in region between hind legs and spreading
on inner surface of thighs. Between these extremes every
gradation may be found, including the most irregular arrange-
ments of isolated dark spots in the regions where the dark cross
bands occur.
Slull and teeth. — Except for their slightly smaller size the
skull and teeth are indistinguishable from those of Miistcia
erminea. The difference is, however, particularly well marked as
compared with the large British M. erminea stahilis, the nearest
geographical ally.
3Ieasure)aents. — External measurements of type (adult male) :
head and body, 228 ; tail, 88 ; hind foot, 42 ; ear from meatus,
21. Average and extremes of four males fi-om Cappagh, Water-
ford : head and body, 271 (256-283) ; tail. 111 (103-117) ; hind
foot, 48 '2 (46-51). Average and extremes of three females
from Powerscourt, Co. Wicklow : head and body, 209 (205-221) ;
tail, 78 (77-81) ; hind foot, 37 "6 (36-40). For cranial measure-
ments see Table, p. 399.
Specimens examined. — Forty-four, from the following localities : —
Ireland : Carrick, Donegal, 1 ; Colebrooke, Fermanagh, 8 ; Enniskillen,
Fermanagh, 2 ; Clandeboj'e, Down, 1 ; Glaslough, Monaghan, 1 ; Board-
mills, Down, 2 ; Castle Hamilton, Cavau, 1 ; ]Mountainstown, Meath, 3 ;
Clonlarock, Galway, 1 ; Woodfair, Galway, 1; Carna, Galway, 1 ; Temple-
more, Tipperary, 1 ; Geashill, Kings, 1 ; Powerscourt, Wicklow, 4 ;
Bagenalstown, "Carlow, 1 ; New Ross, Wexford, 4 ; Arthurstown, Wex-
ford, 2 ; Cappagh, Waterford, 4; Lismore, Waterford, 1.
Isle or Man : Lewaig, Ramsey, 1 ; Santon, 2 ; Tholt-y-Will, Snaefell, 1.
Remarks. — The Irish stoat is strikingly distinct from the
other European species of the sub-genus Mustela, though super-
ficially resembling certain North American members of the
group. It is at once recognizable by the combination of black-
tipped, heavily pencilled tail with entirely dark ear and upper
lip. It is also the only European stoat in which the line of
demarcation on sides of body is irregular and in which the dark
colour of sides tends to invade the light ventral area. In
exceptional instances* the colour pattern of body is like that of
Mustela erminea. While the colouring of the body and head
suggests that of Mustela nivalis, the animal is evidently a true
stoat, as shown by the heavily-pencilled, black-tipped tail and
the form of the skull.
<5 St. Carrick, Donegal, Ireland. Hon. N. C. Roths- 0.5.17.1.
child (P).
3i. Colebrooke, Fermanagh. Sir D. Brooke (c & p). 11. 1. 3. 199-201.
In four of the forty-one skins examined.
MUSTELA 401
S. Enniskillen, Fermanagh. J. E. Harting (c&p). 95.4.5.1.
(Type of species.)
£ St. Clandeboye, Down. Hon. N. C. Roths- 0. 5. 17. 2.
child (p).
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