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http://www.archive.org/details/cu31924003031220
CATALOGUE
OF THE
MAMMALS
OF
WESTERN EUROPE
(EUROPE EXCLUSIVE OF RUSSIA)
IN THE
COLLECTION
OF THE
BRITISH MUSEUM
BY
GEEEIT S. MILLEE
LONDON
PRINTED BY ORDER OF THE TRUSTEES OF THE
BRITISH MUSEUM
SOLD BY
Longmans, Green & Co., 39, Patebnobteb Row, E.C.
B. Quabitch, 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
qL
B56 "j>
'9?
A, 311151
LONDON :
PRINTED BT WILLIAM CLOWES AND SONS, LI5IITED,
DOKE STREET, STAMFORD STREET, S.E., AND GREAT WINDMILL STREET, W.
V>3
PREFACE
Although the idea of a publication on 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 Fund and by the efforts of Major
G. E. H. Barrett-Hamilton, who published many papers on
the subject, and of Mr. Oldfield Thomas, F.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, Bev. 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. Eothschild and Mr. J. I. S. Whitaker. The
Catalogue could hardly have been contemplated if it had
not been for Mr. Thomas' unremitting efforts in developing
the collection. He has not merely regarded these 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 the 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 F. HAEMEE,
Keeper of Zoology.
British Museum (Natural History),
London, S.W.
October, 1912.
INTKODUCTION
The collection of European Land-mammals in the British
Museum consists of about live 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, s^uch 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
Germany, 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 Spain presented by the
late Lord Lilford ; extensive collections made in south-western
France, in southern Italy and in Sicily by A. Robert 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 Mr. 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
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. Rothschild
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 ; (6)
Sicily, Italy and the region of Barcelonnette, Basses-Alpes,
France, by Dane Coolidge ; (c) south-western France, by
Robert T. Young ; (d) north-eastern Germany, the Riesengebirge
and Harz 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
Crespon), Paris (types of Geoffroy and other historic speci-
mens), Genoa (Italian Bats, Microtines and Ungulates), Turin
(Italian mammals, especially Ungulates), Naples (type of Myotis
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
specimens of Lynx), Lausanne Agricultural School (skull of
Ursus "formicarius " from the Alps), Munich (type of Spalax
grzecus Nehring), Berlin Agricultural High School (type of
Arvicola ratticeps stimmingi 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, Lemrnus lemmus
" crassidens ") 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 Ghidini,
of Geneva (Swiss and north Italian mammals), and Dr. Fernand
Lataste, of Cadillac-sur-Garonne, France (carnivores and micro-
tines).
The total number of specimens 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.!
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 Bubalus bubalis in Italy and
Simla sylvanus % on the Rock of Gibraltar, which certainly owe
* In these lists 11,372 specimens are recorded. They are distributed
as follows : Inseotivora, 1,777 ; Chiroptera, 2,210 ; Carnivora, 877 ; Kodentia
Duplieidentata, 379, Rodentia Simplioidentata, 5,854 ; Ungulata, 284.
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.M. 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.
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 fairly 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 be lost.
The literature 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, 1857, and Trouessart, 1910, and
to any other 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
followed 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 drawings 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 prepared all the descriptions, synonymies,
lists of specimens examined, and Tables of cranial measurements.
The external measurements, which are not to be regarded as
INTRODUCTION i x
more than 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,
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.
Insbctivora
7
45
5
Chiroptera
10
33
Oarnivora
15
47
1
EODENTIA DUPDICIDENTATA
2
19
1
„ SlMFI/ICIDENTATA
26
139
6
2
Ungulata
9
31
9
4
Total
69
314
22
6
SYSTEMATIC INDEX
ORDER INSECTIVORA.
Family TALPID^E . . .
2
Sub-Family 1. Talpin^e . .
2
1. Talpa Linnaeus
3
1. europaea Linnseus .
3
2. caeca Savi ....
. 15
3. oocidentalia Cabrera .
. 15
4. romana Thomas
. 18
Sub-Family 2. Desmanum .
. 20
1. Galemys Kaup
. 20
1. pyrenaious Geofiroy .
. 21
a. pyrenaicus Geofiroy
. 26
6. rufulus Graells .
. 26
Family SORICIDiE . . .
. 28
1. Sorex Linnseus
. 29
1. araneus Linnaeus .
. 31
a. araneus Linnaeus
. 35
b. castaneus Jenyns
. 37
c. santonus Mottaz
. 40
d. euronotus Miller
. 41
e. bergensis Miller .
. 41
/. tetragonurus Hermann
. 42
g. pyrenaious Miller
. 44
h, fretalis Miller
45
i. granarius Miller .
52
2. minutus Linnaeus .
53
a. minutus Linnaeus .
55
b. luoanius Miller
60
3. alpinus Schinz .
60
a. alpinus Schinz
62
6. heroynious Miller
63
2. Neomys Kaup ....
65
1. fodiens Sohreber
66
a. fodiens Sohreber.
69
b. bioolor Shaw .
73
2. milleri Mottaz .
78
3. anomalus Cabrera .
'81
3. Paohyura de Selys-Long
champs
81
1. etrusca Savi.
83
4. Crocidura Wagler.
86
1. leucodon Hermann
88
2. mimula Miller . . . .
94
a. mimula Miller .
95
6. iculisma Mottaz .
98
c. cantabra Cabrera
99
PAGE
3. russula Hermann ... 99
a. russula Hermann . . 101
b. pulchra Cabrera . . . 103
c. eintrae Miller. . . . 108
4. sicula Miller .... 108
5. caneae Miller .... 109
6. caudata Miller .... 110
7. cyrnensis Miller . . . Ill
8. balearica Miller . . . 112
Family ERINACEIDiE ... 114
1. Erinaceus Linnaeus . . . 114
1. europaeus Linnaeus . . . 115
a. europaeus Linnaeus . . 120
6. hispanicus Barrett-
Hamilton 122
c, italicus Barrett-Hamil-
ton .. . ... 123
d. consolei Barrett-Hamil-
ton 126
2. roumanicus Barrett-
Hamilton 127
3. nesiotes Bate .... 129
4. algirus Duvernoy and
Lereboullet .... 130
a. algirus Duvernoy and
Lereboullet .... 131
b. vagans Thomas . . . 133
ORDER CHIR0PTERA.
Sub-Obdeb microchiroptera.
Family RHINOLOPHEDiE . . 136
1. Rhinolophus Lacepede . . 137
1. ferrum-equinum Sohreber 139
a. ferrum-equinum Sohre-
ber 142
6. insulanus Barrett-
Hamilton 147
2. hipposideros Bechstein . 147
a. hipposideros Bechstein . 149
6. minimus Heuglin .
c. minutus Montagu .
3. euryale Blasius .
4. mehelyi Matsohie .
151
154
155
159
5. blasii Peters 162
SYSTEMATIC INDEX
Order Chieoptera — Sub - Order
MlCROCHIEOPTEBA — COlltd.
Family VESPERTILIONIDiE . 165
SUB-FAMILY VeSPEBTILIONIM! .
1. Myotis Kaup . .
1. mystacinus Kuhl
2. nattereri Kuhl .
3. emarginatus Geoffroy .
4. bechsteinii Kuhl
5. daubentonii Kuhl .
6. capaccinii Bonaparte .
7. dasyoneme Boie
8. myotis Borkhausen
9. oxygnathus Montioelli
2. Pipistrellus Kaup
1. pipistrellus Schreber .
2. nathusii Keyserling and
Blasius
3. kuhlii Kuhl . . . .
4. savii Bonaparte .
3. Eptesious Rafinesque
1. serotinus Schreber
2. sodalis Barrett-Hamilton.
3. nilssonii Keyserling and
Blasius
4. Vespertilio Linnaeus .
1. murinus Linnaeus .
5. Nyctalus Bowdich
1. maximus Patio.
2. nootula Schreber
3. leisleri Kuhl
4. azoreum Thomas
6. Pleootus Geoffroy .
1. auritus Linnaeus
7. Barhastella Gray .
1. barbastellus Schreber
Sub-Family Miniopteein^j:
1. Miniopterus Bonaparte
1. schreibersii Kuhl
Family MOLOSSIDiE .
1. Nyctinomus Geoffroy
1. teniotis Rafinesque
ORDER CARNIVORA.
165
166
169
174
177
179
184
187
189
192
199
202
204
213
215
219
224
226
231
234
238
238
242
244
245
252
254
256
256
263
263
268
208
269
276
276
277
Family URSID-&! .
284
1. Ursus Linnaeus
285
1. arctos Linnaeus .
285
2. Thalarctos Gray .
297
1. maritimus Phipps .
298
Family CANEDiE . . .
303
1. Oanis Linnaaus
304
1. lupus Linnaeus .
305
a. lupus Linnaeus .
313
b. signatus Cabrera
314
c. deitanus Cabrera
315
2. aureus Linnaeus
315
paob
2. Alopex Kaup . . . 318
1. lagopus Linnasus . . . 319
2. spitzbergenensis Barrett-
Hamilton and Bonhote . 324
3. Vulpes Oken 325
1. vulpes Linnaeus . . . 326
a. vulpes Linnaeus . . . 330
b. crucigera Bechstein . . 331
c. silacea Miller .... 333
2. ichnusae Miller . . 336
Family MUSTELID^l ... 340
Sub-Family Melin.33 . 341
1. Meles Brisson. . . ". 341
1. meles Linnaeus . . 343
a. meles Linnaeus . . . 348
6. marianensis Graells. . 352
2. arcalus Miller .... 352
Sub-Family Luteins .... 354
1. Lutra Brisson . . . . 354
1. lutra Linnaeus . . 355
Sub-Family Mustelines . . . 364
1. Martes Pinel ... .365
1. martes Linnseus . . 366
a. martes Linnaeus . . 372
6. latinorum Barrett-Ha-
milton 373
2. foina Erxleben . . .374
a. foina Erxleben . . . 375
b. mediterranea Barrett-
Hamilton 380
3. bunites Bate . . . 380
2. Mustela Linnaeus. . . . 381
Sub-genus Mustela Linnaeus . 384
1. erminea Linnasus . . . 385
a. erminea Linnaeus . » . 387
b. sestivaKerr .... 389
c. stabilis Barrett-Hamil-
ton 390
d. ricinae Miller . . .397
2. hibernica Thomas and
Barrett-Hamilton . . 398
3. nivalis Linnaeus . . 401
a. nivalis Linnaeus . . . 402
b. boccamela Bechstein . 405
u. iberica Barrett-Hamilton 407
4. africana Desmarest . . 412
5. galinthias Bate . . . 414
Sub-Genus Lutreola Wagner . 415
6. lutreola Linnaeus . . . 415
Sub-Genus Putorius Cuvier . 418
7. putorius Linnaeus . . 419
a. putorius Linnseus . . 423
b. aureolus Barrett-Hamil-
ton 425
3. Vormela W. Blasius ... 428
1. peregusna Gueldenstaedt . 429
Sub-Family Guloninje . . 432
1. Gulo Storr 433
1. gulo Linnaeus . . 434
SYSTEMATIC INDEX
Order Carnivora — continued.
Family VIVERRIDjE . . .
1. Mungos Geoffroy and Cuvier
1. widdringtonii Gray
2. Genetta Oken .
1. genetta Linnaeus .
a. genetta Linnaeus
6. balearica Thomas
c. rhodanica Matsehie
Family FELID^l . . .
1. Felis Linnaeus.
1. silvestris Sohreber .
a. silvestris Sohreber
6. grampia Miller .
c. tartessia Miller
2. sarda Lataste .
3. agrius Bate .
2. Lynx Kerr ....
1. lynx Linnaeus .
2. pardellus Miller
440
440
441
446
447
451
452
452
455
456
457
462
464
465
468
470
470
471
475
ORDER RODENTIA.
Sub-Obdbb duplicidentata.
Family LEPORIDiE .... 484
1. Oryetolagus Lilljeborg . . 484
1. eunioulus Linnaeus . 485
a. eunioulus Linnaeus . . 490
o. huxleyi Haeokel . . . 491
2. Lepus Linnaeus .... 495
1. europaeus Pallas . . 498
a. europaeus Pallas. . . 502
b. oooidentalis de Winton. 504
c. pyrenaious Hilzheimer . 506
d. meridiei Hilzheimer . 506
e. oorsioanus de Winton . 507
/. hybridus Desmarest . . 508
g. transsylvanicus Mat-
sehie 509
2. cretious Barrett-Hamilton 512
3. mediterraneus Wagner . 513
4. granatensis Rosenhauer . 515
a. granatensis Rosenhauer 516
6. gallaecius Miller ... 517
c. iturissius Miller . . . 518
d. parnassius Miller . . 519
5. timidus Linnaeus . . . 522
a. timidus Linnaeus . 526
b. varronis Miller . . . 528
c. seotious Hilzheimer. . 529
6. hibernicus Bell. . . . 531
Sub-Oedeb SIMPLICIDENTATA.
Family ZAPODIDiE .... 535
Sub-Family Sicistin.^
1. Sicista Gray .
1. loriger Nathusius
2. trizona Petenyi .
536
536
537
539
Family HYSTRICIDiE
1. Hystrix Linnaeus ....
1. cristata Linnaeus .
Family MUSCARDINIDiE . .
1. Eliomys Wagner ....
1. queroinus Linnaeus
2. gymnesieus Thomas .
3. pallidus Barrett-Hamilton
4. sardus Barrett-Hamilton .
5. lusitanious Reuvens .
2. Dyromys Thomas.
1. nitedula Pallas .
a. nitedula Pallas .
6. intermedius Nehring .
c. wingei Nehring .
2. robustus Miller.
3. Glis Brisson
1. glis Linnaeus ....
a. glis Linnaeus ....
6. italious Barrett-Hamil-
ton
c. melonii Thomas .
d. pyrenaicus Cabrera .
4. Muscardinus Kaup .
1. avellanarius Linnaeus .
2. pulcher Barrett-Hamilton
Family MURID.S3 ....
Sub-Family Cbicetin^; .
1. Crioetulus Milne-Edwards .
1. atticus Nehring . .
2. Crieetus Leske ....
1. erioetus Linnaeus .
a. crieetus Linnaeus
b. oanesoens Nehring .
c. nehringi Matsohie .
3. Mesooricetus Nehring .
1. newtoni Nehring .
Sub-Family Micbotinje
1. Myopus Miller ....
1. schisticolor Lilljeborg.
2. Lemmas Link
1. lemmus Linnaeus .
3. Evotomys Coues .
1. glareolus Sohreber .
a. glareolus Sohreber
6. britannious Miller
sueoious Miller .
isterious Miller .
norvegious Miller
vaseoniae Miller .
helvetious Miller
h. nageri Sohinz
i. hallucalis Thomas
2. skomerensis Barrett
milton ....
3. oaesarius Miller .
4. rutilus Pallas .
5. rufooanus Sundevall
c.
d.
e.
/■
Ha-
paoe
542
542
543
549
550
551
558
559
560
560
566
567
568
569
570
572
572
573
577
578
579
582
583
583
590
591
592
593
593
596
597
602
603
605
605
606
610
611
611
614
615
623
626
632
634
636
637
638
639
640
641
643
644
645
646
648
SYSTEMATIC INDEX
Order Kodentia — Sub - Order
Simplictdentata — Family
Mubid-e — Sub-Pamily Mi-
cbotin-se —continued.
4. Miorotus Schrank . . . 658
Sub-Genus Miorotus Schrank 659
1. agrestis Linnaeus . . . 662
a. agrestis Linnseus . . 668
b. exsul Miller .... 669
c. levernedii Crespon . . 671
d. bailloni de Selys-Long-
ohamps 672
e. airbus Bellamy . . 673
/. negleotus Jenyns . . . 675
g. rozianus Bocage . . . 680
2. arvalis Pallas . . . 681
a. arvalis Pallas . . . 683
b. meridianus Pallas . . 686
c. duplioatus Rorig and
Borner .... 686
d. levis Miller .... 687
3. incertus de Selys - Long-
champs 690
4. asturianus Miller . . . 693
5. orcadensis Millais . . . 694
6. sandayensis Millais . . 696
a. sandayensis Millais . . 697
b. westrse Miller . 698
7. sarnius Miller , . . 700
8. eabrerse Thomas . . . 701
9. dentatus Miller. ... 703
10. hartingi Barrett-Hamilton 704
11. angularis Miller . . . 706
12. ratticeps Keyserling and
Blasius. . ... 708
Sub-Genus Chionomys Miller 712
13. nivalis Martins ... 713
a. nivalis Martins . . . 716
b. aguitanius Miller . . 717
14. lebrunii Orespon . . 718
a. lebrunii Crespon. . 719
o. leueurus Gertie . . 722
15. ulpius Miller ... 723
5. Arvicola Lacepede . . . 723
1. amphibius Linnseus . . 725
a. amphibius Linnaeus . 730
b. reta Miller . 732
2. sapidus Miller . . . 732
a. sapidus Miller . . . 733
b. tenebrieus Miller . . 735
3. terrestris Linnseus. . 738
4. italicus Savi 740
5. illyricus Barrett-Hamilton 741
6. musignani de Selys-Long-
champs . . . 744
7. scherman Shaw . 744
a. scherman Shaw . . 745
6. exitus Miller .... 746
c. monticola de Selys-
Longchamps .... 749
6. Pitymys McMurtrie . . .
752
1. subterraneus de Selys-
Longchamps ....
755
a. subterraneus de Selys-
Longchamps ....
758
6. capucinus Miller
760
2. daeius Miller
760
3. druentius Miller
762
4. fatioi Mottaz ....
763
5. multiplex Patio
764
6. savii de Selys-Longehamps
768
7. nebrodensis Mina-Palumbo
770
8. pyrenaicus de Selys-Long-
champs
770
a. pyrenaicus de Selys-
Longchamps ....
771
6. brunneus Miller .
772
9. planiceps Miller .
772
10. gerbii Gerbe ....
773
11. lusitanicus Gerbe.
776
12. marise Major ....
777
13. pelandonius Miller .
778
14. depressus Miller
779
15. ibericus Gerbe
780
a. ibericus Gerbe .
782
6. centralis Miller
782
c. pascuus Miller .
783
d. regulus Miller
784
16. duodecimcostatus de
Selys-Longchamps
784
17. provincialis Miller
785
18. thomasi Barrett-Hamil-
786
19. atticus Miller .
787
Sub-Family Musing .
791
1. Apodemus Kaup .
791
1. epimelas Nehring .
794
2. sylvaticus Linnseus
797
a. sylvaticus Linnseus .
803
6. callipides Cabrera
809
c. dichrurus Rafinesque .
810
d. creticus Miller .
813
3. hebridensis de Winton
824
4. hirtensis Barrett-Hamil-
ton ....
825
5. fridariensis Kinnear .
825
6. flavicollis Melchior
828
a. flavicollis Melchior .
829
b. wintoni Barrett-Hamil-
ton .
831
7. agrarius Pallas ....
836
2. Micromys Dehne .
840
1. minutus Pallas .
841
a. soricinus Hermann .
844
6. pratensis Ockskay .
846
3. Epimys Trouessart .
848
1. rattus Linnseus .
849
a. rattus Linnseus .
853
6. alexandrinus Geofiroy .
854
2. norvegicus Erxleben .
858
SYSTEMATIC INDEX
Order Rodentia — Sub - Order
Sihplicidentata — Family
Mubid^; — Sub-Family Mu-
BiNai — continued.
4. Mus Linnaeus 863
1. musculus Linnaeus . . 865
a. musculus Linnaeus . . 869
6. azorious Schinz . . . 871
2. muralis Barrett-Hamilton 874
3. faeroensis Clarke . . . 875
4. spioilegus Petenyi . . 877
a. spioilegus Petenyi . 878
6. hispanicus Miller . . 879
c. lusitanious Miller . 882
5. Aoomys Geoffroy . . . 883
1. minous Bate .... 883
Family SPALACIDJE ... 887
1. Spalax Gueldenstaedt . . 887
1. dolbrogeae Miller ... 889
2. hungaricus Nehring . . 894
3. graeous Nehring . . . 895
Family SCIURIDiE .... 897
1. Sciurus Linnaeus .... 898
1. vulgaris Linnaeus . . . 898
a. vulgaris Linnaeus . . 905
6. varius Gmelin . . . 906
c. leucourus Kerr . . . 907
d. russus Miller. ... 909
e. fuscoater Altum . . . 910
/. italious Bonaparte . . 912
g. lilaeus Miller .... 913
h. alpinus Desmarest . 914
i. numantius Miller . . 914
j. infuscatus Cabrera . 916
k. segurae Miller . . . 917
I. baetious Cabrera . . . 923
2. Citellus Oken 924
1. citellus Linnaeus . . . 924
2. suslica Gueldenstaedt . . 929
3. Marmota Blumenbach . . 931
1. marmota Linnaeus . . 932
2. bobak Miiller .... 937
Family PETAUBJSTID^ . . 940
1. Sciuropterus F. Cuvier . 941
1. russicus Tiedemann . . 941
Family CASTORIDJE ... 947
1. Castor Linnaeus .... 947
1. fiber Linnaeus .... 947
OBDEB UNGULATA
PAGE
Family SUIDjE 956
1. Sus Linnaeus 956
1. scrofa Linnaeus. . . 957
[attila Thomas] .... 960
2. meridionalis Major . . 960
Family CERVIDiE ....
1. Cervus Linnaeus .
1. elaphus Linnaeus .
a. germanicus Desmarest .
b. elaphus Linnaeus
c. atlanticus Lonnberg
d. scoticus Lonnberg .
e. hispanicus Hilzheimer .
/. corsicanus Erxleben.
2. Dama Hamilton Smith
1. dama Linnaeus .
3. Gapreolus Gray .
1. capreolus Linnaeus .
a. capreolus Linnaeus .
b. transsylvanicus Matschie
c. canus Miller ....
d. thotti Lonnberg.
4. Alces Gray
1. alces Linnaeus ....
5. Bangifer Hamilton Smith .
1. tarandus Linnaeus .
2. f ennicus Lonnberg .
3. platyrhynchus Vrolik .
Family BOVID.-E ....
1. Ovis Linnaeus
1. musimon Pallas
2. Capra Linnaeus .
1. ibex Linnaeus
2. pyrenaica Schinz .
a. pyrenaica Schinz
b. lusitanica Franca .
c. victoriae Cabrera.
d. hispanica Schimper .
3. aegagrus Erxleben
3. Rupricapra Blainville
1. rupicapra Linnaeus
2. ornata Neumann .
3. pyrenaica Bonaparte .
4. parva Cabrera
962
963
964
965
967
967
968
969
969
970
970
972
973
974
975
975
975
976
978
979
980
981
9S5
986
986
987
988
989
989
990
991
991
991
992
992
993
994
995
995
APPENDIX.
Crocidura ichnusae Festa . . . 998
Evotomys glareolus norvegicus —
cranial measurements . . . 999
Spalax — Forms recognized by
Mehely 1000
CATALOGUE
OP THE
LAND-MAMMALS OF WESTERN
EUROPE.
I
Order INSECTIVORA.
1827. Insectivora Gray, Griffith's Guvier, Anim. Kingd., v, p. 100.
Geographical distribution. — Africa (including 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 posterior upper premolar and anterior lower molar
never specially modified as carnassials.
Remarks. — 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) Erinaceidx, 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 large lateral vacuities ;
no auditory bulla ; general form mouse-like, the
neck evident; external ear present (Shrews) Soricidas, p. 28.
B
£ INSECTIVORA
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) Talpidse, 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) Talpinte, 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) Desmaninx, p. 20.
Family TALPIDEE.
1825. Talpidse Gray, Thomson's Annals of Philosophy, xxvi, p. 339.
Geographical distribution. — 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 flattened 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,
terete or depressed ; no external ear.
Remarks. — The members of the family Talpidse are at once
recognizable among European mammals by the great development
of the shoulder girdle and apparent absence of neck, the auditory
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 TALPINTE.
Geographical distribution. — Temperate portions of Europe and
Asia, from England to Japan : in Europe south to the Mediter-
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 upper
canine large, strongly trenchant ; external form highly modified
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, composed 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 Linnaeus.
1758. Talpa Linnseus, Syst. Nat., i, 10th ed., p. 52.
Type species. — Talpa europsea Linnseus.
Geographical distribution. — Europe and western and central
Asia. Eastern limits of range not known.
Characters. — Dental formula : i ,-Ej, c j5j, pin jtj, m ^ 3 = 44.
Upper 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
slightly inflated, its outlines usually indistinct, the meatus small,
sub-circular ; external form strictly talpine ; ear-conch absent ;
eye minute, often covered by the skin.
Remarks. — The genus Talpa contains half a dozen or more
species. Pour of these occur in Europe, one of them generally
distributed, the three others confined to the Mediterranean
region.
KEY TO THE EUROPEAN SPECIES OE TALPA.
Greatest diameter of m 1 about 4 mm. ; three lower
molars together about 8 mm. (vicinity of
Borne) T. romana, p. 18.
Greatest diameter of m l about 3 mm. ; three lower
molars together about 7 mm. or less.
Oondylobasal length of skull 33 to 37 mm. (dis-
tribution general) T. europsea, p. 3.
Oondylobasal length of skull 29 to 32 mm.
(southern).
Posterior border of anteorbital foramen over
front of last molar (Italian) T. cseca, p. 15.
Posterior border of anteorbital foramen over
middle of second molar (Iberian) T. occidentalis, p. 15.
TALPA EUROP.ffi:A Linnseus.
1758. [Talpa} europsea Linnaeus, Syst. Nat., i, 10th ed., p. 52 (Sweden).
1766. Talpa frisius P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band,
p. 36 (Ostfriesland).
1772. [Talpa] caudata Boddaert, Kortbegrip van het zamenstel der Natuur,
i, p. 50 (Renaming of europsea).
B 2
4 INSECTIVOEA
1777. [Talpa europsea] a albo-maculata Erxleben, Syst. Begni Anim., I,
p. 117 (Ostfriesland).
1785. [Talpa] vulgaris Boddaert, Elenohus Anim., i, p. 126 (Europe).
1789. [Talpa europsea] |8 variegata Gmelin, Syst. Nat., i, 13th ed., p. 110
(Sweden).
1789. [Talpa europsea] y alba Gmelin, Syst. Nat., I, 13th ed., p. HO
(Sweden).
1789. Talpa europsea e cinerea Gmelin, Syst. Nat., i, 13th ed., p. 110 (Eifel,
Germany).
1792. Talpa europ[sea] nigra Kerr, Anim. Kingd., p. 200 (Renaming of
europsea).
1797. Talpa europsea rufa B[orkhause]n, Der Zoologe (Compendiose Biblio-
thek gemeinniitzigsten Kenntnisse fur alle Stande, pt. xxi),
Heft v-viu, p. 13 (Southern Prance).
1836. Talpa europica flavescens Reichenbach, Praeht.-gemoinn. der
Saugeth. des In- und Auslandes, fig. 473 (Saxony).
1852. Talpa europsea albida Reichenbach, Vollstandigste Naturgesch. des
In- und Auslandes, iv, p. 336 (Germany).
1852. Talpa europsea lutea Beiehenbaoh, Vollstandigste Naturgesch. des
In- und Auslandes, IV, p. 336 (Germany).
1857. Talpa europsea Blasius, Saugethiere Deutschlands, p. 109.
1869. Talpa europsea, flavescens Pitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, dix, pt. i, p. 400.
1869. Talpa europsea, maculata Pitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lix, pt. I, p. 401
(Renaming of albo-maculata).
1869. Talpa europsea 5 grisea Pitzinger, Sitzunsgber. kais. Akad. Wissensch.
Wien, Math.-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. [Talpa] scalops Schulze, Abh. u. Vortr. Gesammtb. Naturw. iv,
no. 10, p. 19. (Substitute for europsea.)
1910. Talpa europsea Trouessart, Paune 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 first premolar 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 ro 1 about
3 mm. ; mesostyle of m 2 and m 3 entire or with apex slightly
notched.
External characters. — General form highly modified for sub-
terranean burrowing habits, the neck so short that the conical
head with much produced snout appears 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 narrow 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 near
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. Mammae :
p2-2; »2-2 = 8.
Colour. — Fur everywhere dark slaty grey, sometimes almost
blackish, at others more nearly a dark smoke-grey, the hairs
everywhere witb a noticeable metallic or purplish iridescence ;
underparts sometimes a little less dark than back, and occa-
sionally with a yellowish brown suffusion ; 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.
* Much confusion has been caused by the supposition that the presence
or absence of this minute aperture is a specific or racial character.
INSBCTIVOEA
Outline when viewed from the side long wedge-shaped, rounded
off posteriorly 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 forward and outward
from antero-external angle of interparietal along edge of slightly
inflated mastoid region, and terminating anteriorly in a slightly
pointed projection. Brain-case with posterior margin 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 slightly 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 anteropos-
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
lambdoid crest. Base of brain-
case smooth, without conspicuous
ridges, furrows or open spaces,
the bones for the most part some-
what inflated ; two ill-defined pits
in surface of basioccipital in front
of foramen magnum ; a shallow,
broadly triangular median furrow
between the low, flattened bullae,
the sub-circular, slightly triangu-
lar outline of which is sometimes
distinctly indicated, but more
often very obscure ; auditory
meatus small, nearly circular.
Inner 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
between 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 sub-cylindrical, dis-
tinctly expanded at middle. Rostrum narrower and somewhat
abruptly lower than interorbital region, its narrowest point just
behind canines ; nares with evident posterior emargination.
Anteorbital foramen moderately large, its posterior border over
metastyle of to' 2 , the plate forming outer wall of canal usually
much narrower than foramen Lachrymal foramen above anteor-
¥10. I.
Talpa, europsedf. Nat. size.
bital foramen and slightly in front of its middle ; its orifice over
metastyle of in 1 . 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 m 3 , in a slightly
raised crescentic ridge ; a vacuity about as large as the minute
incisive foramen on each side of palate opposite space between
m 1 and m 2 . 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 equal to least breadth of
posterior 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.
Teeth. — In proportion to the size of the skull the teeth are
moderately large, their general
aspect noticeably trenchant. Upper
incisors simple, chisel - shaped,
perpendicular, 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 obliquely for-
ward, essentially similar to
the upper teeth in form,
but smaller and narrower.
Upper canine large, two-
rooted,* the height of the
shaft greater than that of any
of the other teeth, and fully
equal to anterior breadth of
palate ; shaft wider ante-
riorly than posteriorly, with
antero-internal longitudinal
groove, and highly deve-
loped, slightly concave pos-
terior cutting edge. Lower
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 in old individuals.
t'IG. 2.
Talpa europvea. Teeth X 5.
canine very small, resembling a
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 i , 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 passing 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 2 .
Paracone and metacone sub-equal in m 2 and m 3 , the metacone
the larger in the former, the paracone in the latter. In m 1 the
metacone is about double 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 2 . Styles and outer commissures well
developed in m 2 and forming a distinct VV"P a ttern ; mesostyle
entire or with apex slightly notched. In m 1 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 3 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 anteriorly, second and third
slightly broader anteriorly than posteriorly, the two triangles
essentially alike in form. Metaconid of m 1 low, scarcely more
* The thickenings vary considerably in different individuals. Occasion-
ally they are 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 V, the outer surface of paracone
essentially resembling that of pm' 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.
Measurements. — 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 body, 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-Spey, Elgin, 13 (Wilson) ;
Cortachy, Forfar, 1 (Wilson) ; Stockbriggs, Lanarkshire, 1.
England : Bowdon, Cheshire, 1 ; Altrincham, 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 ; Eulbourn, Cambridge, 1 ; Holloway,
Somersetshire, 2 ; Somersetshire, no exact locality, 1 ; Banstead, Surrey, 2 ;
Coombe, 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: Waremme, Liege, 3 (U.S.N.M.).
Prance : Guines, Pas-de-Calais, 1 ; Pont-de-Briques, Pas-de-Calais, 1 ;
Trinity, Jersey, 2; St. Lawrence, Jersey, 1; Barhizon, Seine-et-Marne, 2;
Melun, Seine-et-Marne, 1 (Mottaz) ; Lignieres, Charente, 1 (Mottazl ;
Huelgoat, Brittany, 1 ; Cadillac-sur-Garonne, Gironde, 1 (U.S.N.M.) ;
Poret de Bouconne, Gers, 13; Solferino, Landes, 10; Caterille, Haute-
Garonne, 10; Legouvin, Haute-Garonne, 3; Luchon, Haute-Garonne, 3;
Bareges, Hautes-Pyrenees, 5; l'Hospitalet, Ariege, 1; Porte, Pyrenees-
Orientales, 3; St. Gilles, Gard, 1; Valescure, Var, 1; Agay, Var, 1;
Etupes, Doubs, 3 (Mottaz) ; Lucinges, Haute-Savoie, 9 ; Montauban,
Haute-Savoie, 15 ; Cranves-Sales, Haute-Savoie, 12.
Spain: Pajares, Leon, 2; Castrillo de la Beina, Burgos, 9; Castauares,
Burgos, 8 ; Lerida, 1 ; Barracas, Castellon, 24 ; Catarroja, Valencia, 4 ;
Dehesa de Valencia, Valencia, 17.
Germany : K6nigsberg,4 (U.S.N.M.) ; Moritzburg, Saxony, 8 (U.S.N.M) ;
Ummerstadt, Thiiringen, 3 ; Brunswick, 15 (U.S.N.M.) ; Aachen, 9
(U.S.N.M.); Bremen, 1; Ingelheim, Kheinhessen, 4; Strass, near Burg-
heim, Bavaria, 7; Niesky, Silesia, 1; Kalbe, Saale, 2; Magdeburg,
Saxony, 1 ; Strassburg, 3.
Austria-Hungary : Csallokfiz-Somorja, Pressburg, Hungary, 2 ; Hatszeg,
Hunyad, Transylvania, 2.
10
INSECTIVORA
Boumania : 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.) ; Mflrren, 1 ; Thurgau, 1 ; Oberhasli Valley
1; St. Gallen, 3; Bheinthal, St. Gallen, 2; Degersheim, St. Gallen, 3
Gossau, St. Gallen, 3; Untervatz. Grisons, 1; Breganzona, Tieino, 1
Cortivallo, Tieino, 6 (B.M. and U.S.N.M.) ; Comano, Tieino, 2 (U.S.N.M.)
Lugano, Tieino, 4 (B.M. and U.S.N.M.) ; Muzzano, Tieino, 2 (B.M. and
U.S.N.M.) ; Sorengo, 1 (U.S.N.M.) ; Cremignone, Tieino, 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, Pinalborgo, 10 (Genoa) ;
Florence, 10 (U.S.N.M.).
Remarks. — With the possible exception of certain bats, the
common mole shows less tendency to vary geographically than
any other European mammal of equally wide range.
2 S. Black Isle, Cromarty, Scot-
land.
?. Gordonstown, Elgin.
rf. Stockbriggs, Lanarkshire.
9. Parsop, Herefordshire, Eng-
land.
2 6. Bugby, Warwickshire.
2. Warwickshire.
6. Eulbourn, Cambridgeshire.
2 al. Holloway, Somerset.
2. Somerset. (Hilgel.)
2 al. Cobham, Surrey.
1 al. 1. Egham, Surrey.
9. Knockholt, Kent.
2 $. al. Bromley, Kent.
3 S. Devonshire.
2 c5. Trinity, Jersey, Channel
Islands. (R. H. Bunting.)
9. St. Lawrence, Jersey.
6. Guines, Pas-de-Calais, 80 ft.
Prance.
S. Pont - de - Briques, Pas - de-
Calais.
5. Huelgoat, Brittany, 600 ft.
2d, 2 9. Poret de Bouconne, Gers,
250 m. (A. Robert.)
2 <5, 2 9. Solferino, Landes. (A. Ro-
bert.)
2 c?, 2 9. Caterille, Haute - Garonne,
900-1000 m. (A. Robert.)
2 <5, 9. Luchon, Haute - Garonne.
(A. Robert.)
2 6, 1 9. Bareges, Hautes-Pyrenees.
6. L'Hospitalet, Ariege. (A.
Robert.)
S. Porte, Pyrenees-Orientales.
2 S Porte, Pyrenees - Orientales,
1600-1700 m. {A. Robert.)
i. Valesoure, Var.
W. B. Ogilvie-
Grant (p).
W. B. Ogilvie-
Grant (p).
E. B. Alston (p).
E. A. Denny (p).
E. E. Austen (p).
Tomes Collection.
H. King (p).
E. B. Alston (p).
Dr. Leach (p).
P. Heiss (p).
W. Blaekwell (p).
H. E.Bawson (p).
Oxley Grabham
O. Thomas (p).
Mrs. Power (p).
O. Thomas (c&p).
11. 1. 3. 62-63.
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.
O.Thomas (c&p).
O. Thomas (p).
O. Thomas (p).
O. Thomas (p).
0. Thomas (p.)
G. S. Miller (c).
0. Thomas (p).
G. S. Miller (c).
0. Thomas (p).
92. 9. 5. 1.
6. 4. 1. 28-31.
6. 4. 1. 32-35.
6. 4. 1. 24-27.
6. 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. Montaubah, Haute - Savoie, A. Eobert (o & p). 97. 1. 9. 1.
1000 m.
3 6, 1 9. Cranves-Sales, Haute-Savoie, A. Eobert (o & p). 5. 4. 4. 1. 5. 4,
900 m. 9. 3-5.
5(5, 3 9. Cranves-Sales, Haute-Savoie, O. Thomas (p). 5. 11. 18. 1-8.
909-1200 m. Prance. (A.
Bobert.)
6, 9. Pajares, Leon, Spain. (N. O. Thomas (p). 8. 2. 9. 26-27.
Gonzales.)
2. Castrillo de la Reina, Burgos, G. S. Miller (o). 8.8.4.20-21.
Spain.
2 6, 5 9. Castrillo de la Reina, Burgos. N. Gonzalez (c.) 8. 7. 7. 1-7.
<5. Lerida, Spain. (N. Gonzales.) O. Thomas (p). 8. 2. 9. 28.
6 6, 6 9. Barraoas, Castellon. (N. Gon- O. Thomas (p). 8. 2. 9. 29-40.
sales.)
3 <J, 9. Catarroja, Valencia. (N. Gon- O. Thomas (p). 8.2.9.12-15.
sales.)
4<5, 5 9. Dehesa de Valencia, Val- O. Thomas (p). 8.2.9.17-25.
encia. (N. GonsaUs.)
3(5,9. Ingelheim,Rheinhessen,Ger- C. Hilgert (c). 8.11.2.11-14.
many.
6,9. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.115-116.
(Schuchardt.)
6. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.137.
(Schuchardt.)
6, 3 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11. 1. 1. 1-4.
(KOrbits.)
2 6, 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.132-134.
(KOrbits.)
6. Niesky, Silesia. (W. Boer.) Dr. B. Hamilton 97. 12. 4.18.
(*)■
9. Kalbe, Saale. (W. Bauer.) Lord Lilford (p). 11. 1. 1. 5.
6. Kalbe, Saale. (W. Bauer!) Lord Lilford M. 11. 1. 1. 136.
6. Magdeburg, Saxony. (Wolter- Lord Lilford (p). 11. 1. 1. 135.
storff.)
6, 9. Strassburg, Alsace. (C. Mot- O. Thomas (p). 8. 8. 10. 10-11.
tas.)
2. Csall6k<Sz-Somorja, Pressburg, Budapest Museum 94. 3. 1. 27-28.
HuDgary. (p).
2(5. Hatszeg, Hunyad, Transyl- C. G. Danford (c). 3.2.2.20-21.
vania, Hungary.
5. Bustenari, Prahova, Rouma- Lord Lilford (p). 4. 4. 6. 13.
nia. (W. Dodson.)
6. Comana, Vlasoa, 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.
(E. H. Zollikofer.)
1. Oberhasli Valley, Switzerland. Tomes Collection. 7.1.1.130.
(Koeserman.)
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. (E. H. Zollikofer).
5. Cortivallo, Ticino. (E. H. 0. Thomas (p). 2. 8. 4. 15.
Zollikofer).
9. Lugano, Ticino, 300 m. 0. Thomas (c & p). 2. 7. 1. 1.
6. Muzzano, Ticino. (E. H. 0. Thomas (p). 2. 8. 4. 16.
Zollikofer).
9. Cremignono, Ticino. (E. H. 0. Thomas (p). 2. 8. 4. 20.
Zollikofer) .
12
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TALPA CMCk Savi.
1822. Talpa cseca Savi, Nuovo Giom. de' Letterati, Pisa, I, p. 265.
1857. Talpa cmca Blasius, Saugethiere Deutsehlands, p. 115.
1906. Talpa caeca Camerano, Boll. Mus. Zool. ed Anat. Comp. della K.
Univ. di Torino, xxi, No. 530, p. 1, June 22, 1906.
1910. Talpa cseca Trouessart, Paune Mamm. 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 europsea (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 l and m 3 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, Italy (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 : —
Switzebland : Agmizzo, Ticino, 1 ; Bellinzona, Ticino, 1 (U.S.N.M.) ;
Breganzona, Ticino, 1 (U.S.N.M.) ; Canabbio, 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.
Itals: Regello, Tuscany, 4 (U.S.N.M.); N.S. della Vittoria, Ligurian
Appenines, 1 (Genoa) ; no exact locality, 2.
<?. Agmizzo, Ticino, Switzerland. 0. Thomas (p). 2. 8. 4. 18.
(E. H. Zollikofer.)
{. Stabio, Ticino. (E. H. Zol- 0. Thomas (p). 2. 8. 4. 19.
likofer.)
6, ¥ al. Italy. Dr. Riippell (c). 45. 7. 22. 32-33.
TALPA OCCIDENTALS Cabrera.
1907. Talpa cseca occidentalis Cabrera, Ann. and Mag. Nat. Hist., 7th ser.,
xx, p. 212, September, 1907. Type in Cabrera collection.
1907. Talpa cseca occidentalis Cabrera, Bol. Ileal. Soc. Espafl. Hist. Nat.,
Madrid, vn, p. 222, October, 1907. (For date see Cabrera, Ann.
and Mag. Nat. Hist., 8th ser., I, p. 189, February, 1908.)
1910. Talpa cseca occidentalis Trouessart, Faune Mamm. d'Europe, p. 63.
Type locality. — La Granja, Province of Segovia, Spain.
Geographical distribution. — Iberian Peninsula.
16
INSECTIVORA
Diagnosis. — Like Talpa caeca but skull rather robust, the
breadth of rostrum over roots of canines usually more than
4 mm. ; anteorbital foramen contracted, its posterior border over
mesostyle of m 2 , 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 Talpa caeca.
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
ia noticeably smaller than in Talpa
caeca, and its posterior border 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 Talpa caeca. Zygoma
about as long as in Talpa caeca, but
owing to the different position of pos-
terior border of anteorbital foramen, the
distance from this point 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 caeca, 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. — Ten, from the following localities : —
Spain : Galicia, 1 ; La Granja, Segovia, 5.
Portugal : Cintra, 4.
Fig. 3.
Talpa cseca (upper figure), and
T. occidentalis (lower figure).
> T at. siz2.
1 al. Galicia, Spain.
2 S, 2 Sal. La Granja, Segovia,
Spain.
Skull. La Granja, Segovia.
2 S, 1 al. Cintra, Portugal.
1 skull.
Prof.Seoane (c&p). 94.1.1.23.
M.delaEscalera(c). 8. 7. 30. 19-22.
M.de la Escalera(c)
0. Thomas (c&p).
6. 11. 4. 2.
98.2. 2.8 9,58,
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TALPA
17
Observations.
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TALPA ROMANA Thomas.
1902. Talpa romana Thomas, Arm. and Mag. Nat. Hist., 7th ser., x, p. 517,
December, 1902. Type in British Museum.
1904. Talpa romana Camerano, Mem. Reale Aeoad. Soi. di Torino, 2nd ser.,
liv, p. 81.
1910. Talpa romana Trouessart, Paune Mamm. d'Europe, p. 64.
Type locality. — Ostia, Rome, Italy.
Geographical distribution. — Vicinity of Rome, Italy. •
Diagnosis. — Externally similar to Talpa europeea but a trifle
larger ; skull and dentition more robust tban in the related
animal (greatest diameter of m 1 about 4 mm. ; length of upper
tooth-row, exclusive of incisors, 14 to .15 mm.) ; mesostyle of all
three upper molars bifid at tip.
Sleull 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 difference particularly noticeable in the first upper
molar and in the large lower cheek-teeth (the combined length of
fig. 4.
Crown of molars in Talpa europsea (a\ and T. romana (b). X 5.
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 1 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 developed than in T. europsea, its edge frequently
forming an evident cusp, especially in m 2 .
Measurements. — External measurements of type (from
Thomas) : head and body, 126 ; tail, 29 ; hind foot, 19 (all
measurements from skin). Pour males from the vicinity of
Rome (in alcohol) : head and body, 138 '3 (130-145); tail, 28-6
(27-30) ; hind foot, 19 • 6 (19-20). Three females from the same
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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 (B.M. 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
large teeth.
1. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 8.
(Type of species.)
1. Fraseati, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 9.
t, 9 al. Rome. Genoa Museum (e). 3. 1. 31. 1-2.
2 S. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 1-2.
Sub-Family DESMANINJE.
Geographical distribution. — South-western Prance 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 Talpinse, 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.
Remarks. — The sub-family Desmaninse contains the genera
Desmana of Russia and Siberia, and Galemys 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 Talpinse.
Genus GALEMYS Kaup.
1829. Galemys Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, I,
p. 118 (Mygale pyrenaica Geoff roy).
1846. Galomys Agassiz, Nom. Zool., Index Univ., p. 149 (Emendation of
Galemys).
1849. Mygalina I. Geoff roy, D'Orbigny's Diet. Univ. d'Hist. Nat., iv, p. 709
(Mygale pyrenaica Geoff roy).
Type species. — Mygale pyrenaica Geoffroy.
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 Desmana* in which the tail is flattened
laterally throughout, the unieuspid 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 distribution. — 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 orifice
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 dense and velvety than in the moles, and of the peculiar
quality characteristic of aquatic 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., depth 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, j The median
* The synonymy of this genus is as follows : —
1777. Desmana Gueldenstaedt, " Beschaft. Berliner Gesellsoh. Naturforsch.
Freunde, in, p. 108." (Castor moschatus Linnseus.)
1799. Desman Lacepede, Tabl. Mamm., p. 7. Same type.
1800. Mygale Cuvi,er, Lee. d'Anat. Comp., i, Tabl. 1. Same type. (Described
in Tabl. Elem. d'Hist. Nat. des Anim., p. 109.)
1815. Desmanus Bafinesque, Analyse de la Nature, p. 59 (Emendation of
Desman).
1829. Myogalea Fischer, Synops. Mamm., p. 250 (Substitute for Mygale).
1830. Gaprios Wagler, Nat. Syst. Amphibien, p. 14 (Substitute for Mygale).
1836. Myogale Brandt, Wiegmann's Archiv fur Naturgesch., 1834, i, p. 176.
t The actual narial opening is situated at extreme inner portion of
nostril, and is capable of complete closure by the combined action of
antero-internal 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 finely 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 Talpa, though probably never
covered by the integument. No external ear, the meatus about
4 mm. in diameter. Eront feet rather large and broad, readily
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,
fifth and third sub-equal and
slightly shorter, second and first
still shorter. Palms naked, their
surface finely tuberculo - rugose,
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 front 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 larger ■ fourth digit longest,
third, fifth, second and first suc-
cessively shorter, the first extending nearly to end of first
phalanx of second ; surface of sole like 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
fig. 5.
Galemys pyrenaicus. Nat. size.
GALEMYS 23
a rudimentary dorsal and ventral keel. Mammas : pi — l,al — 1,
* 2 - 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 effect 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 europeea,
but the brain-case is shorter and squarely truncate posteriorly,
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 more
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 posteriorly,
completely hidden when viewed from above. Interparietal
projecting further forward than in Talpa, its anteroposterior
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 Talpa ; no pits in basioccipital
in front of foramen magnum. Bullae 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 reduced 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. Nares 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 middle of m 1 . Posterior border of
anteorbital foramen over parastyle of m 2 . Palate essentially as
in Talpa, but vacuities smaller and incisive foramina large, their
24
INSECTIVORA
longitudinal diameter about equal to width of palate in same
region. Posterior palatal ridge much as in Talpa, but the
extremities produced as distinct backward-curved processes.
Mandible rather robust, the ramus nearly straight, the angular
process much below alveolar line Coronoid process high and
narrow, slightly recurved at tip, its height above alveolar line
considerably 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 europsea,
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, theformer 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 antero-internal faces
ridge ; height of shaft
Fig. 6.
Galemys pyrenaicus. Teeth x 5.
marked by a slight though evident
about equal to width of palate ; first 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 forming a straight transverse cutting edge which
GALEMYS 25
acts in opposition to combined posterior surface of large upper
incisors. Upper unicuspids forming an unbroken row continuous
posteriorly with series of cheek teeth, but separated anteriorly
from large incisor by distinct space into which the apex of
second lower incisor fits when jaws are closed. Two anterior
unicuspids (i 2 and i 3 ) 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 1 ) single-rooted, subterete,
scarcely larger than second. Fifth and sixth unicuspids (pm 2
and pm 3 ) 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 pm 2 and pm? but with crowns lower and
longer, slanting a little forward, each with a faintly developed
antero-internal lobule. These teeth are slightly imbricated and
their form approximates that of the unicuspids of the Soricidie.
First and second (i s and c ) 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 4 ) largest of the
series, distinctly two-rooted, its cusp nearly on level with
main cusps of molars, its an tero- 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 developed anterior and posterior
cutting edges, its antero-internal cusp small but evident, its
postero-internal cusp about equal to protocone of m 3 . 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 to 1 , 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
Talpa europsea, but contrast in height of outer and inner
cusps very slight.
26
INSECTIVORA
Galemys pyeenaicus pyeenaicus Geoffroy.
1811. Mygale pyrenaica Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii,
p. 193.
1910. Myogale pyrenmca Trouessart, Paune Mamm. d'Europe, p. 60.
Type locality. — Near Tarbes, Hautes-Pyrenees, France.
Geographical distribution. — Pyrenees and adjacent portion of
southern Prance ; probably also north-eastern Spain to the
Ebro ; Asturias 1
Diagnosis. — Hind foot, 32 • 4 to 34 • 6 mm. ; condylobasal
length of skull about 33 to 34 mm.
Colour. — Back and sides intermediate between prouts-brown
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-buff, clouded by slaty under colour. Front feet dull
ochraceous-buff 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 body, 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,
Specimens examined. — Fifteen, from the following localities :—
Prance : Ax-les-Thermes, Ariege, 5 ; Pyrenees, no exact locality, 8
(B.M. andTJ.S.N.M.).
Spain : Pajares, Leon, 2.
2 al. Ax - les - Thermes, Ariege, Toulouse Museum 1. 7. 27. 1-2.
720 m. Prance. (p).
<5, 9. Ax-les-Thermes, Ariege. V. Builles (p). 8. 3. 27. 4-5.
2. Pyrenees. (Verreaux.) Tomes Collection. 7. 1. 1. 15-16.
1. Unknown. P. Maxwell Lyte(p). 62.1.13.2.
1 al. Pyrenees. (No history.)
1. Pyrenees. Purchased (Parzu- 41. 918.
daki).
1. Pyrenees. Dr. J. E. Gray (p). 43. 10. 14. 1.
£, ?. Pajares, Leon, Spain. O. Thomas (p). 8. 2. 9. 47-48.
(N. Gonzalez.)
Galemys pyeenaicus eufulus Graells.
1897. Myogalea rufula Graells, Mem. Eeal. Acad. Sci., Madrid, xvii, p. 460.
1910. Myogale pyrenaica rufula Trouessart, Paune Mamm. d'Europe, p. 61.
Tyle locality. — Rio Balsain, above the Venta de los Mosquitos,
Sierra de Guadarrama, Segovia, Spain.
Geographical distribution. — Central Spain, south of the Ebro
Valley.
Diagnosis. — Hind foot, 36 to 38 mm. ; condylobasal length
27
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28
INSECTIVORA
of skull about 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 Pyrenean form.
Measurements. — 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 from Silos, province of Burgos, Spain,
and one from Buitrago, province of Madrid (U.S.N. M.).
Remarks. — 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 (Nos. 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 water-shrew, 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 general form to be an active swimmer.
Its defective vision, inherited from the same source, would also
tend to a like result.
2 9 sks. Silos, Burgos ; Spain. Rev. S. Gonzalez (c). 8. 7. 7. 8-9.
6, 2 sks. Silos. G. S. Miller (c). 8. 8. 4. 43-45.
Family SORICID^E.
1821. Soricidie Gray, London Med. Repos., xv, p. 800, 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
rudiment of the zygomatic process 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 produced in axis of mandible,
the other anterior teeth forming a series of small " unicuspids,"
differing from each other chiefly in size ; crowns of upper molars
SORBX 29
low, sub-quadrate 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.
Bemarhs. — 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 OP SORIGIDM.
Posterior lower molar with five cusps ; teeth pigmented
at tips ; tail without sprinkling of elongated hairs.
Upper unicuspid teeth 5-5 ; 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 sprinkling of
elongated hairs.
Upper unicuspid teeth 4-4 Pachyura, p. 81.
Upper unicuspid teeth 3-3 Crocidura, p. 86.
Genus SOREX Linnaeus.
1758. Sorex Linnaeus, Syst. Nat., i, 10th ed., p. 53.
1829. Oxyrhin Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 119 (Included the undeterminable Sorex constrictus Hermann
and S. tetragonurus Hermann; the latter may be chosen as type).
1835. Amphisorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,
ii, p. 23 (hermanni = Neomys fodiens skull -f- Sorex araneus
tetragonurus animal) Part.
1838. Oorsira Gray, Proc. Zool. Soc. London, 1837, p. 123. June 14, 1837
(vulgaris = araneus).
1842. Otisorex De Kay, Zool. of New York, I, Marnm., p. 22 (platyrhinus
1857. Sorex Blasius, Saugethiere Deutschlands, p. 124.
1890. Somalurus Schulze, Schriften Naturwiss. Vereins Harzes in Werni-
gerode, v, p. 28 (alpinus).
Type species. — Sorex araneus Linnaaus.
Geographical distribution. — Northern portion of both hemi-
spheres ; in Europe west to Ireland and south to central Spain
and southern Italy.
30
INSECTIVORA
3-3
1-1'
Characters. — Upper unicuspid teeth 5-5 (dental formula :
c i=I> .P m f=i! m t=j~ 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 which occur in Europe.
KEY TO THE EUROPEAN POEMS 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 2 ; tail about
as long as head and body ; colour uniform dark
slaty grey. (Alpine Shrews) S. alpinus, p. 60.
Condylobasal length of skull 19-4 to 20 - 6 mm.;
upper tooth-row 8'6 to 9'0 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. hereynicm, p. 63.
Anterior mandibular incisor with high, distinct lobes
on cutting edge ; first lower unicuspid single-
pointed ; lachrymal foramen in front of point of
contact between m' and m 2 ; tail shorter than
head and body ; colour never uniform dark slaty
grey, the under parts usually much lighter than
back.
Pirst, second and third upper unicuspids sub-equal ;
condylobasal length of skull 14 • 8 to 16 ■ 6 mm. ;
bead and body usually about 50 to 60 mm.
(Pigmy Shrews) S. minutus, p. 53.
Molars and anterior upper incisor normal (Dis-
tribution general) s. m. minutw, p. 55.
Molars and anterior upper incisor enlarged
(Southern Italy) 5, m . lucanius, p. 60.
Pirst and second upper unicuspid much larger than
third ; condylobasal length of skull 1Y • 4 to
20 mm. ; head and body usually about 65 to
80 mm. (Common Shrews) S. araneus, p. 31.
SOEBX 31
Palate broad anteriorly, its width at level of first
uniouspid nearly equal to that of tooth.
Oondylobasal length of skull 18 • 4 to 19 • 2 mm. ;
anterior teeth enlarged (Island of Jersey) ... S. a. fretalis, p. 45.
Oondylobasal length of skull about 17 '5 mm. ;
anterior teeth not enlarged (Mountains of
central Spain) S. a. granarius, p. 52.
Palate narrow anteriorly, its width at level of
first uniouspid barely more than half that
of tooth.
Oondylobasal 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. tergensis, 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. tetragonurus, p. 42.
Back rarely if ever blackish in summer
pelage (Pyrenees) S. a. pyrenaimis, p. 44.
Oondylobasal 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) 5. u. santonus, 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 Prance) 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. aranew, 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 Linnaeus.
(Synonymy under subspecies.)
Geographical distribution. — Northern portions of Europe and
Asia from England and Scotland eastward ; exact limits of
ran^e unknown ; in western Europe south to central Spain, and
central Italy, north to northern Scandinavia.
Diagnosis. — Size medium, head and body usually about
65-80 mm, the tail 20-35 mm. shorter, oondylobasal length of
skull, 17-8-20 mm.; posterior lobe of anterior upper incisor
32 INSECTIVORA
compressed laterally, 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 4 mm. in summer, 8 mm. in winter, its
texture not specially modified ; no elongated hairs on flanks 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 surface ; 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 apparently 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 from 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, rather closely
appressed, and nearly concealing the annulation ; pencil usually-
well developed, 4-6 mm. in length, but occasionally 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. Mammae :
i 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 santonus. Underparts smoky grey washed
with wood-brown, or occasionally suffused 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 this pattern is often the most convenient
character by which to recognize shrunken ill-prepared specimens,
which might otherwise be mistaken for Sorex minutus.
Skull. — 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 remaining
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 first
confined to lateral portions of occiput, rarely
extending to median line. Dorsal profile
usually with evident concavity in inter-
J . J . Sorex amneus.
orbital region (more marked than in aorex Nat. size.
minutus and S. alpinus). 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 to 1 . 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
diminish 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 equal 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
FIG. 8.
Sorex aran&us. Anterior
teeth in profile. X 5.
and posterior borders sub-equal, slightly longer than anterior
border. Cusp rounded on antero-external face, squarely truncate
postero-internally along 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 unicuspid essentially
similar to first 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 representing 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. Protocone long and rather
^EHZIu
Fie. 9.
Sorex aratieus. Teeth x 10.
SOBEX 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.
Measurements. — 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 mm., 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.
Remarks. — 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.
Sorex araneus araneus Linnseus.
1758. [Sorex] araneus Linnaeus, Syst. Nat., I, 10th ed., p. 53 (Sweden).
1828. Sorex coronatus Millet, Faune de Maine-et-Loire, I, p. 18 (Blou,
Maine-et-Loire, France).
1828. Sorex personatus Millet, Faune de Maine-et-Loire, I, p., 18, foot-
note (Rejected MS. name for coronatus). Not Sorex personatus
I. Geoffroy, 1827.
1832. Sorex concinmts Wagler, Isis, p. 54 (Bavaria).
1832. Sorex rhinolophus Wagler, Isis, p. 54 (Bavaria).
1832. Sorex melanodon Wagler, Isis, p. 54 (Bavaria).
1838. Sorex vulgaris Nathusius, Wiegmann's Archiv fur Naturgesch.
iv, I, p. 45.
1839. S\orex~\ macrotrichus de Selys-Longchamps, Etudes de Micromamm.,
p. 20. (Specimen of S. araneus briefly described as agreeing with
the S. macrotrichus Mehlis MSS. No locality given.)
1839. S\prex~] labiosus Jenyns, Ann. Nat. Hist., n, p. 326, January, 1839.
(Frankfurt a/.\I., Germany.)
1857. Sorex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part).
1895. Sorex araneus Thomas, The Zoologist, 3rd ser., xrx, p. 63, February,
1895.
1910. Sorex araneus Trouessart, Faune Mamm. d'Europe, p. 51 (part).
Type locality. — Upsala, Sweden.
Geographical distribution. — Western Continental Europe,
d 2
36
INSECTIVORA
araneus.
tetragonurus.
•o%
.. 38-7%
22-6%
.. 93-7%
21-2%
.. 88-7%
45-3%
.. 100-0 %
54-6%
■o%
•o%
.. 37-2%
except Atlantic watershed of south-western Norway, from
Finland to central France, central Germany, and northern Hun-
gary. Exact southern and eastern limits of range not known.
Diagnosis. — Size small (condylobasal 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 1 and m 2 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 pigmentation on all of the cusps is less
extensive than in the Alpine and Pyrenean races, the differences
are best seen in the hypocones of the three large upper cheek
teeth and protocone of m 3 , 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 tetragonurus gives the following results : —
Large premolar with pigment on hypocone
First molar with pigment on hypocone .
Second molar with pigment on hypocone
Third molar with pigment on protocone
None of the small cusps pigmented
All of the small cusps pigmented .
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, central Gudbrandsdal, Norway
(dry): hind foot, 12-3 (12 "2-12 '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, 12 • 7 (12 • 2-13 • 0). 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 • 8-12 - 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 Christiania, 4 (U.S.N.M.) ; Holme,
Mandal, 7.
Sweden: Upland, 1; Upsala, 97 (U.S.N.M.); Skaane, 3 (U.S.N.M.).
Denmark: Hillerbd, Zealand, 6; Nystad, Lolland, 3 (U.S.N.M.);
Skansen, Lolland, 1 (U.S.N.M.).
Holland: Oosterbeek, Guelderland, 6; Leiden, 4 (U.S.N.M.).
Belgium: Hastiere, Namur, 1; Waremme, Liege, 10 (U.S.N.M.).
France : Guines, Pas-de-Calais, 4 ; Manonville, Meurthe-et-Moselle, 2 ;
Barbizon, Seine-et-Marne, 3.
SOREX
37
Germany : Brunswick, 35 (B.M. and U.S.N.M.) ; Bodethal, Harz Mts.,
15 (U.S.N.M.) ; Mauseklippe, Harz Mts., 2 (U.S.N.M.) ; Bahrenberg, Harz
Mts., 9 (U.S.N.M.); Tharand, Saxony, 1; Magdeburg, Saxony, 5; Moritz-
burg, Saxony, 1 (U.S.N.M.); Ingelheim, Rheinhessen, 3; Nuremberg,
Bavaria, 10 (U.S.N.M.) ; Marxbeim, near Monheim, Bavaria, 15 ; Strass,
near Burgheim, Bavaria, 1 ; Eulengrund, Riesengebirge, Silesia, 2
(U.S.N.M.) ; Wolfsbau, near Sneekoppe, Riesengebirge, Silesia, 8 (U.S.N.M.);
Niesky, Silesia, 5 ; near Konigsberg, 6 (U.S.N.M.) ; no exact locality, 1.
Austria-Hungary : Haida, Arva, Bohemia, 9.
6,9.
2 <5,2 9.
i.
6, 5 9.
c5al.
9.
4<5,2 9.
i 6, 2 9.
9.
3 6, 1 9.
2 i, 2 9.
6, 3 9, 2.
9.
Sal.
<5,2 9.
3 3, 4 9.
2 6.
i.
3<J,2 9.
2al.
1.
3 £, 4 9.
2.
Molmen, Gudbrandsdal. R. J. Cuninghame
Norway.
Holaaker, Gudbrands-
dal, 1900 ft.
Lesjevark, Gudbrands-
dal.
Holme, Mandal, 200 ft.
Norway.
Upland, Sweden.
(<?. Kolthqff.)
Hillerod, Zealand, 10 m.
Denmark.
Oosterbeek,Guelderland, O. Thomas (c & p
10-15 m. Holland.
Hastiere, Namur, Bel-
gium.
Guines, Pas-de-Calais.
10 m. France.
Barbizon, Seine - et
Marne.
Auerum Forest, Bruns-
wick, Germany.
Tharandt, Saxony.
Magdeburg, Saxony.
R. J. Cuninghame
(Pi-
Miller Collection.
R. J. Cuninghame
Lord Lilford (p).
O. Thomas (c&p).
G. A. Boulenger
(o & p).
O. Thomas (c & p).
G. S. Miller (c).
G. Barrett-Hamil-
ton (o & p).
Lord Lilford (p).
Dr. W. Wolterstorff
Ingelheim, Rheinhessen. C. Hilgert (o).
Marxheim, Bavaria. Lord Lilford (p).
Bayreuth, Bavaria. Miller Collection.
Strass, Burgheim, Ba- Lord Lilford (p).
varia.
Niesky, Silesia, 200 m. Lord Lilford (p).
(W. Baer.)
No exact locality. Zool. Soc. Collec-
tion.
Germany. Stockholm Museum
(■)•
Haida, Bohemia. Lord Lilford (p).
Haida, Bohemia. 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. 6-8.
94. 7. 9. 1.
94. 6. 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. 6-8.
9. 9. 8. 7-13.
7. 7. 7. 2S68-
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.
Sobex aeaneus castaneus Jenyns.
1838. S[orex] tetragonurus var. $ S[prex] 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.
Diagnosis. — Similar to Sorex araneus araneus, 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 frequenth*
seen in the typical race.
Skull and teeth. — The skull and teeth resemble those of true
Sorex araneus.
Measurements. — Average 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 specimens 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, 16 ; 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 ; Stockbriggs, 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 ; Biding Mill-on-
Tyne, Northumberland, 3; Newby Bridge, Lake Windermere, Cumber-
land, 1 ; Grimsby, Lincolnshire, 17 ; Whitnash, Warwickshire, 1 ; Bugby,
Warwickshire, 2 ; Piley, Yorkshire, 3 ; Wellersey Hill, Broadway, Wor-
cestershire, .1 ; West Cheshire, 1 ; Shropshire, 1 ; Staffordshire, 1 ; Swith-
land, Leicestershire, 10 ; Bishopstoke, Herefordshire, 1 ; Leominster, Here-
fordshire, 1 ;• Graffconbury, Herefordshire, 19 ; Lilford, Northamptonshire,
2 ; Drinkstone Park, Bury St. Edmunds, Suffolk, 2 ; Lowestoft, Suffolk, 6 ;
Wormsley, Oxfordshire, 1 ; Stokenchurch, Oxfordshire, A ; Cambridge-
shire, 1 ; Kensington Gardens, London, 1 ; Hillingdon, Middlesex, 3 ;
Bletchingley, Surrey, 4 ; Godalming, Surrey, 2 ; Merstham, Surrey, 1 ;
Bichmond Park, Surrey, 1 ; Crowborough, Sussex, 3 ; St. Leonard's,
Sussex, 1 ; Tunbridge Wells, Sussex, 1 ; Bastwell, Kent, 3 ; Lyndhurst
Boad, Hampshire, 2; New Forest, Hampshire, 17; Basingstoke, Hamp-
shire, 1 ; Alum Bay, Isle of Wight, 3 ; Clifton Bridge, Gloucestershire, 2 ;
Leigh Woods, Clifton, Gloucestershire, 3 ; Blandford, Dorsetshire, 1 ;
Combmartin, Devonshire, 4 ; Chagford, Devonshire, 5 ; Northlew, Devon-
shire, 6 ; no exact locality, 8.
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
S0KEX 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 (bergensis, tetragonurus and pyrenaicus), 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(5. Black Isle, Cromarty. W. R. Ogilvie-Grant 11. 1. 3. 103-
(c & p). 107. 109-112.
2c5. South Sutor, Cromarty. W.R. Ogilvie-Grant 11. 1. 3. 108,
(c & p). 113.
2 (5, 2 9. Nairn, Morayshire. W. R. Ogilvie-Grant 11. 1. 3. 99-102.
(c & p).
6. Morayshire. E. R.Alston (c&p). 79.9.25.79.
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).
<5. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.98.
shire. (c & p).
3 <J, 7 9. Cromlix, Stirlingshire. W. R. Ogilvie-Grant 11.1.3.114-123.
(c & p).
May. H. Russell (c & p). 9. 9. 11. 1-3.
Stockbriggs, Lanark- E. R. Alston (c &p). 79.9.25.11-12.
shire.
Kirtle Bridge, Dumfries- Miss D. Bate (c&p). 11.1.3.88.
shire. 11. 1. 3. 92-94.
Wyseby, Dumfriesshire. Miss D. Bate (c&p). 11.1.3.89-91.
Aberia, Merionethshire, G. H. Caton Haigh 11. 1. 3. 156-
Wales. (c & p). 163.
Bridgend, Glamorgan- R. I. Pocock (c&p). 11. 1. 3. 164-
shire. 169.
Berwick-on-Tweed, J. H. Fryer (c & p). 47.11.11.6-7.
Northumberland,
England.
Newby Bridge, Cumber- J. Paul (c&p). 94. 9. 3. 2-3.
land.
Grimsby, Lincolnshire. G. H. Caton Haigh 11. 1. 3. 124-
(c & p). 131.
Swithland, Leicester- P. A. Butler (c&p). 11. 1. 3. 132-
shire. 138.
Shropshire. T. 0. Eyton (c & p). 63. 10. 12. 4.
Bishopatoke, Hereford- S. O. Ridley (c&p). 84.10.6.1.
shire.
Graftonbury, Hereford- W. de Winton 11. 1. 3. 72-87.
shire. (c&p).
Graftonbury, Hereford- W. de Winton 96. 4. 28. 17-19.
shire. (c&p).
Bury St. Edmunds, J. H. Powell (c & p)'. 80. 5. 22. 1-2.
Suffolk.
Lowestoft, Suffolk. O. Thomas (c & p). 11. 1. 3. 139-
144.
Stokenchurch, Oxford- W. R. Ogilvie-Grant 11. 1. 3. 145-
shire. (c&p). 148.
Hillingdon, Middlesex. Miller Collection. 7. 7. 7. 3595-
(O. Thomas.) 3597.
3al.
6,9.
6, 3 9.
3 9.
5 6,39,
3 6, 3 9.
2al.
lal.
■5,5 9,2.
<5,6 9.
lal.
9al.
8 <5, 8 9.
3 st.
2al.
3=5,3 9.
4 9.
3 9.
40 INSECTIVOEA
1. Kensington Gardens, N. Churton (c & p). 161. a.
London.
4 9. Bletohingley, Surrey. W. R, Ogilvie-Grant 11. 1. 3. 149-
(p). 152.
1. Crowborough, Sussex. Miller Collection. 7. 7. 7. 3593.
(W.R.Ogilvie-Grant.)
Sal. Richmond Park, Surrey. Prof. Owen (p). 75.9.17.1.
7 S, 10 9. NewForest,Hampshire. Miller Collection. 7. 7. 7. 2851-
2867. 3023.
9. Basingstoke, Hamp- Miller Collection. 7. 7. 7. 3611.
shire. (W. P. Stark.)
5,2 9. Alum Bay.Isle of Wight. 0. Thomas (c & p). 11.1.8.153-155.
9 (albino) Winscombe, Somerset. P. A. Knight (o. & p). 4. 8. 9. 1.
6, 4 9. Northlew, Devonshire. P. C. Wroughton 11. 1. 3. 170-
(c & p). 174.
2<J, 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.
SoKEX ARANEUS SANTONUS Mottaz.
1908. Sorex santonus Mottaz, Bull. Soc. Zool. de Geneve, i, p. 118,
April 30, 1908. Type in Mottaz Collection.
1910. Sorex santonus Trouessart, Paune Mamm. d'Europe, p. 54.
Type locality. — Lignieres-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 dark 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-defined
line of demarcation is present. Tail blackish throughout. Feet
scantily clothed with inconspicuous brownish hairs.
Shull and teeth. — The skull 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).
Remarks. — The Charente shrew is so different in aspect from
the other European races of Sorex araneus that it needs no
special comparison with any of them. Its peculiar colour is,
however, almost exactly duplicated by that of a larger, longer -
tailed form from Asia Minor.
41
SOREX ARANEUS EURONOTUS Miller.
1901. Sorex araneus euronotus Miller, Proo. Biol. Soc, Washington, xiv,
p. 44, April 25, 1901 (Montrejeau, Haute-Garonne, Prance). Type
in U.S. National Museum.
1910. Sorex araneus euronotus Trouessart, Faune Mamm. d'Europe, p. 53.
Type locality. — Montrejeau, Haute-Garonne, France.
Geographical distribution. — Probably the plains between the
Pyrenees and the Garonne. At present known from the type
locality only.
Diagnosis. — Similar to Sorex 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. Skull 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 type 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. — Nine, 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.,
m, p. 416, May, 1909. Type in U.S. National Museum.
1910. Sorex araneus bergensis Trouessart, Faune Mamm. d'Europe, p. 52.
Type 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 obscurely
bicolor, brownish above, dull yellowish below.
Skull 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 INSECTIVOKA
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.
Specimens examined. — Twenty-seven, from the following localities : —
Norway: 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.).
BemarJcs. — 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.
E. G. B. Meade 5. 7. 1. 2.
Waldo (c & p).
0. H. Stephenson 8. 1. 5. 1-5.
(c & p).
Miller Collection. 7. 7. 7. 4453-
4456.
Miller Collection. 7. 7. 7. 4457-9.
G. Barrett-Hamil- 8. 9. 21. 1.
ton (p).
Soeex araneus tetragonurtjs Hermann.
1780. Sorex tetragonurus Hermann in Zimmermann, Geogr. Gesch., n,
p. 383 (Strassburg, Germany).
1792. Sorex guadricaudatus Kerr, Anim. Kingd., p. 208 (Strassburg,
Germany). (Based on Pennant's account of S. tetragonurus
Hermann.)
1835. Sorex fodiens Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Stras-
bourg, ii, p. 17. Part : skull, not animal.
1835. Sorex hermanni Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Stras-
bourg, ii, p. 23 (Near Strassburg, Germany). Part: animal, not
skull.
1857. Sorex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part).
1868. ? Sorex araneus, pallidus Fitzinger, Sitzungsber. kais. Akad. Wis-
sensch., Wien, Math. Naturwiss. Classe, lvii, pt. I, p. 488
(Based on specimen from unknown locality, probably in Italy,
figured by Bonaparte, Iconogr. Paun. Ital., fasc. xxix, pi., fig. 5.)
1869. [Sorex vulgaris] var. nuda Patio, Paune Vert. Suisse, I, p. 127
(Bernese Oberland, Switzerland).
1.
Vefsen, Nordland, Nor-
,5,4?.
way.
Skjserdal, Nordfjord.
2(5,2 9.
Opheim, Bergen.
6,2 9.
9.
Bergen.
Bergen, 2,700 ft.
SOEBX 43
1869. [Sorex vulgaris] var. nigra Patio, Faune Vert. Suisse, i, p. 127
(Lucerne, Switzerland).
1900. S[orexr\ vulgaris var. vel subsp. mollis Patio, Bev. Suisse de Zool.,
viii, p. 471 (Substitute for nigra).
1901. Sorex araneus alticola Miller, Proc. Biol. Soc, Washington, xiv,
p. 43, April 25, 1901 (Meiringen, Switzerland). Type in U.S.
National Museum.
1905. S[orex] vulgaris crassicaudatus Patio, Arch. Sci. Phys. et Nat.,
Geneve,' 4th ser., xix, p. 201, February 15, 1905 (Zermatt, Switzer-
land). Cotypes in Geneva Museum.
1905. Crossopus ou Sorex ignotus Patio, 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, Huny ad, Hungary) .
Type in British Museum.
1910. So7-ex araneus tetragonurus and S. araneus carpathicus Trouessart,
Paune Mamm. d'Burope, pp. 52, 54.
Type locality. — Strassburg, Germany.
Geographical distribution. — Alps and neighbouring portions of
Germany, Prance and Italy ; eastward through Tirol to the
mountains of Transylvania.
Diagnosis. — Similar to Sorex araneus bergensis, but colour in
summer pelage not so dark, the contrast between back and
sides often not conspicuous, though back is frequently blackish ;
pigmentation of teeth more extensive than in the northern races,
the hypocone of m 1 and m 2 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 body, 73-1 (70-81); tail, 51-1 (50-55); hind foot, 13-3
(13-14). Measurements of an adult female from Zermatt, Valais,
Switzerland (in alcohol) : * head and body, 74- 6 ; tail, 51 ; hind
foot, 13-6. Average and extremes of three specimens from
Vitznau, Switzerland : f head and body, 67 -3 (62-72); tail,
46-3 (44-49); hind foot, 13-2 (12-8-13-6). Type of 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 : J
head and body, 74-0 (70-77); tail, 52-5 (47-57); hind foot,
13-3 (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.
* Ootype of Sorex vulgaris crassicaudatus Patio,
t Essentially topotypes of Sorex vulgaris nigra and mollis Patio.
X Topotypes of Sorex araneus alticola Miller and essentially topotypes
of Sorex vulgaris nuda Patio.
44
INSBCTIVORA
Specimens examined. — Two hundred and sixty-two, from the following
localities : —
France : Etupes, Doubs, 10 ; Barcelonnette, Basses-Alpes, 3 ; Cha-
monix, Haute-Savoie, 10 (U.S.N.M.).
Germany : Strassburg, 8.
Switzerland : 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.) ;
Andermatt, Uri, 49 (U.S.N.M.) ; Hospenthal, Uri, 1 (U.S.N.M.) ; Zurich, 1 ;
Murgsee region, St. Gallen, 16 (U.S.N.M.) ; Degersheim, St. Gallon, 3
(U.S.N.M.); Uzwil, St. Gallen, 1 (U.S.N.M.); Sitterwald, St. Gallen, 5
(U.S.N.M.); Ziiberwangen, St. Gallen, 5 (U.S.N.M.); Wildkirchli, Appen-
zell, 1 (U.S.N.M.) ; Albulapass, Grisons, 1 (U.S.N.M.) ; Untervatz, Grisons,
5 (U.S.N.M.); Grisons, no exact locality, 4 (U.S.N.M.); Faido, Ticino, 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.
Austria-Hungary : Hatszeg, Hunyad, Transylvania, 10 ; Csall6k8z-
Somorja, Pressburg, Hungary, 3; Schwaz, Tirol, 2 (U.S.N.M.).
Italy : Near Turin, 4 (Turin) ; Unerzio, Cuneo, 1 ; Vallombrosa, near
Florence, 2.
0. Thomas (p). 8. 8. 10. 14-24.
O. Thomas (p). 8. 8. 10. 25-27.
0. Thomas (p). 8. 8. 10. 28-32.
Dr. J. Anderson 91. 10. 15. 26.
(c & p).
0. Thomas (o & p). 5. 8. 2. 32-34.
C. Mosch (p). 89. 11. 8. 3
0. Thomas (c & p). 5. 8. 2. 19.
0. Thomas (o & p). 5. 8. 2. 2-5.
E. B. Alston (p). 79. 9. 25. 10.
C. G. Danford (o). 3. 2. 2. 3-10.
3. 11. 8. 15.
Budapest Museum 94. 3. 1. 22-25.
(«)■
Dr. G. Ceeconi 1. 8. 2. 2-3.
(o & p).
8<5,2 9.
Etupes, Doubs, 350 m.
France. (0. Mottaz).
3 6.
Barcelonnette, Basses-
Alpes. (0. Mottaz)
4 6, 1 9.
Strassburg, Alsace.
(C. Mottaz.)
1 al.
Zermatt, Valais, Swit-
zerland.
3 6.
Vitznau, Lake Lucerne.
1 al.
Zurich.
9.
Faido, Ticino.
2.5,2 9.
Locarno, Ticino.
9.
Switzerland.
8 <J, 1 9.
Hatszeg, Hunyad, Tran-
sylvania, 1500-2000ft.
Hungary.
3.
Csall6koz-Somorja,
Pressburg, 400 ft.
Hungary.
2al.
Vallombrosa, Florence,
Italy.
SOEEX AEANEUS PYRENAICUS Miller.
1909. Sorex araneus pyrenaicus Miller, Ann. and Mag. Nat. Hist., 8th ser.,
in, p. 416, May, 1909. Type in British Museum.
1910. Sorex araneus pyrenaicus Trouessart, Faune Mamm. d'Europe, p. 53.
Type locality. — L'Hospitalet, Ariege, France. Altitude
4,700 feet.
Geographical distribution. — 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
SOEEX 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
body, 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, Pyrenees-Orientales (Spanish watershed), 9 ; l'Hospitalet, Ariege,
12 ; Ax-les-Thermes, Ariege, 2 ; Bareges, Hautes-Pyrenees, 9.
Remarks. — 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.
S, 9. Porte, Pyrenees-Orien- G. S. Miller (o). 8. 8. 4. 142-143.
tales, 1600m. France.
2 <5, 9. Ported Pyrenees-Orien- 0. Thomas (p). 8. 9. 1. 49-51.
tales, 1600-1700 m.
(A. Robert).
S 4 9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.151-154.
4700 ft. 301.
(8. 8. 4. 301. Type of subspecies.)
9. Ax-les-Thermes, Ariege, G. S. Miller (c). 8. 8. i. 141.
2600 ft.
<5 6 9. Bareges, Hautes-Pyre- G. S. Miller (c). 8. 8. 4. 144-150.
nees, 1300-1500 m.
SOEEX AEANEUS FRETALIS Miller.
1909. Sorex arcmeus fretalis Miller, Ann. and Mag. Nat. Hist., 8th ser., in,
p. 416, May, 1909. Type in British Museum.
1910. Sorex araneus fretalis Trouessart, Faune Mamm. d'Burope, p. 52.
Type locality. — Trinity, Jersey, 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
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INSECTIVORA
teeth (incisors both above and below, and first and second upper
unicuspids) enlarged.
Colour. — Summer 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, buffy grey, decidedly lighter than in
any skins of the other races yet examined.
Skull. — In size and general form the skull does not differ
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
breadth is especially noticeable from below, the space between
the anterior unicuspids about equalling diameter of these teeth,
while in all the other races (except S. a. 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 type (adult female) :
head and body, 63; tail, 48-2; hind foot, 13. For cranial
measurements see Table, p. 51.
Specimens examined.— Five, all from the island of Jersey.
Remarks. — The Jersey shrew is easily distinguishable from
the other races by its enlarged anterior teeth. It is probable
that a sufficient series of skins will show that there is an average
difference in colour as well.
?, <$. Jersey, Channel Islands. 0. Thomas (p). 8.9.2.1-2.
(B. H. Bunting.) (8. 9. 2. 1. Type of subspecies.)
SoBEX AEANEUS GRANARIUS Miller.
1910. Sorex araneus granarius Miller, Ann. and Mag. Nat. Hist., 8th ser.,
vi, p. 459, November, 1910. Type in British Museum.
Type locality. — La Granja, Segovia, Spain.
Geographical distribution. — 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 only 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 araneus.
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 portion 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 fretalis. 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 type locality.
Bemarlcs. — In its broad palate the Guadarrama 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.delaEsoalera(c). 6.11.4.3-4.
Spain. (6. 11. 4. 4. Type of subspecies.)
SOREX MINUTUS Linnaeus.
(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.
External 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.
Upper parts between sepia and wood-brown in summer, more
nearly hair-brown in winter, the hairs slate-grey at base and
with faint silvery sub-terminal annulations more visible in some
lights than in others, and giving rise to the greyish effect already
alluded to. Underparts smoke-grey of varying depth, some-
54
INSECTIVOEA
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.
Skull. — Apart from its conspicuously smaller size (condylo-
basal length, 14-8 to 16 mm. instead of 17 "8
to 20 mm.) the skull diners notably from
that of Sorex araneus and S. alpinus in the
narrower, more elongate brain-case, the outline
of which when viewed from above is distinctly
oval instead of sub-circular. Antero-external
portion of border flattened, though less notice-
ably than in Sorex araneus. Owing to its
different form the brain-case is less abruptly
marked off from interorbital region than in
Sorex i. > ;tw>. ^he °tber 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 minutus 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.
When viewed from below the posterior lobe appears, however,
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 Vla ' n '
square. In lateral view the outline is quite S °tSTin ^SSie^x *"
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 cusp, its form when viewed from
* Sometimes equal to fourth or slightly larger.
SDEEX 55
the side thus noticeably different from that of first upper
unicuspid. Second unicuspid, together with other mandibular
teeth, essentially as in S. araneus. Maxillary cheek-teeth as in
S. araneus, except that hypocones are less developed, that on
large premolar obsolete.
Remarks. — 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.
Soeex minutus minutus Linnseus.
1766. [Sorex] minutus Linnseus, Syst. Nat., I, 12th ed., p. 73 (Siberia).
1769. Sorex pygm&us Laxmann, Sibirische Briefe, p. 72 (Barnaul, Tomsk,
Siberia).
1789. {Sorex] exilis Gmelin, Syst. Nat., i, 13th ed., p. 115 (Yenesei Biver,
Siberia).
1806. Sorex canaliculatus Ljungh, Kongl. Vetensk. Akad. Nya Handl.,
xxvii, p. 263 (Lommaryd Vicarage, northern Vedbo district,
Jornkoping, Sweden).
1811. Sorex pygmseus Pallas, Zoogr. Bosso-Asiat., I, p. 134 (Ob and Yenesei
Bivers, Siberia).
1811. Sorex minimus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvn, p. 186
(Accidental renaming of minutus).
1832. Sorex pumilio Wagler, Isis, p. 54 (Bavaria).
1838. S[orex] rusticus Jenyns, Ann. Nat. Hist., i, p. 423, August, 1838
(England).
1838. S[prex] rusticus var. B S[orex] hibernicus Jenyns, Ann. Nat. Hist., i,
p. 423, August, 1838 (Dublin, Ireland).
1844. Sorex pumilus Nilsson. Ofversigt af Kongl. Vetensk.-Akad. Forhandl.,
Stockholm, I, p. 33, March 20, 1844 (North-eastern Skaane,
Sweden).
1857. Sorex pygmmus Blasius, Saugethiere Deutschlands, p. 133.
1895. Sorex minutus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,
1895.
1910. Sorex mimttus Trouessart, Faune Mamm. d'Europe, p. 55.
Type locality. — Vicinity of the Yenesei River, Siberia.
Geographical 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 Man :
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
INSECTIVORA
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 • (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,
Ndrth Uist, Hebrides, 1 ; Barra Island, Hebrides, 1 ; Stockbriggs, Lanark-
shire, 1; Kirtle Bridge, Dumfriesshire, 1; Wyseby, Dumfriesshire, 2.
Wales : Aberia, Merionethshire, 2 ; near Bridgend, 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, 3 ;
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 Gudbrandsdal, 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.).
Prance : Barbizon, Seine-et-Marne, 1 ; Montrejeau, Haute-Garonne, 1
(U.S.N.M.); Ax-les-Thermes, Ariege, 2; l'Hospitalet, Ariege, 3; Porte,
Pyrenees-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 Kbnigsberg, 2 (U.S.N.M.).
Austria-Hungary: CsaU6k8z-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, Moray- G. Denson (c & p). 8. 9. 17. 1.
shire, Scotland.
9,1. Lhanbryde,Elgin,200ft. W. Taylor (c & p). 8.9.3.1-2.
¥. Dunblane, Stirlingshire. W.E. Ogilvie-Grant 8.9.6.1.
(c & p).
1. Aberdeen. Dr. Macgillivray 52. 7. 10. 13.
(c & p).
<J. Dunvegan, Skye. J. S. Elliott (c&p). 8. 9. 4. 1.
<5. Stornoway, Lewis. R. M. D. Hawker 8. 9. 18. 1.
(c & 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
<5. 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.Harmer.)
1. Clifton, Gloucestershire.
(R. I. Pocock.)
6. Lundy Island, Devon-
shire.
2 <5. Chagford, Devonshire,
856ft. (W.P.Stark.)
6, 4 9. Eamsay, Isle of Man.
i. Sulby Glen, Isle of Man.
9. Clonbroch, Galway, Ire-
land.
1 al. Longford.
$ st. Knock, Down.
2 al. Dunkettle, Cork.
2 S, 1 9. Duncannon, Wexford.
(Bev. Dr. Martin.)
1 al. Ireland.
tj. Holaaker, Gudbrandsdal,
1900 ft. Norway.
9. Barbizon, Seine-et-
Marne, France.
2 9. Ax-les-Thermes, Ariege.
i. 1' Hospitalet, Ariege,
4700 m.
(5. 1' Hospitalet, Ariege,
1450 m. (A. Robert.)
9. Porte, Pyrenees-Orien-
tales, 1600 m.
2. Querum Forest, Bruns-
wick, Germany.
1 al. Frankfort - on Main,
Nassau.
9. Niesky, Silesia, 181 m.
(W. Boer.)
6. Marxheim, Bavaria.
1. Csall6k8z-Somorja, Press-
burg, 400 ft. Hungary.
1. Hatszeg, Hunyad, Tran-
sylvania, 2000 -
5500 ft.
3. Hatszeg, Transylvania,
2000-5500 ft.
3 6. Hatszeg, Hunyad, 1500-
2000 ft.
E,R. Alston (c&p).
B.I. Pocock (c&p).
G. Stokes (o&p).
G. H. Caton Haigh
(o & p).
G. H. Caton Haigh
(c & p).
W. de Win ton
(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. F. Witherby
(c & p).
Dr. G. E. Dobson
(c & p).
Hon. N. C. Roths-
child (c&p).
MissO'Keefe(c&p).
G. Barrett-Hamilton
Zoological Society's
Collection.
B. J. Cuninghame
(c & p).
G. S. Miller (c).
V. Builles (c & p).
G. S. Miller (c)\
0. Thomas (p).
G. S. Miller (c).
G. Barrett-Hamilton
Tomes Collection.
Lord Lilford (p).
Lord Lilford (p).
Budapest Museum
(b).
C. G. Danford (c).
C. G. Danford (c).
C. G. Danford (c).
79. 9. 25. 13.
8. 9. 5. 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. i46.
94. 3. 1. 25.
3. 2. 2. 11.
3. 2. 2. 13-15.
3. 11. 8. 16-18.
58
INSECTIVORA
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60
INSECTIVORA
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 minutus lucanius Miller.
1909. Sorex minutus lucanius Miller, Anrj. and Mag. Nat. Hist., 8th ser.,
in, p. 417, May, 1909. Type in British Museum.
1910. Sorex minutus lucanius Trouessart, Fauna Mamm. d'Europe, p. 55.
Type locality. — Monte Sirino, Lagonegro, Italy.
Geographical distribution. — At present known from the type
locality only.
Diagnosis. — Similar to Sorex minutus minutus, 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
lucanius 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 eye the greater 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. Monte Sirino, Lagonegro, O. Thomas (p). 8. 9. 1. 5.
S. Italy. (A. Robert.) {Type of subspecies.)
SOREX ALPINUS Schinz.
(Synonymy under subspecies.)
Geographical distribution. — Pyrenees, Alps, Harz Mountains,
JRiesengebirge and Carpathians.
Diagnosis. — Size essentially as in Sorex araneus (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 anterior 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 depth 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 very
SOEEX
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.
Shull. — 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 slightly
larger than in S. araneus. Lachrymal fora-
men opening over point of contact between
first and second molars.
Teeth. — Relatively 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. minutus, 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 differs 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
INSBCTIVORA
are all longer in proportion to their height than in Sorex
araneus, and the posterior border 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-
lp\ 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
"Pin 1^
peculiarities of form. The first, how-
Sorex alpinus. Anterior L . . ., . , ,.~, . e ,,
teeth, x s. ever, is strikingly dinerent from 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 different 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. Sohweiz. Gesellsch.
Naturwiss., Neuchatel, I, p. 13 (St. Gothard Pass, Switzerland).
1840. ? Sorex antinorii Bonaparte, Iconogr. Faun. Ital., I, fasc. 29 (No exact
locality, and probably not a European species).
1857. Sorex alpinus Blasius, Saugethiere Deutschlands, p. 126.
1870. ? Sorex intermedins Cornalia, Catal. Descrit. Mamm. Ital., p. 27 (Hills
of Brianza, Como, Italy). Part : body ; see Sordelli, Atti Soc. Ital.
Sci. Nat. e del Mus. Civ. Stor. Nat., Milano, xxxvm, p. 364, 1899.
1899. ? Sorex alpinus var. longobarda Sordelli, Atti Soc. Ital. Sci. Nat. e del
Mus. Civ. Stor. Nat., Milano, xxxvm, p. 363 (MS. synonym of
intermedins).
1910. Sorex alpinus Trouessart, Faune Mamm. d'Europe, p. 50.
Type locality. — St. Gothard Pass, Uri, Switzerland.
Geographical distribution. — From the Jura and Alps through
Tirol to Transylvania ; Pyrenees.f
* In an adult female from Briinig, Switzerland (No. 85830, U.S.N.M.),
the third and fourth unicuspids on right side are fused into a single
2-cusped tooth, while those on left side are normal.
t I have seen no Pyrenean specimens of Sorex alpinus. For record of
its occurrence see Trutat, Bull. Soc. Hist. Nat., Toulouse, xn, p. 100, 1878
("massif de la Maladetta"). The animal is probably less common in the
Pyrenees than in the Alps, as I was unable to find it in several localities
resembling those where it regularly occurs in Switzerland.
SOEEX 63
Diagnosis. — 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 : —
Ebance : Chamonix, Haute-Savoie, 1 (U.S.N.M.).
Switzerland : St. Gerguee, 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-Hungaby : Hatszeg, Hunyad, Transylvania, 4.
i. Vitznau, 440 m. Swit- 0. Thomas (c & p). 5. 8. 3. 15.
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. 13.
46. 2. 13. 14.
1. St. Gothard, Uri. Purchased (Stock- 46. 6. 2. 32.
holm Museum).
1. Switzerland. E. B. 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.
6, 2 ?. Hatszeg, Hunyad, Tran- 0. G. Danford (c). 3. 11. 8. 12-14.
sylvania, 1500-2000 ft.
SOEEX ALPINUS HBHCYNICUS Miller.
1909. Sorex alpinus hercynicus Miller, Ann. and Mag. Nat. Hist., 8th ser.,
in, p. 417, May, 1909. Type in U.S. National Museum.
1910. Sorex alpinus liercynicus Trouessart, Eaune Mamm. d'Europe, p. 51.
Type locality. — Mauseklippe, Bode Valley, Harz Mountains,
Germany.
Geographical distribution. — Harz Mountains and Riesen-
Diagnosis. — Skull and teeth not so large as in typical
Sorex 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
INSBCTIVORA
Observations.
Teeth slightly worn.
„ not worn.
, slightly worn.
„ not worn.
»j jj
jj »j
>J J!
slightly worn.
„ not worn.
jj jj
jj »
j jj
jj jj
jj jj
„ slightly worn.
j j>
„ not worn.
„ slightly worn.
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NEOMYS 65
(72-77); tail, 66-6 (65-68); hind foot, 15- 1 (14-8-15-8).
Average and extremes of five specimens from the Riesengebirge :
head and body, 73-4 (72-77); tail, 65-4 (59-68); hind foot,
14-9 (14-8-15-0). For cranial measurements see Table, p. 64.
Specimens examined. — Eleven (all in U.S.N.M.), from the following
localities in Germany: Bahrenberg, Harz Mts., 2; Bodethal, Harz
Mts., 1 ; Mauseklippe, Harz Mts., 3 ; Eulengrund, Riesengebirge, Silesia, 3 ;
Wolfshau, Eiesengebirge, Silesia, 2.
Remarks. — In external measurements the Alpine shrew of
the Harz Mountains 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. Neomys Kaup, Entw.-Gesoh. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 117 (Sorex daubentonii Erxleben).
1829. Leucorhynchus Kaup, Entw.-Gesch. u.. Natiirl. Syst. Europ. Thier-
welt, i, p. 117 (Sorex lineatus Geoffroy and S. leucodon Hermann ;
the first chosen as type by Thomas, The Zoologist, 4th ser., n,
p. 102, March, 1898).
1829. Hydrogale Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,
p. 119 (Sorex remifer Geoffroy).
1832. Crossopus Wagler, Isis, p. 275 (Sorex fodiens Schreber).
1835. Hydrosorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,
ii, p. 19 (Sorex fodiens Pallas, see p. 17).
1835. Amphisorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,
ii, p. 23 (hermanni = fodiens skull + Sorex araneus tetragonurus
animal). Part.
1838. Pinalia Gray, Proc. Zool. Soc, London (1837), p. 126, June 14, 1838
(MS. synonym of Crossopus).
1848. Galemys Pomel, Archiv Sci. Phys. et Nat., Geneve, ix, p. 249,
November, 1848 (part, included Brachysorex Duvernoy, Crossopus
Wagler, and Pachyura de Selys-Longchamps).
1854. Myosictis Pomel, Catal. Meth. Vert. Poss. Bassin de la Loire, p. 14
(based on a fossil not distinguished specifically from Neomys
fodiens).
1857. Crossopus Blasius, Saugethiere Deutschlands, p. 119.
1898. Neomys Thomas, Zoologist, 4th ser., n, p. 100, March 15, 1898.
Type species. — Sorex daubentonii Erxleben =S. fodiens Schreber.
Geographical distribution. — Palsearctic region north of the
Mediterranean from Spain and England to the Caucasus and
Asia Minor, north to northern Scandinavia.
Characters. — Upper unicuspid teeth 4-4, their form more pre-
hensory than in Sorex; dental formula : i £=?, eg, pm ?=?, m |=| = 30 ;
posterior lobe of anterior upper incisor less than half as high
as first ; anterior lower incisor with one ill-defined lobe on cutting
edge near middle of tooth ; third lower molar and second lower
unicuspid as in Sorex ; points of all the teeth pigmented ; fur
unusually soft and dense, entirely concealing ears ; feet with a
noticeable fringe of elongated hairs at edges of soles and toes,
66
INSEOTIVOEA
the hind feet enlarged and turned somewhat outward ; in one
species a longitudinal ridge of stiffened 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.
Remarks. — 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 Sorex.
They retain, however, the primitive form of the third lower
molar and lower premolar characteristic of Sorex, these teeth
showing no tendency to the more highly modified structure
found in Crocidura. The members of the genus Neomys are the
only European shrews specially adapted to aquatic life. They
are at once recognizable by 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 European shrew.
KEY TO THE EUROPEAN FORMS OP NEOMYS.
Tail with median keel of stiffened hairs extending
entire length of under surface N. fodiens, p. 66.
Underparts whitish or yellowish, occasionally
brownish (Continental Europe except Iberian
Peninsula) X. f. fodiens, p. 69.
Underparts usually brownish (England and Scot-
land) X.f- bicolor, 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) N. anomalus, p. 81.
NEOMYS fodiens Schreber.
(Synonymy under subspecies.)
Geographical distribution. — From Norway and England to the
Pyrenees, northern Italy and western Siberia. Northern and
eastern limits of range not known.
Diagnosis. — Tail with keel always present ; feet very con-
spicuously fringed; hind foot usually more than 17 mm. ; lachry-
mal foramen opening over posterior half of m 1 ; anterior upper
incisor robust.
External characters. — Fur very dense, its depth at middle of
back about 6 mm. in summer, 9 mm. in winter, its texture soft
and velvety, this particularly 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 Sorex araneus, the median ridge on
upper lip with distinct wart-like outgrowth which fits between
points of upper incisors when mouth is closed. Feet larger than
NEOMYS 67
in species of Sorex of approximately the same size, noticeably
broadened ; digits proportioned as in Sorex araneus but gradua-
tion less ; soles and palms completely bare, their surface strongly
tuberculo-reticulate ; pads 6-6, much as in Sorex araneus but
more widely spaced and better defined. At edge of soles and
toes the hairs are elongated to form a dense, conspicuous fringe.
Tail terete or slightly four-sided, its hairs minute, flattened, and
closely appressed, nearly concealing the rather indistinct annula-
tions, of which there are about 25 to the centimeter at middle ;
pencil evident, usually about 3 to 5 mm. in length. Along
median line of tail below, from extreme base to tip, the hairs
are so crowded, elongated, and directed inward as to form a
distinct, low keel, the presence of which is often made more
evident by the nearly bare condition of the skin immediately
at each side of it. Mammas : a 2-2, i 2-2 = 8.
Colour. — Entire underparts a dark slaty grey, more bluish
in summer, more blackish in winter, the individual hairs blackish-
slate with seal-brown tips and a sub-terminal lighter area of
varying distinctness. Underparts usually whitish in strong
contrast with back, the line of demarcation everywhere evident,
and on head passing just below base of ear and extending along
upper lip to nostril. The underparts, while never so dark as
back, are often strongly suffused with bufly, smoky, or wood-
brown, and occasionally tinged with salmon-colour. A small
whitish tuft usually springs from inner surface of ear, and a
minute speck of the same colour is generally present behind eye.
Feet thinly sprinkled with silvery grey hairs. Tail a dark
indefinite brown above, slightly less dark below, the keel usually
a silver grey, sometimes standing out in strong contrast against
the dark surrounding parts.
Skull. — The skull differs from that of Sorex araneus in larger
size, relatively 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 in outline, though distinctly flattened
antero-externally and postero-externally ; in
front it is marked off from interorbital
region by a conspicuous, well defined angle.
In lateral view the profile of brain-case is
high and rounded posteriorly, low anteriorly
where it passes by a distinct, often conspicu-
ously abrupt curve into the nearly flat dorsal
outline of interorbital region and rostrum.
Anteorbital foramen relatively larger and more l °' t j-
widely open than in the European species of .Nat. size.
Sorex. Lachrymal foramen over posterior
half of m 1 . Mesopterygoid space decidedly more than half as
wide as long ; hamular processes shorter than in Sorex araneus
f 2
68
INSECTIVOEA
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 particularly 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-
teriorly and terminating in a slight
though evident point. In 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
Xeomys fodiens. Anterior teeth i , ,. 2 . - n L .
in profile, x 5. between tips 01 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 postero-internal fourth of crown. The con
cave area of cusp 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 araneus araneus, 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, 19'6 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.
Bemarhs. — 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 greatly 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 aquations P. L. S. Muller, Natursyst. Suppl. u. Regist.-Band,
p. 36 (Prance ; based on Buffon, vin, pi. 11). Not Sorex aguaticus
Linnaeus, 1758.
1777. Sorex fodiens Schreber, Saugthiere, in, p. 571 (Berlin, Germany).
1777. [Sorex] daubentonii Erxleben, Syst. Regni Anim., i, p. 124 (Burgundy,
France ; based primarily on the Musaraigne d'eau of Daubenton,
Hist, de l'Acad. Boy. des Sci., Paris, 1756, p. 42).
1780. Sorex carinatus Hermann in Zimmermann, Geogr. Gesch., II, p. 382
(Strassburg, Germany).
1792. Sorex liricaudatus Kerr, Anim. Kingd., p. 208 (Strassburg, Germany ;
based on Pennant's account of Sorex carinatus Hermann).
1793. [Sorex] fluviatilis Bechstein, Gemeinn. Naturgesch. Deutschlands,
in, 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[orex] f[pdiens~] albus Bechstein, Thomas Pennant's Allgem. Uebers.
vierfiiss. Thiere, n, p. 723.
1811. Sorex hydrophilus Pallas, Zoogr. Eosso-Asiat., p. 130 (Berlin,
Germany).
1811. Sorex lineatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvn, p. 181
(Paris, Prance). Type in Paris Museum.
1811. Sorex remifer Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 182
(Abbeville, Somme, Prance). Type in Paris Museum.
1818. Sorex collaris Desmarest, Nouv. Diet. d'Hist. Nat., xxn, p. 65
(Holland: islands at mouth of Escaut and Meuse). Described
but not named by Geoffroy, Mem. Mus. d'Hist. Nat., Paris, i,
p. 309, 1815.
* In an adult male from Geneva, Switzerland (No. 1046 Mottaz), the
right upper incisor and first unicuspid and right lower incisor are white
throughout. All other teeth normal.
70
1NSECTIVORA
1822. Sorex macrourus Lehmann, Observ. Zoologies in faunam Hamburg-
ensem,i,p. 5 (Saehsenwald, near Friedrichsruh, Sohleswig-Holstein,
Germany).
1826. Sorex amphibius Brehm, Ornis, n, p. 44 (Renthendorf, Tbiiringen,
Germany).
1826. Sorex natans Brebm, Ornis, II, p. 44 (Benthendorf, Tbiiringen,
Germany).
1826. Sorex stagnatilis Brehm, Ornis, n, p. 47 (Renthendorf, Thiiringen,
Germany) .
1830. Sorex rivalis Brehm, Isis, p. 1128 (Renthendorf, Thiiringen, Germany).
1832. Sorex musculus Wagler, Isis, p. 54 (Bavaria, Germany).
1832. Sorex psilurus Wagler, Isis, p. 54 (Bavaria, Germany).
1834. S[orex] nigripes Melchior, Den Danske Stats og Norges Pattedyr,
p. 68 (Sielland, Denmark).
1835. Sorex fodiens Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,
II, p. 17. Part : animal, not skull.
1835. Sorex hermanni Duvernoy, Mem. Soo. du Mus. d'Hist. Nat., Stras-
bourg, ii, p. 23 (Strassburg, Germany). Part : skull, not animal.
1838. Amphisorex linneana Gray, Ann. Nat. Hist., n, p. 287, December,
1838 (North Bothnia, Sweden).
1839. [ Sorex fodiens] var. leucotis de Selys-Longohamps, Etudes de Mioro-
mamm., p. 142 (described on p. 25) (St. Gervais, at foot of Mt.
Blanc, Haute-Savoie, France).
1839. ? [Sorex fodiens] var. albiventris de Selys-Longchamps, Etudes de
Micromamm., p. 142 (nomen nudum).
1845. ? [Sorex fodiens] var. nigricans Nilsson, Atti della sesta Riunione
degli Scienziati Italiani, Torino, 1844, p. 357 (Sweden). Nomen
nudum.
1857. Crossopus fodiens Blasius, SiLugethiere Deutschlands, p. 120 (part) .
1868. Sorex fimbriatus and Crossopus fimbriatus Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Clasre, lvii, pt. 1,
p. 610 (Synonyms of " Crossopus daubentonii nigripes," wrongly
attributed to Giebel, Saugethiere, p. 899). Not Sorex fimbriatus
Wagler, 1832.
1868. Crossopus ciliatus, griseogularis Pitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lvii, pt. 1, p. 623
(Chartres, Eure-et-Loire, Prance).
1870. Sorex intermedius Cornalia, Catal. Descrit. Mamm. Ital., p. 27 (Hills
of Brianza, Como, Italy). Part: tail only (see Sordelli, Atti
Soc. Ital. Sci. Nat. e del Mus. Civ. Stor. Nat., Milano, xxxvin,
p. 364, 1899).
1895. Neomys fodiens Thomas, The Zoologist, 4th ser., n, p. 100, March,
1908.
1899. Sorex alpinus var. longobardus Sordelli, Atti Soc. Ital. Sci. Nat. e
del Mus. Civ. Stor. Nat., Milano, xxxvin, p. 363 (MS. synonym
of intermedius).
1901. Neomys fodiens minor Miller, Proc. Biol. Soc, Washington, xiv,
p. 45, April 25, 1901 (Montrejeau, Haute-Garonne, Prance). Type
in U.S. National Museum.
1905. Crossopus ou Sorex ignotus Patio, Arch. Sci. Phys. et Nat., Geneve,
4th ser., xix, p. 202, February 15, 1905 (Switzerland). Part:
skull, not mandible. Type in Geneva Museum.
1905. Neomys fodiens naias Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., xv, p. 507, May, 1905 (Hatszeg, Hunyad, Hungary). Type
in British Museum.
NEOMYS 71
1906. Neomys fodiens nanus Lydekker, Zoological Record, xlu (1905),
Mamm., p. 34, August, 1906 (Accidental renaming of naias).
1910. Neomys fodiens, N. fodiens naias and N. fodiens minor Trouessart,
Paune Mamm. d'Europe, pp. 56, 57.
Type locality. — Berlin, Germany.
Geographical distribution. — Continental Europe from Norway
to Italy and the Pyrenees.
Diagnosis. — Underparts buffy 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, 18 '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, 61 (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, Hunyad, 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 : Guinea, Pas-de-Oalais, 2 ; Abbeville, So rime, 2 (B.M. and Paris ;
the latter type of remifer Geoffroy) ; near Paris, 1 (Paris ; type of lineatus
Geoffroy) ; Dinan, C6tes-du-Nord, 1 ; Cadillae-sur-Garonne, Gironde, 2
(U.S.N.M.) : Montrejeau, Haute-Garonne, 3 (U.S.N.M.) ; Porte, Pyrenees-
Orientales, 5 ; l'Hospitalet, Ariege, 2 ; Luchon, Haute-Garonne, 7 ; Bareges,
Hautes-Pyrenees, 3 ; Barcelonnette, Basses-Alpes, 1 ; Chamonix, Haute-
Savoie, 1 (U.S.N.M.) ; Cranves-Sales, Haute-Savoie, 1 ; Scientrier, Haute-
Savoie, 1 (Mottaz) ; Etupes, Doubs, 6 (Mottaz) ; no exact locality, 1.
Germany : Brunswick, 10 (U.S.N.M.); Saxony, no exact locality, 1
(U.S.N.M.); 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) ; Ohesieres, Vaud, 4 (Mottaz) ; Les
Plans, Vaud, 3 (U.S.N.M.) ; Meiringen, Bern, 9 (U.S.N.M.) ; Grindelwald,
* " Neomys fodiens minor " Miller.
f "Neomys fodiens naias " Barrett-Hamilton.
I .i INSECTIVORA
Bern, 1 (U.S.N.M.) ; Miirren, 2 ; Gosehenen, 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. Gallon, 2 (U.S.N.M.); Murgsee region, St. GaUen, 5
(U.S.N.M.) ; Au, St. GaUen, 1 (U.S.N.M.) ; Uzwil, St. GaUen, 1 (U.S.N.M.) ;
Sitterwald, St. GaUen, 1 (U.S.N.M.) ; Faido, Tioino, 1 ; Gordola, Locarno,
Tioino, 3 ; Porlezza, Tioino, 1 (Mottaz) ; Muzzano, Tioino, 1 (Mottaz) ; no
exact locality, 1.
Italy : Busalla, Liguria, 3 ; Vallombrosa, Florence, 1.
Bemarhs. — 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. While much
material has been examined it is not wholly satisfactory, con-
sisting chiefly of small lots taken b}' many different 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
race.
1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 1.
way. Waldo (c & p).
S. Kvikne, Hedemarken, G. Barrett-Hamilton 11. 1. 2. 88.
2200 ft. (N. F. Tice- (p).
hurst.)
9. Brekkebygden, Trond- G. Barrett-Hamilton 11. 1. 2. 89.
b/jem (N. F. Ticehttrst.) (p) .
1. Bothnia, Sweden. Purchased. (Wahl- 38. 9. 24. 14.
berg.)
9. Medstugan, Jemtland, Lord Lilford (p). 11. 1. 1. 147.
550 m. (G.Kolthoff.)
2 9. Guines, Pas-de-Calais, 0. Thomas (c & p). 94. 6. 6. 1-2.
France.
<J, 2 9. Porte, Pyrenees-Orion- 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.
<5.
L'Hospitalet, Ariege,
1450 m. (A. Bobert.)
?.
L'Hospitalet, Ariege.
<J,2. 3 9.
Luchon.Haute-Garonne,
600-900m. (A.Robert.)
2<5.
Bareges, Hautes-Pyre-
nees, 1300-1500 m.
9.
Baroelonnette, Basses-
Alpes. (C. Mottaz.)
1.
Cranves-Sales, Haute-
Savoie.
1.
Abbeville, Somme.
1. -
Frankfort, Wiesbaden,
Germany.
9.
Niesky, Silesia, 181 m.
(W. Baer.)
1.
Strassburg, Alsace.
(O. Mottaz.)
2 9.
Haida, Bohemia,
Austria.
1.
Hatszeg, Hunyad, Tran-
sylvania, 1500 ft.
Hungary.
6<J,49.
Hatszeg, Transylvania,
1500-2000 ft.
2.
Miirren, Bern, Switzer-
land.
4 9.
Ziiberwangen, St.Gallen.
(E. Zollikofer.)
9.
Faido, Ticino.
3 6.
Locarno, Ticino.
].
Switzerland.
(J, 2 9.
Busalla, Liguria, Italy.
lal.
Vallombrosa, Florence.
NEOMYS
0. Thomas (p).
G. S. Miller (c).
0. Thomas (p).
G. S. Miller (c).
0. Thomas (p).
A. Robert (p).
V. Baillon (c).
Dr. Dieffenbach (p).
Lord Lilford (p).
0. Thomas (p).
Lord Lilford (p).
G. G. Danford (c).
0. G. Danford (c).
W. Gurtner (p).
0. Thomas (p).
0. Thomas (c & p).
0. Thomas (c & p).
Purchased (Parrey s) .
0. Thomas (o & p).
Dr. G. Cecconi (p).
73
8. 9. 1. 42.
8. 8. 4. 139.
6. 4. 1. 12. 14-
18.
8. 8. 4. 136-
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 bicoloe Shaw.
1791. Sorex bicolor Shaw, Naturalists' Miscellany, n, pi. 55, named in
index (Oxford, England).
1805. Sorex ciliatus Sowerby, British Miscellany, xlix, p. 103 (Norfolk,
England).
1837. Amphisorexpennanti Gray, Proc. Zopl. Soc, London, p. 125 (England).
1840. Crossopus sowerbyi iBonaparte, Iconogr. Faun. Ital., i, fasc. 29, in
text under C. fodiens.
1857. Crossopus fodiens Blasius, Saugethiere Deutschlands, p. 120 (part).
1905. Neomys fodiens ciliatus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., xv, p. 508, May, 1905.
1910. Neomys fodiens ciliatus Trouessart, Faune Mamm. d'Europe, p. 56.
Type locality. — Oxford, England.
Oeographical distribution. — Great Britain.
Diagnosis. — Similar to Neomys 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 Gortachy, Forfar) : head
74
INSEOTIVORA
and body, 83 and 71 ; tail, 57 ■ 5 and 53 ■ 5 ; hind foot, 18 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 fool;, 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) ; Cortaohy, Elgin, 1 (Wilson) ; Cromlix, Stirling-
shire, 1 ; Knibruok, Stirlingshire, 1 ; Aberdeen, 1 ; Penkill, Ayrshire, 1.
Wales : Fishguard, Pembrokeshire, 1 ; no exact locality, 1.
Enqland : Mill-on-Tyne, Northumberland, 1 ; Leeds, Yorkshire, 1 ;
Wellington, Somerset, 1 ; Great Grimsby, Lincolnshire, 2 ; Shaftesbury,
Dorset, 1; Halesworth, Suffolk, 1; Cambridgeshire, 2; Thornhaugh,
Northampton, 1 ; Podington, 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 ; BasiDgstoke, Hampshire, 2 ;
Hampshire, no exact locality, 2 ; Yalding, Kent, 1.
6.
Dunphail, Elgin.
Miller Collection.
7. 7. 7. 2848.
lal. 9
Aberdeen, Scotland.
Hon. N. C. Roths-
child (p).
10. 11. 28. 1.
1.
Penkill, Ayrshire.
E. R. Alston (p).
79. 9. 25. 8.
9.
Fishguard, Pembroke-
shire, Wales.
Y. H. Mills (c & p).
11. 1. 3. 376.
1.
Wales.
S. Stokes (c & p).
48. 9. 24. 4.
9.
Mill-on-Tyne, North-
Bev. H. H. Slater
11. 1. 3. 375.
umberland, England.
(c & P).
lal.
Wellingborough, North-
amptonshire.
R. R. Orlebar (c&p).
84. 4. 21. 1.
1.
Halesworth, Suffolk.
Bev. W. R. Tate
(c & p).
87. 6. 7. 1.
9.
Cambridgeshire.
3. Baker (c&p).
E. B. Alston (p).
39. 9. 29. 26.
1.
Cambridgeshire.
79. 9. 25. 7.
(J. Baker.)
1.
Epping, Essex.
J. Baker (p).
40. 4. 11. 9.
2al.
Henley, Oxfordshire.
W. Boyal Dawson
(c & p).
11. 1. 3. 377.
lal.
Bury St. Edmunds,
Suffolk. »
J. H.Powell (c&p).
83. 8. 8. 1.
3al.
Bury St. Edmunds,
Suffolk.
Duncan Parker (p).
84. 5. 16. 1-3.
cSal.
Esher, Surrey.
E.Dalgleish(c&p).
11. 1. 3. 378.
9al.
Godalming, Surrey.
W. T. Blanford
(c & P).
94. 8. 5. 2.
<5,3 9.
New Forest, Hampshire.
Miller Collection.
7. 7. 7. 2845-
2849.
2 9.
Basingstoke, Hampshire.
Miller Collection.
7. 7. 7. 4470-
4471.
2al.
Hampshire.
E. Bartlett (p).
74. 11. 24. 1-2.
75
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78
INSECTIVOKA
NEOMYS MILLERI Mottaz.
1907. Neomys milleri Mottaz, Mem. Soo. Zool. de France, xx, p. 22,
September 20, 1907.
1910. Neomys milleri Trouessart, Faune Mamm. d'Europe, p. 58.
Type locality. — Chesieres, Alpes Vaudoises, Switzerland.
Altitude, 1,230 m.
Geographical distribution. — Pyrenees and Alps ; Hungary.
Details of distribution not yet known.
Diagnosis. — Slightly smaller than Neomys fodiens ; tail with
keel absent, or 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 1 and m 2 ; anterior upper incisor
slender.
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 Neomys fodiens. Mammse : a 1—1,
i 4-4 = 10.
Colour.' — The oolour is similar to that of typical specimens of
Neomys fodiens fodiens. Among the skins examined none shows
any noticeable wash of brown 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 m 2 . 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, 76; tail, 59; hind foot, 16; hind foot, including claws,
17. Average and extremes of ten specimens from the type
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,
NEOMTS 79
France: head and body, 78 (75-80); tail, 50-6 (47-53); hind
foot, 15 - 7 (15-4—16). Adult male from Bareges, Hautes-
Pyrenees : head and body, 76; tail, 56; hind foot, 16 '4. For
cranical measurements see Table, p. 80.
Specimens examined. — Thirty-three, from the following localities : —
Switzerland : Near Geneva, 1 ; Lausanne, Vaud, 2 (U.S.N. II.) ;
Chesieres, Alpea Vaudoises, 10 (B.M. and Mottaz) ; Meiringen, Bern, 2
(U.S.N.M.) ; Untervatz, Grisons, 6 (U.S.N.M.) ; Zuberwangen, St. Gallen, 3
(B.M. and U.S.N.M.) ; Sorengo, Ticino, 1 (U.S.N.M.) ; Locarno, Ticino, 2.
Italy : Porlezza, Como, 1 (Mottaz).
Austbia-Hungaby : Zuberec, Northern Hungary, 1.
Frakce : L'Hospitalet, Ariege, 3 ; Luchon, Haute-Garonne, 1 ; Bareges,
Hautes-Pyrenees, 1.
Remarks. — Though superficially resembling Neomys fodiens
the round-tailed water-shrew is easily recognizable by the
complete absence of any true keel on the tail. The slight
elongation of the hairs in median region which sometimes occurs
on terminal third of tail never results in the formation of a
definite keel similar to that of the related species ; while no
trace of such a structure is ever present on basal half of tail.
The smaller hind foot is usually diagnostic, though in this
character there is a slight overlapping between the two species.
Though in all essential respects true water-shrews, the members
of the round-tailed group (N. anomalus of Spain, N. milleri of
the Pyrenees, Alps and northern Hungary, and N. teres of
Asia Minor) are less highly specialized than Neomys fodiens.
Their habits, however, appear to be essentially similar to those
of the common species, as the two animals are found together in
localities where their ranges coincide.*
2 9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.138,140.
Prance.
9. Chesieres, Alpes Vau- C. Mottaz (p). 6. 2. 6. 4.
doises, 1200 m. Swit-
zerland.
9. Ziiberwangen.St.Gallen, O. Thomas (p). 4. 4. 5. 52.
Switzerland.
(E. H. Zollikofer.)
2 1. Locarno, Ticino, Swit- O. Thomas (c & p). 5. 8. 2. 6-7.
zerland.
1. Zuberec, Hungary. Budapest Museum 94. 3. 1. 26.
(e).
* At two localities in the Pyrenees (l'Hospitalet, Ariege and Bareges,
Hautes-Pyrenees), where I found the two animals occurring together, both
were taken alternately in the same traps set at the edge of mountain
streams. In the original description of the species, however, Mr. Mottaz
observes (p. 23) that according to his observations the round-tailed animal
is the more inclined to wander away from the immediate vicinity of water.
The habits of N. anomalus in North Central Spain appear to be as strictly
aquatic as those of N. fodiens.
80
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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 Soo. Espafi. Hist. Nat., Madrid,
vii, p. 224. Published November, 1907.
1910. Neomys fodiens anomalus Trouessart, Eaune Mamm. d'Europe, p. 57.
Type locality. — San Martin de la Vega, Madrid, Spain.
Geographical 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.).
Mammse : 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 specimens 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. — Twenty-two, from the following localities in
Spain : Silos, Burgos, 9 ; La Granja, Segovia, 2 ; Barracas, Castellon, 2 ;
Lerida, 9.
"Remarks. — 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 S, 7 ?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 22-30.
2 al. La Granja, Segovia. M. delaEscalera (c). 8.7.30.6-7.
6,9. Barracas, Castellon. O. Thomas (p). 8.2.9.41-42.
(JV. Gonzales.)
Genus PACHYURA de Selys-Longchamps.
1839. Pachyura de Selys-Longchamps, Etudes de Micromamm., p. 32
(Sorex etruscus Savi). Sub-genus of Crocidura.
1857. Pachyura Blasius, Saugethiere Deutschlands, p. 147 (Sub-genus of
Crocidura).
1897. Plerodus Schulze, Mamm. Europ., in Helios, Abhandl. n. Vortraga
Gesammtb. Naturwiss., xiv, p. 90 (Crocidura suaveolens Blasius
= Sorex etruscus 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 j=J, pm ^, »y= 30.
Remarks. — Although perhaps not forming a natural group,
the 30-toothed species of Crocidura may for convenience be
6
82
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PACHYURA 83
treated as members of a distinct genus. The only European
species is immediately recognizable by its excessively small size.
PACHYURA ETRUSCA Savi.
1822. Sorex etruscus Savi, Nuovo Giorn. de' Letterati, Pisa, I, v. 60 (Pisa
Italy). '* K
1835. [Sorex] pachyurus Kiister, Isis, p. 77 (Cagliari, Sardinia).
1841. Pachyura etrusca Bonaparte, Ioonogr. Faun. Ital., i, Indice distrib.
1857. Grocidura suaveolens Blasius, Saugethiere Deutsohlands, p. 147.
1910. Pachyura etrusca Trouessart, Faune Mamm. d'Europe, p. 43.
Type locality. — Pisa, Italy.
Geographical distribution. — Italy and adjoining portions of the
Mediterranean region.* Limits of range not known.
Diagnosis. — Much smaller than Grocidura russula (head and
body, 35-40 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 condylobasal 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
Grocidura leucodon and C. mimula ; teeth not conspicuously
different 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.).
r External characters. — In general external form Pachyura
etrusca does not differ notably from Crocidura russula 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 G. 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.
Mammse, i 3 — 3 = 6.
Colour. — Upper parts a uniform slaty brown, perhaps best
described as drab-grey washed with light bister, the individual
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 essentially straight from nares to occipital,
usually a little concave in interorbital region, while in the
European forms of Crocidura it is usually a little convex
throughout, and never evidently concave in interorbital region.
* Spain. See footnote, p. 86.— O. T.
G 2
84 INSECTIVOKA
The brain-case is excessively flattened, relatively more so than in
Crocidura mimula, 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
a^Z^Psi which to condylobasal length is about 46,
Jil^j^ slightly less than in Crocidura mimula. Ante-
"* orbital foramen relatively large, its position
normal. Lachrymal foramen over posterior
half of m 1 . Palate relatively short and wide.
Mesopterygoid space relatively longer than in
the European forms of Crocidura, its anterior
border on line with posterior edge of third
jfio. le. molar instead of distinctly behind it, its lateral
FaCh xT£™ USCa ' borders nearly parallel, 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.
Teeth. — While in general, and aside from the presence of
the fourth unicuspid, the teeth do not differ materially from
those of the smaller European Crociduree except in size, they
show several peculiarities 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 unicuspid relatively
larger and higher than in Crocidura russula, its cingulum less
curved, and its width distinctly exceeding that of palate instead
of barely equalling it. Contrast in both height and crown area
of first and second unicuspids greater than in C. russula, 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. leucodon, 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. russula, 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
PACHYURA
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86
IXSECTIVORA
1 al.
2al.
lal.
Pisa, Italy.
Tuscany.
Italy.
Skeleton.
<5, 9 al.
Italy.
Sicily.
lal.
Sardinia.
lal.*
Lamia, Greece,
Genoa, Italy : head and body, 41 and 42 ; tail, 26 and 28 ; hind
foot, 7 ■ 6 and 8 ' 0. Adult female from Sassari, 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.M. and Mottaz) ; Florence, 6
(U.S.N.M. andilottaz) ; Tuscany, 2; Borne, 1 (Genoa) ; no exact locality, 2.
Sicily : No exact locality, 2.
Sakdiuta: Assuni, 1 (U.S.X.M.). No exact locality, 1.
Greece : Lamia, 1.
Marquis G. Doria (p). 69. 3. 4. 1.
(No history).
Zoological Society's 55. 12. 26. 295.
Collection.
Purchased. 58. 10. 21. 13.
Purchased (Parzu- 52. 2. 26. 22-23.
daki).
Hon. N. C. Boths- 11. 10. 1. 1.
child (p).
E. MacDoneU (p). 8. 7. 22. 1.
Genus CROCIDURA Wagler.
1832. Crocidura Wagler, Isis, p. 275 [Sorex leucodon Hermann).
1857. Crocidura Blasius, Saugethiere Deutschlands, p. 137.
1869. Leucodon Fatio, Faune Vert. Suisse, I, p. 132 (substitute for
Crocidura).
1897. Paurodus Schulze, Mamm. Europ., in Helios, Abhandl. n. Vortrage
Gesammtb. Naturwiss., xiv, p. 90 (Sorex leucodon Hermann and
S. araneus Schreber = S. russulus Hermann).
Type species. — Sorex leucodon Hermann.
Geographical distribution. — Africa and warmer portions of
Europe and Asia, including the Malay Archipelago ; in Europe,
north to northern Holland and central Germany, west to the
Atlantic coast and the Channel Islands. Absent from Great
Britain and Ireland.
Characters. — Upper unicuspid teeth 3-3 (dental formula :
i !t 3 , c J^J, pm !=1, m jj=| = 28 ; 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 rudimentary second
cusp and commissure ; teeth white throughout ; skull heavier and
more robust than in Sorex and Neomys, 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
* i, 2 9. llarismas, Lower Guadal- A. Chapman (c&p). 11.12.19.1-3.
quivir, Spain.
Beceived December, 1911.— 0. T.
CROGIDURA o I
above fur, the meatus closed by two valves as in Neomys ; tail
somewhat thickened, its surface covered with short hairs, among
which are sprinkled numerous longer ones ; habits terrestrial.
Remarks. — Though not highly modified in external pecu-
liarities the genus Grocidura 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 Soricidee the species of Grocidura 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 Sorex 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 OP GROCIDURA.
Tail decidedly more than half as long as head and
body, the ratio normally varying from 70 to 80.
Hind foot about 14 mm. (Sicily) C. caudata, p. 110.
Hind foot 11 to 12' 5 mm.
Upper tooth-row about 8 - 6 mm. (Corsica) C. cyrnensis, 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 uniouspid with crown area about
equal to that of third (Crete) C. canese, p. 109.
Second upper unicuspid with crown area distinctly
less than that of third C. mimuia, 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,
Prance) C. m. iculisma, p. 98.
Brain-case decidedly flattened (entire range
of species, western portion excepted)... C. m. mimiUa, p. 95.
Condylobasal length of skull 17 '6 to 20 • 4 mm.
(usually 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) G. leucodm, p. 88.
Animal obscurely bicolor ; condylobasal length
of skull usually 18 to 19 mm. (Sicily) C. sicula, p. 108.
Brain-oase more than half as high as wide C. russula, p. 99.
Condylobasal length of skull usually 19 to
20-4 mm. (Central Europe, south into
Italy) C. r. rusmla, p. 101.
Condylobasal length of skull usually 18 to 19 mm.
Colour a dark, coppery-brown (Portugal) C. r. cintne, p. 108.
Colour a pale drab-brown (Spain and south-
western Prance) C. r. pttlchra, p. 103,
88
INSECTIVORA
CROCIDURA LEUCODON Hermann.
1780. Sorex leucodon Hermann in Zimmermann, Geogr. Gesoh., n,
p. 382 (vicinity of Strassburg, Germany).
1781. Sorex leucodon Hermann in Schreber, Saugthiere, pi. clix.
1782. Sor[ex~] leucodon Hermann, Tabula Affinitatum Animalium, p. 79
(footnote).
1792. Sorex albipes Kerr, Anim. Kingd., p. 208 (based on Pennant's account
of S. leucodon Hermann).
1832. Croc[idura] leucodon Wagler, Isis, p. 275.
1857. Crocidura leucodon Blasius, Saugethiere Deutschlands, p. 140.
1869. Leucodon microurus Patio, Paune Vert. Suisse, i, p. 173 (Substitute
for leucodon).
1897. [Crocidura] leucodus Schulze, Mammalia Europaea, p. 18 (Substitute
for leucodon).
1910. Crocidura russula leucodon Trouessart, Paune Mamm. d'Europe, p. 44.
Type locality. — Vicinity of Strassburg, Germany.
Geographical 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 1 3 mm. ; condylobasal length of skull, 1 8 • 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
Sorex 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 Sorex 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 Sorex and
Neomys, 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 (6) loosely spreading hairs about 5 mm. long, rather
thickly sprinkled among the others ; annulation ill-defined,
about 35 to the centimeter at middle. Mammas, i 3 — 3 = 6.
CROCIDUEA
89
Colour. — Upper parts varying from a slaty drab to dull
russet, the hairs with a slight metallic gloss and with silvery
reflections, which in certain lights produce an evident effect of
fine speckling. Underparts and inner surface of limbs buffy
white, dulled to a varying degree by the slaty under colour, the
chin and throat often suffused with cream-buff. Line 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. Most
of the specimens examined are in some degree intermediate.
Feet dull whitish, often irregularly clouded with drab. Tail
sharply bicolor, whitish below, concolor with back above, the
longer hairs silvery grey.
Skull. — The skull is slightly larger than that of Sorex araneus
and noticeably more heavily built, particularly that portion
lying in front of brain-case, all of which is both broadened and
deepened, so that the general outline tapers less conspicuously
from behind forward, whether skull is viewed from above or
from the side. Brain-case less well marked off from interorbital
region than in the European species of Sorex and Neomys, its
main sutures closing early in life. Brain-case
slightly longer than broad, its posterior out- /7 ~~~~ — —
line rounded but broken by the slightly Lk^-i^S^p^
projecting points of the condyles, its antero-
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 fig. 20, page 100). Dorsal profile
with a slight concavity at front of brain-case
and slight convexity over middle of rostrum. v 17
Nares squarely truncate posteriorly, the Cm idu.m lencoHon.
lateral wall abruptly angled near middle. Nat. size.
Anteorbital foramen relatively smaller than in
Sorex. araneus, and region between it and edge of alveolus dis-
tinctly wider. Lachrymal foramen over metastyle of m 1 Plate
forming outer wall of anteorbital canal nearly three times as wide
as lachrymal foramen. Angular shelf -like region over posterior
molars broader and more prominent than in the European species
of Sorex, but rudimentary zygomatic process of maxillary reduced
to the merest trace. Mesopterygoid fossa as in Sorex araneus ;
floor of brain- case between tympanic bones narrower and with
distinct median ridge.
Teeth. — Dentition noticeably heavier than in the European
species of Sorex and Neomys, the difference in general aspect
heightened by the absence of brown colouring matter on points
of cusps. Anterior upper incisor with main cusp long and
slender, abruptly hooked downward; basal lobe low and
90
INSECTIVOBA
FIG. 18.
Croeidura 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 unicuspid tooth. General
form of first incisor not unlike that of
Neomys fodiens, but with even more
contrast between height of the two
cusps. Anterior lower incisor simple,
rather robust, the shaft slightly taper-
ing, the point slightly bent upward, 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 oblique 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 developed
and reaching posterior border of
tooth somewhat outside of middle.
Crushing surface well developed,
occupying 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 differen-
tiated than in first.
Second smaller than
third, its crown about
as broad as long. Third
separated from large pre-
molar by a slight space,
its crown longer than
broad, its cusp about
equal in height to para-
cone of large premolar.
Lower unicuspids essen-
tially alike in form, the
second differing from that
of Sorex and Neomys 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 Sorex and
Fig. 19.
Croeidura leucodon. Teeth X 10.
CROCIDURA 91
Neomya, the emargination of posterior border deeper, particularly
in pm*, 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 Sorex
araneus, but containing the same elements. Lower cheek-teeth
essentially as in Sorex araneus except for the peculiarities of m 3
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, Grisons, 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, 12-5(11- 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, Liege, 2; Waremme, Liege, 1 (U.S.N.M.).
Prance: Dinan, 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-
ringen, Bern, 4 (U.S.N.M.) ; Untervatz, Grisons, 15 (B.M. and U.S.N.M.) ;
Grisons, no exact locality, 1 ; Lugano, Ticino, 4 (U.S.N.M.) ; Zuber-
wangen, St. Gallon, 1.
Italy : Porlezza, Como, 1 (Mottaz) ; Boccadassa, Genoa, 1 (Genoa) ;
Pisa, 1 (Mottaz) ; Florence, 3 (U.S.N.M. and Mottaz) ; Rome, 1.
Remarks. — 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
russula, 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
animal.
92
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94 INSBCTIVORA
2 9, 1. Esneux, Liege, Belgium Lord Lilford (p). 95. 1. 1. 1-2.
(H. Gronvold.)
i. Dinan, Brittany, Prance. G. Barrett-Hamilton 11. 1. 2. 158.
( p )-
6. Manonville,Meurthe-et- Lord Lilford (p). 8.9.8.1.
Moselle.
5 6, 4 9. Etupes, Doubs, 350 m. 0. Thomas (p). 8. S. 10. 33-41.
(C. Mottaz.)
2 9. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.2-3.
Germany.
1. Frankfort, Hessen- Dr. Dieffenbach (p). 47.1.8.50.
Nassau.
i. Nussberg, Brunswick. G.Barrett-Hamilton 11. 1. 2. 90.
(*)■
1 al. Germany. Tomes Collection. 7. 1. 1. 45.
9. Hatszeg, Transylvania, C. G. Danford (o). 3. 2. 2. 16.
1500 ft. Hungary.
3<J,4 9. Untervatz, Grisons, O. Thomas (p). 4.4.5.21-22.
Switzerland. 10. 8. 16. 9-13.
(E. H. ZolUkofer.)
9 sk. Grisons, 550 m. O. Thomas (p). 4. 4. 5. 20.
(E. H. ZolUkofer.)
9. Lugano, Ticino. 0. Thomas (p). 10. 8. 16. 15.
(E. H. ZolUkofer.)
9. Zilberwangen.St.Gallen. 0. Thomas (p). 10. 8. 16. 14.
(E. H. ZolUkofer.)
i. Rome. (G. Goli.) G. Barrett-Hamilton 11. 1 2. 103.
CROCIDURA MIMULA Miller.
(Synonymy under subspecies.)
Geographical distribution. — Central Europe from north-
western Spain through Prance, Switzerland, central Germany
and northern Hungary to Roumania and Bulgaria, south into
Italy and Greece.
Diagnosis. — Size less than in Grocidura russula and C. leuaodon
(hind foot, 10 to 12 mm. ; condylobasal length of skull, 16 to
17 • 6 mm.) ; skull and teeth resembling those of G. 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. — Upper parts varying from a dull russet tinged with
sepia to a dark hair-brown with a tinge of drab, the hairs with
metallic gloss and silvery reflections ; underparts varying from
dull ochraceous-buff to greyish cream-buff, never sharply defined
from colour of back and sides. Feet buffy whitish, usually with
some dark clouding. Tail g-reyish or brownish, very obscurely
bicolor.
Skull and teeth. — Except for their noticeably smaller size the
skull and teeth closely resemble those of Grocidura leucodon,
though the brain-case is less constantly flattened, sometimes
assuming a form essentially like that in 0. russula. Plate forming
CB0CIDT7RA 95
outer wall of anteorbital canal narrower than in C. leucodon, its
width scarcely twice that of lachrymal 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. leucodon.
Measurements. — According to measurements made by various
collectors, the head and body varies from 55 to 72 rum., tail
from 28 to 40 mm., and hind foot from 10 to 12 mm. The
condylobasal length of skull ranges between 16 - and 17-6 mm.,
and length of upper tooth-row between 7 • 2 and 8 ■ mm.
Remarks. — Grocidura mimula 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. russula, 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 clearly understood.
Ceociduea mimula mimula Miller.
1839. ?? [Crocidura aranea] var. minor de Selys-Longchamps, Etudes de
Micromamm., p. 35 (Silesia).
1901. Crocidura mimula Miller, Proc. Biol. Soc, Washington, xiv, p. 95,
June 27, 1901 (Ziiberwangen, 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 G. antipai Trouessart, Paune Mamm. d'Europe,
pp. 46, 48.
Type locality. — 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, Orisons, 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(11-12). A male from Gageni,
Roumania, and female from Bustenari, Roumania : head and
body, 64 and 66 ; tail, 31 and 34; bind foot, 11 and 11. Two
adult males from Agay, Var, France : head and body, 64 and 66 ;
tail, 35 and 34 ; bind foot, 11 and 11. Adult 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, 1 1 ■ 8 and 11*4. For cranial
measurements see Table, p. 96.
Specimens examined. — Thirty-seven, from the following localities : —
France : Abbeville, Somme, 1 ; Agay, Var, 2.
Germany : Marxheim, Bavaria, 2.
Austria-Hungary: Haida, Arva, Bohemia, 4; Hatszeg, Hunyad,
Transylvania, 1; Tatra Mts., Hungary, 1.
Boumania : Gageni, Prahova, 1 ; Bustenari, Prahova, 1.
Bulgaria: Sofia, 1 (Andersen) ; Varna, 1 (Andersen).
Switzerland : Ziiberwangen, St. Gallen, 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 ; Borne, 1.
Greece : Corfu, 2.
1. Abbeville, Somme, Baillon Collection. 56. b.
France.
2 S. Agay, Var. G. S. Miller (c). 8. 8. 4. 162-163.
c5. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.4.
Germany. ( Wolter-
storff.)
<*, 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 Mountains. Dr. B. Collett (p). 91. 1. 21. 2.
9. Gageni, Prahova, Bou- 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.
<*. Locarno, Ticino. 0. Thomas (c & p). 5. 8. 2. 1.
<5, 9. Untervatz, Grisons. 0. Thomas (p). 4.4.5.21-22.
(E. H. Zollihofer.)
9. Lugano, Ticino. 0. Thomas (p). 4.4.5.57.
(E. H. Zollikofer.)
9. Viareggio, Lucca, 5 m. 0. Thomas (c & p). 5. 8. 2. 21.
Italy.
6. Borne. (Goli.) G. Barrett-Hamilton 11. 1. 2. 97.
(*)■
6. Corfu, 50 m. Greece. J.I. S.Whitaker (p). 8.10.1.8.
(0. Mottaz.)
Crocidura mimula iculisma Mottaz.
1908. Crocidura mimula iculisma Mottaz, Bull. Soc. Zool. de Geneve, i,
p. 119, April 30, 1908. Type in Mottaz Collection.
1910. Crocidura mimula iculisma Trouessart, Faune Mamm. d'Europe, p. 47.
Type locality. — Lignieres-Sonneville, Charente, France.
Geographical distribution. — Known from the type locality
only.
Diagnosis. — Size as in C. mimula mimula or slightly smaller
(hind foot, 10; condylobasal length of skull, 16); brain-case
deep, nearly as in C. russula.
CROCIDURA 99
Measurements. — 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).
Remarks. — While this race appears to be distinct from true
mimula, the material seen is insufficient to form the basis of any
final opinion as to its status.
Ceociduka mimula cantabra Cabrera.
1908. Crocidura cantabra Cabrera, Bol. Eeal Soo. Espafi. Hist. Nat., vm,
p. 239, May, 1908. Type in Madrid Museum.
1910. Crocidura cantabra Trouessart, Eaune Mamm. d'Europe, p. 46.
Type locality. — Basque 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.)
Remarks. — 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 G. russula pulchra to true russula.
CROCIDURA RUSSULA Hermann.
(Synonymy under subspecies.)
Geographical distribution. — 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
depressed, its height always at least half greatest width and
usually more ; large upper premolar with antero-external cusp
small, its height usually less than that of third unicuspid ;
colour of underparts not strongly contrasted with that of back,
the line of demarcation along side vaguely defined.
External characters. — In external characters, aside from the
relatively longer tail, Crocidura russula agrees with C. leucodon.
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 peculiar 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.
Skull 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 behind. Teeth essentially as in the
related species, but large upper premolar with antero-external
Fis. 20. Fig. 21.
Posterior view of skull of Crocidura russula. Anterior
Croeidura leucodon (upper teeth in profile. X 5.
figure), and C. russula (lower
figure), x It.
cusp (paracone) low, its height often much less than that of
third unicuspid, the distance from its point to ill-defined 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 different races the head and body
ranges from 64 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 differences 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 thoracicus
and Dehne's S. chrysothorax.
CROCIDUEA 101
Orociduea russula eussula Hermann.
1777. Sorex araneus Sohreber, Saugthiere, in, p. 573 (not of Linnaeus, 1758).
1780. Sorex russulus Hermann in Zimmermann, Geogr. Gesch., n, p. 382
(vicinity of Strassburg, Germany).
1780. 11 Sorex constrictus Hermann in Zimmermann, Geogr. Gesch., n,
p. 383 (vicinity of Strassburg, Germany). Based on young in nest.
1792. 11 Sorex unicolor Kerr, Anim. Kingd., p. 208 (Strassburg, Germany.
Based on Pennant's account of Sorex constrictus).
179S. Sorex musaraneus Ouvier, Tabl. Elem. de l'Hist. Nat. des Anim.,
p. 109 (Prance).
1800. 11 Sorex leucurus Shaw, Gen. Zool., I, pt. 2, p. 538 (Strassburg,
Germany. Based on Schreber, pi. clixc, S. constrictus Hermann).
1801. S\prex'] a[raneus] cinereus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
1801. S[orex'\ a[raneus] candidus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
1832. Sorex fimbriatus Wagler, Isis, p. 54 (Bavaria, Germany).
1832. Croc[idura] moschata Wagler, Isis, p. 275 (Substitute for Sorex
fimbriatus).
1832. C[rocidura] major Wagler, Isis, p. 1218 (Bavaria, Germany).
1832. Orocidura 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. 1 Sorex inodorus de Selys-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 aranea] var. minor de Selys-Longchamps, Etudes de
Micromamm., p. 35 (Silesia).
1839. [Crocidura aranea"] var. albiventris de Selys-Longchamps, Etudes de
Micromamm., p. 36. (No locality given.)
1839. ? Crocidura hydruntina Costa, Fauna del Kagno di Napoli, Mamm.,
p. 6 (Otranto, Calabria, Italy).
1855. Sorex chrysothorax Dehne, Allg. deutsche Naturhist. Zeitung, Neue
Polge, 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, Faune Mamm. d'Europe, p. 43.
Type 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 ? * ; Guernsey and
Alderney, Channel Islands.
Diagnosis. — Size rather large (hind foot, ll - 7 to 14, condy-
lobasal length 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, Guelderland, 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 Corsican C. cymensis (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" 7-1 3). Average of ten specimens from 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.
Specimens examined. — One hundred and two, from the following
localities : —
Holland : Oosterbeek, Guelderland, 4.
Belgium : Esneux, Liege, 9 ; Waremme, Liege, 8 (U.S.N.M.) ; no exact
locality, 1.
France : Boulogne-sur-Mer, Pas-de-Calais, 8 ; Guines, Pas-de-Calais, 2 ;
Abbeville, Somme, 2 (B.M. and Mottaz) ; Guernsey, Channel Islands, 4 ;
Alderney, Channel Islands, 1 ; St. Briac, Brittany, 1 ; Lignieres,
Charente, 1 (Mottaz) ; Nancy, Meurthe-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, Thilringen, 2 ; Ingelheim, Rheinhessen, 2 ;
Strassburg, 1.
Switzerland : Geneva, 10 (U.S.N.M . and Mottaz) ; St. Cergues, Vaud, 9
(U.S.N.M. and Mottaz) ; Grosjoan, Vaud, 1 (Mottaz) ; Chesieres, Vaud, 1
(Mottaz) ; Lucerne, 1 ; Vitznau, Lake of Lucerne, 3 ; Thurgau, Roggwil, 1 ;
St. GaUen, 3 (U.S.N.M.) ; Ziiberwangen, St. Gallon, 5 (B.M. and U.S.N.M.) ;
Degersheim, St. GaUen, 5 (B.M. and U.S.N.M.) ; Engelberg, Unterwalden, 1.
Italy : Ceresole d'Alba, Turin, 2 (Turin).
3 <5, 1 ?. Oosterbeek, Guelderland, 0. Thomas (c & p). 98. 2. 1. 9-12.
50 m. Holland.
Oosterbeek, Guelderland.
(0. Thomas.)
AS, 49,1. Esneux, Liege, Belgium.
1. Belgium. (H. Gronvold.)
2<5,9,1. Guernsey, Channel Is-
lands. (B.H.Bunting.)
Alderney.
Boulogne, Pas-de-Calais,
10 m. Prance.
Guines, Pas-de-Calais.
Abbeville, Somme.
St. Briac, Brittany.
Montauban, Haute-
Savoie.
Montauban, Haute-
Savoie, 900 m.
(A. 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.
<5,9.
9.
5c5,3 9.
2(5.
1.
1.
2.
2(5.
(5,9.
9.
2 9.
9.
3 9.
lal.
Miller Collection.
Lord Lilford (p).
Tomes Collection.
0. Thomas (p).
W.Eagle Clarke (p).
O. Thomas (c & p).
O. Thomas (c & p).
Baillon Collection.
W. M.Daly (c&p).
A.Robert (c&p).
0. Thomas (p).
G. S. Miller (c).
V. Builles (c&p).
0. Thomas (p).
C. Hilgert (c).
0. Thomas (p).
0. Thomas (c&p).
7. 7. 7. 3851-
3852.
95. 1. 1. 3-11.
7. 1. 1. 30.
8. 9. 2. 18-21.
9. 3. 28. 1.
98. 1. 9. 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. 3 .
CROCIDURA 103
Ceocidura eussula pulchea Cabrera.
1907. Crocidura russula pulchra 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. EspaS. Hist.
Nat., Madrid, vu, 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 paler
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, ll - 9 (10 • 8-1 2 - 4). Average and extremes of seven
specimens from Granada, 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 : —
Fbance : Cadillac-sur-Garonne, Gironde, 19 (U.S.N.M.) ; Montrejeau,
Haute-Garonne, 3 (U.S.N.M.) ; St. Genies, near Nimes, 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 cintree).
9. St. Genies, Gard, 102 m. 0. Thomas (p). 8. 8. 10. 42.
France. (C. Mottaz.)
4 S. Silos, Burgos, 980 m. G. S. Miller (c). 8. 8. 4. 38-41.
Spain.
?. Dehesa de Valencia. O. Thomas (p). 8. 2. 9. 46.
(N. Gonzalez.)
2 <5, 1 9. Alcoy, Alicante. 0. Thomas (p). 8. 2. 9. 43-45.
(N. Gonzalez.)
6, 9. Venta del Baul, Granada. G. S. Miller (c). 8.8.4.31-32.
4 6, 1 9. Granada. G. S. Miller (c). 8. 8. 4. 33-37.
dal. Sierra de Gerez.Portugal. Dr. H. Gadow 87.3.28.1.
(c & p).
* Intermediate between pulchra and true russula.
104
INSECTIVOEA
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Locality.
C. russula russula (continued).
Switzerland : Ziiberwangen, St. \
Gallen . . ./
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>>
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C. russula pulchra.
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11 )!
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1 08 INSECTIVORA
Ceociduea eussula cinte^; Miller.
1907. Crocidura russula cintrse Miller, Ann. and Mag. Nat. Hist., 7th ser.,
xx, p. 390, November, 1907. Type in British Museum.
1910. Crocidura russula cintrse Trouessart, Faune Mamm. d'Europe, p. 45.
Type locality. — Cintra, near Lisbon, Portugal.
Geographical distribution. — At present known only from the
type locality.
Diagnosis. — Size as in C. russula pulchra (hind foot, 1 1 ■ 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 Ridgway, the hairs with a peculiar, strong, coppery
lustre rarely indicated in the typical race.
Measurements. — External measurements of type : head and
body, 64 ; tail, 33 ; hind foot, 1 1 • 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 <$, 5 ?. Cintra, Estremadura, O. Thomas (c & p). 98. 2. 2. 10-20.
300-350 m. Portugal. (Type of subspecies 98. 2. 2. 11.)
CROCIDURA SICULA Miller.
1879. ? Crocidura sicula Giglioli, Wiegmann's Archiv fur Naturgesch.,
1879, I, p. 96. Nomen nudum : " Crocidura sicula (Giglioli MSS.
sp. nov. ?) Castelbuono, 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, Faune 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 "6 to 19 mm.); brain-case nearly as much flattened as in C.
leucodon, its depth usually a little less than half greatest width ;
crown area of molars slightly reduced.
Colour. — Upper parts a light bluish drab tinged with sepia ;
underparts faintly constrasted pale smoke-grey, with or without
a buffy cast. Feet dull whitish grey. Tail obscurely bicolor,
brownish above, whitish grey below.
Skull and teeth. — Except for its distinctly flattened brain-case
CROCIDURA 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 antero-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 Sioily. Exact localities :
Palermo, 5 (B.M. and . TJ.S.N.M.) ; Marsala, 2 ; San Giuglielmo, Castel-
buono, 6 ; Picuzza, 1.
2 9. Palermo, Sicily. J.I. S. Whitaker (p). 98.10.6.2-2*.
1. Marsala. (A. Robert.) 0. Thomas (p). 6. 8. 4. 26.
3 <5, 1 9. San Giuglielmo, Castei- O. Thomas (p). 8. 9. 1. 6-9.
buono. (A. Robert.)
CROCIDURA CANEffi Miller.
1909. Crocidura caneee Miller, Ann. and Mag. Nat. Hist., 8th ser., in,
p. 418, May, 1909. Type in British Museum.
1910. Crocidura canese Trouessart, Faune Mamin. d'Europe, p. 48.
Type locality. — Crete.
Geographical distribution. — Island of Crete.
Diagnosis. — Size and general appearance as in the smaller
forms of CrocidMra russula, and skull with similarly deep cranium ;
but second upper premolar as large as third, and entire anterior
portion of upper tooth-row unusually long relatively to cheek-
teeth.
Colour. — The colour does not differ appreciably from that of
dark individuals of C. russula.
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.
Teeth. — 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 unicuspid. 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-row. In
C. canese 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 INSECTIVORA
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 (Linnsea, 84. 3. 14. 2.
Frankfort).
(Type of species.)
i. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 1.
(C. H. B. Grant.)
CROCIDURA CAUDATA Miller.
1901. Crocidura caudata Miller, Proc. Biol. Soc. Washington, xiv, p. 42,
April 25, 1901. Type in U.S. National Museum.
1910. Crocidura caudata Trouessart, Faune Mamm. d'Europe, p. 49.
Type locality. — Palesmo, Sicily.
Geographical distribution. — Sicily.
Diagnosis. — Size about as in large specimens of G. russula
(hind foot, 14 mm.). Tail very long, its ratio toihead and body
about 80, and so thickened that its diameter at middle is 3 mm.
(in other European species 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 differ 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 sicula. After
eight years more in the same fluid the back appears to have
assumed a somewhat more brownish cast.
Skull and teeth. — The only known skull is so injured that the
details of its form cannot be seen. The rostral portion does not
differ appreciably from that of C. russula. Teeth essentially as
in C. russula, but first upper unicuspid larger, third unicuspid
more crowded against large premolar, and cutting edge of large
premolar 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.
CROCIDTJRA HI
Measurements. — External measurements of type : head and
body, 63 ; tail, 52 ; hind foot, 14. For cranial measurements
see Table, p. 113.
Specimen examined. — The type.
Remarks. — 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., xx,
p. 390, November, 1907. Type in British Museum.
1910. Crocidura cyrnensis Trouessart, ITaune 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. — While 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 j hind foot,
12. For cranial measurements see Table, p. 113.
Specimens examined. — Three, all from Corsica.
"Remarks. — 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 INSBCTIVORA
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.
S 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. Grocidura russula Thomas, Proc. Zool. Soc. London, p. 39.
1907. Grocidura balearica Miller, Ann. and Mag. Nat. Hist., 7th ser., xx,
p. 391, November, 1907. Type in British Museum.
1910. Crocidura balearica 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 cyrnensis at present known, though the
actual elements of the colour are the same. Feet with the same
colour pattern.
Skull 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.
Specimens examined. — Three, all from the type locality.
£, 2? San Cristobal, Minorca; O. Thomas and R. I. Pocock 0.7.1.40-42.
Balearic Islands. (c & p).
(0. 7. 1. 42. Type of species.)
CKOCIDUBA
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114 INSBCTIVORA
Family ERLTSTACEID^E.
1821. Erinaceidse Gray, London Med. Repos., xv, p. 300, April 1, 1821.
Geographical distribution. — 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 ; floor 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 pterygoid
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 cusps 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, stiff spines.
BemarJcs. — 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 Linnaeus.
1758. Erinaceus Linnaeus, Syst. Nat., i, 10th ed., p. 52 (E. europseus).
1857. Erinaceus Blasius, Saugethiere Deutsehlands, p. 152.
1868. Herinaceus Mina-Palumbo, Ann. Agrio. Sicil., 2nd ser., xii, p. 37.
Type species. — Erinaceus europseus Linnseus.
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 about equally of ectopterygoid and a broad horizontal
outgrowth from palatine; dental formula: i jjr*, c^l, pm 3z3
m 3 1 3 = 36 ; canines not differentiated by form from the contiguous
teeth ; third upper molar consisting of a large protocone and
minute paracone, all trace of crushing surface absent ; body
short and heavy, the back covered with stiff, sharply pointed
bristles of uniform length ; tail shorter than hind foot.
EEINACEUS 115
Bemarks. — About twenty -five species are currently referred
to this genus, four of them occurring in Europe.
KEY TO THE EUROPEAN FORMS OP EEINACEUS.
Third upper incisor 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 Prance)... E. algirus, p. 130.
Underpart8 clouded with brown, at least in inter-
ramial and intercrural regions (Spain and
southern Prance) E. a. algirus, p. 131.
Underparts entirely whitish (Balearic Islands).... E. a. vagans, p. 133.
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. ; condylobasal 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 skull 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, buffy grey, but usually with notice-
able blackish markings (Central-western
Europe) E. e. europseus, p. 120.
Average colour lighter, the face usually clear,
pale, buffy grey without noticeable black-
ish markings (Iberian Peninsula) E. e. hispanicus, 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. eonsolei, p. 126.
ERINACEUS EUROPSEUS Linmeus.
(Synonymy under subspecies.)
Geographical distribution. — Europe from the Mediterranean
coast to Scotland and southern Scandinavia ; west to Ireland •
eastern limits of range not known.
Diagnosis. — Size large (head and body about 225 to 275 ; 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 posterior 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 (fig. 25 a).
External characters.— General form short, thick and clumsy,
the legs short, the feet large, plantigrade. Legs, tail, underparts
and head, except crown, clothed with coarse fur, the finer more
woolly hairs of which are about 15 mm. long, the longer, straight
hairs about 40 mm. in length. The rest of the body is covered
by a densely-set mass of sharply pointed bristles about 25 mm. in
length and 1 mm. in diameter. * The skin on which these bristles
are set is loosely attached to body, and provided 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 complete 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-confluent 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 hind foot. Bar 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 edges finely scalloped, its lower border continued
as a pair of parallel ridges extending inward to palate. Eye
well developed but rather small. Mammae : p 1-1, a 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. Spines buffy at base, then with
* Barely the bristles are replaced by coarse hair like that on under-
parts. See Natural Science, xm, p. 156, pi. n, September, 1898.
BRIKACBUS 117
a slaty area of variable width, followed by a narrow but sharply
denned buffy annulation and an obscurely darker tip. The
general effect is a coarse grizzle, the exact tone of which varies
considerably in the different geographical races as well as in
individuals of the same race.
Shull. — General form of skull rather short, heavy and deep,
the zygomatic breadth about £ upper length, the brain-case not
rig. 22.
Erinaceus europseus. Nat. size.
conspicuously wider than interorbital region, the rostrum short
and deep (distance 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 INSEOTIVOEA
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 occipital 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 pointed 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 postero-externally, its
greatest breadth (antero-internal) about 3 mm. Inner and outer
pterygoid plates broadly triangular, approximately alike in size
and form, each containing more of the pterygoid than palatine
element. Hamular short, strongly curved. Mesopterygoid space
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 ridge 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 anterior border
over anterior root of large premolar. 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 : nearly square, broadly or narrowly
cuneate, rounded, or rounded with supplemental inner spicule.
Mandible short and heavy, the greatest depth of ramus 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, heavy 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 equal 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 third sub-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 very
rudimentary secondary cusplet. First and second incisors invari-
ably single-rooted. Third incisor single-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 often showing traces of longitudinal division, and perhaps
rarely double. Anterior lower incisor essen-
tially like the corresponding upper tooth
but not so high. Its shaft is directed
obliquely forward in line with symphysis,
the teeth of opposite sides parallel, separated
throughout by a narrow space.
Three succeeding teeth unicuspid,
the crowns similar in outline to
those of upper unicuspids, but an-
terior cusps obsolete and posterior
cusplets relatively better developed.
Second upper premolar scarcely
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 faintly indicated commissures, that
extending outward and backward from metacone to rudimen-
tary metastyle the most distinct. Parastyle and mesostyle
no. 23.
Erinaceus europseus. Teeth.
1 20 INSBCTIVOEA
absent.* A small but evident metaconule. Second molar like
first but smaller, its crown area about equal to tbat 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 bueopjeus europ^eus Linnseus.
1758. [Erinaceus] europseus Linnaeus, Systema Naturse, I, 10th ed., p. 52
(Sweden).
1779. [Hystrix] erinaceus Blumenbach, Handbuoh d. Naturgesch., p. 72
(Germany).
1803. Erinaceus suillus Geofiroy, Catal. Mammif. du Mus. Nat. d'Hist.
Nat., p. 67 (Prance).
1803. Erinaceus caninus Geofiroy, Catal. Mammif. du Mus. Nat. d'Hist.
Nat., p. 68 (France).
1857. Erinaceus europseus Blasius, Saugethiere Deutschlands, p. 153 (part).
1858. E[rinaceus] caniceps Hamilton Smith, Jard. Nat. Libr., 2nd ed., xv
(Mammalia I), p. 148 (near Brussels, Belgium).
1897. [Erinaceus] echinus Schulze, Abh. a. Vortr. Gesammtb. Naturw. iv,
No. 10, p. 19 (Substitute for europssus).
1900. Erinaceus europseus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., v, p. 362, April, 1900.
1900. Erinaceus europssus occidentalis Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., v, p. 362, April, 1900 (Haddingtonshire,
Scotland). Type in British Museum.
1910. Erinaceus europseus and E. europseus occidentalis Trouessart, Faune
Mamm. d'Europe, p. 38.
Type locality. — Upsala, Sweden.
Geographical 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 mm., 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.
ERINACBDS 121
female from Oundle, Northampton, England : head and body,
249 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 and 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 ; Innerwiok,
Haddingtonshire, 1 ; Glendoo, Inverness, 1 (Wilson) ; Lanarkshire, 1.
England: Kelnsea, Spurn, Yorkshire, 1; Leeds, Yorkshire, 1 (U.S.N.M.);
Somersetshire, 4; Shrewsbury, Shropshire, 3; Oundle, Northampton, 2;
Graftonbury, Herefordshire, 1 ; Saffron Walden, Essex, 2 ; Banstead,
Surrey, 1 ; Elstead, near Godalming, Surrey, 1 ; Oekley, Surrey, 1 ;
Wandsworth Common, Surrey, 1 ; Hampshire, 1 (U.S.N.M.).
Wales : Cardiff, 2.
Ireland : Castle Hamilton, 1 ; Nenagh, Tipperary, 2 ; Ennis, Co.
Clare, 2 ; Glenmore, Co. Donegal, 1 (U.S.N.M.) ; Kilmanook, Wexford, 2.
Norway : Asker, near Christiania, 1.
Sweden : Upsala, 5 (U.S.N.M.) ; Upland, 1.
Denmark: Copenhagen, 5 (Andersen).
Holland : No exact locality, 1.
Belgium : Maredsous, Namur, 2.
Prance : Eoret de Bouconne, Gers, 1 ; Cranves-Sales, Haute-Savoie, 1.
Germany: Brunswick, 1 (U.S.N.M.) ; Heidelberg, 1 (U.S.N.M.) ; Strass,
near Burgheim, Bavaria, 3 ; Ingelheim, Bheinhessen, 1.
Switzerland : Geneva, 2 (Mottaz) ; St. Gallen, 7 (B.M. and U.S.N.M.) ;
Uzwil, St. Gallen, 1 (U.S.N.M.) ; Heresau, St. Gallen, 1 (U.S.N.M.) ; Wolf-
halden, Appenzell, 1 (U.S.N.M.) ; Thurgau, 2.
Remarks. — 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.*
<5, 9. Gordonstown, Elginshire, W. R. Ogilvie-Grant 11. 1. 3. 379-
Scotland. (c & p). 380.
<5. Innerwick, Haddington- W. Eagle Clarke (c & 0. 3. 13. 1.
shire. p).
(Type of E. e. occidentalis B.-Ham.)
9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 76.
i, 9. Oundle, Northampton, Hon. N. C. Roths- 11. 1. 3. 381-
England. child (c & p). 382.
S. Graftonbury, Hereford- W. E. de Winton (c 11. 1. 3. 383.
shire. & p).
* For discussion of this character see Lonnberg, Ann. and Mag. Nat.
Hist., 7th ser., 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
INSBCTIVORA
2 6.
6.
6.
6.
3 juv.al.
S, 9.
6, 9 al.
9.
6 al. <5.
2 6.
Saffron Walden, Essex. G. Barrett-Hamilton 11, 1. 2. 91-92.
(Wright.)
Banstead, Surrey.
Godalming, Surrey.
Ockley, Surrey.
Shrewsbury, Shropshire.
Somerset.
Somerset.
Cardiff, Glamorganshire,
Wales.
Nenagh, Tipperary, Ire-
land. (W. Smithwick.)
Ennis, Glare.
Kilmanock, Wexford.
C.H.B.Grant(c&p).
W.T.Blanford(o&p).
Hon. Ella Scarlett
(c & p).
H. E. Forrest (c & p).
Dr. J. Anderson (s).
Dr. J. Anderson (p).
E. Drane (c & p).
11. 1. 3. 386.
11. 1. 3. 384.
11. 1. 3. 385.
Asker, Christiania, Nor-
way.
Upland, Sweden. (G.
Eolthoff.)
Holland. {Seba Coll.)
Maredsous, Namur, Bel-
gium.
Foret de Bouoonne, Gers,
250 m. Prance. (A.
Robert.)
Cranves - Sales, Haute-
Savoie. (A. Robert.)
Burgheim, Bavaria, Ger-
many. (Wolterstorff.)
Ingelheim, Rheinhessen.
2 <5, 9 juv. St. Gallen, 500 m.
Switzerland. (E. H.
Zollikofer.)
6, 9. Thurgau, 400 m. (E. H.
Zollikofer.)
6 juv.
lal.
2 6.
6, 9.
6.
0. 9. 23. 1-3.
93. 7. 81. 1-2.
93. 7. 31. 3-4.
11. 1. 3. 387.
G. Barrett-Hamilton 11. 1. 2. 95.
(*)■
J. W. Scott (o & p). 93. 10. 30. 1-2.
G. Barrett-Hamilton 11. 1. 2. 93-94.
(o & p).
Christiania Museum 93. 3. 1. 7.
(e).
LordLilford (p). 11. 1. 1. 151.
Lidth de Jeude Coll. 67. 4. 12. 555.
Bev. G. Pournier (c 1. 6. 2. 1-2.
&p).
O. Thomas (p). 6. 4. 1. 10.
O. Thomas (p).
Lord Lilford (p).
O. Hilgert (c).
0. Thomas (p).
O. 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.
Ekinaceus euroPjEUS hispanicus Barrett-Hamilton.
1900. Erinaceus europseus hispanicus Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., v, p. 363, April, 1900. Type in British Museum.
1910. Erinaceus europseus hispanicus Trouessart, Faune Mamm. d'Europe,
p. 39.
Type locality. — Seville, Spain.
Geographical distribution. — Iberian Peninsula.
Diagnosis. — Size large, essentially as in E. europseus europseus
(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 ;
spines 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 scarcely speckled, 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 following localities in Spain :
Arreohavaleta, Vitoria, 1 ; Pajares, Leon, 7 ; Burgos, 5 ; Palacios de la
Sierra, Burgos, 1; Bejar, Salamanca, 2; Seville, 4.
Remarks. — The Spanish hedgehog is a moderately well
differentiated form. Extreme specimens are easily distinguish-
able from typical europaeus ; but in general the difference between
the two races must be regarded as an average one.
6. Arreohavaleta, Vitoria, 0. Thomas (p). 8. 2. 9. 9.
Spain. (N. Gonzalez.)
<5, 2 9, ¥ juv. Pajares, Leon. (N. Gon- O. Thomas (p). 8. 2. 9. 2-6.
S, 9, ? juv. Burgos. G. S. Miller (c). 8. 8. 4. 17-19.
2 <5. Burgos. (N. Gonzalez.) O. Thomas (p). 8. 2. 9. 10-11.
2 c5 juv. Bejar, Salamanca. (N. O. Thomas (p). 8. 2. 9. 7-8.
Gonzalez.)
6, 2 ?, 1. SeviUe. (Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 1-4.
(Type of suhspecies 95. 3. 3. 2.)
Erinaceus eueop^eus italicus Barrett-Hamilton.
1857. Erinaceus europseus Blasius, Saugethiere Deutschlands, p. 153 (part).
1900. Erinaceus europseus italicus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., v, p. 364, April, 1900. Type in British Museum.
1910. Erinaceus europseus italicus Trouessart, Faune Mamm. d'Europe,
p. 39.
Type locality. — Siena, Italy.
Geographical distribution. — Italian Switzerland, Italy and
Sardinia ; Corsica 1
Diagnosis. — 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 individuals of
typical europseus, though the speckling of the spiny area seems in
general to be finer, and the underparts usually lack all heavy
dark clouding.
Measurements. — Adult female from Curoggio, Ticino, Switzer-
land : head and body, 250; tail, 29 ; hind foot, 41. Two adult
124
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males from Siena, Italy : head 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 : —
Switzebland: Bigorio, Ticino, 1 (U.S.N.M.); Curoggio, Tieino, 1
(U.S.N.M.) ; Gentilino, Ticino, 1 (U.S.N.M.).
Italy : Empoli, Florence, 1 ; Siena, 4 ; Ostia, Borne, 1 ; Borne, 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 europseus. 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 specimens 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 Corsica, but
no specimens have yet been compared with the Italian race.
¥. Empoli, Florence, Italy. A. H. Savage Landor 97.3.7.1.
(c & p).
2 6, 2 ?. Siena. (S. Brogi.) Dr. E. Hamilton (p). 98. 10. 2. 5-8.
(Type of subspecies 98. 10. 2. 5.)
1. Ostia, Borne. Dr.L. Sambon(c&p). 1. 1. 2. 7.
3 i. Borne. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 3-4. 96.
(*)■
£. Su Gramu, Sardinia. (W. O. Thomas (p). 0. 12. 3. 5.
Wolterstorff.)
t. Bare. (W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 3.
9. Marusei. (W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 6.
? juv. Trecorgia. (W. Wolterstorff.) O. Thomas (p). 0.12.3.4.
Erinaceus europseus consolei Barrett-Hamilton.
1900. Erinaceus europseus consolei Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., v, p. 366, April, 1900. Type in British Museum.
1910. Erinaceus europseus consolei Trouessart, Faune Mamm. d'Europe,
p. 40.
Type locality. — Palermo, Sicily.
Geographical distribution. — Sicily.
Diagnosis. — Size apparently as in JS. europseus italicus (only
known specimen imperfect) ; colour differing from that of
italicus in the uniform dusky brown head and shoulders ; quills
unusually robust, their dark and light markings strikingly con-
trasted.
Colour. — Whole head and sides of neck and shoulders a
uniform dark brown between hair-brown and sepia, sprinkled
with buffy grey hairs. The dark brown continues back along
* Type.
ERINACEUS 127
edge of spiny 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 middle of back than on those of sides, thus producing a slight
though evident darker median dorsal area. Feet so injured that
colour cannot be determined.
Skull and teeth. — The imperfect skull shows no peculiarities.
Teeth as in specimens from the mainland.
Measurements. — Type (sex not known) : head and body, 252 ;
tail, 50 ; hind foot, 40. For cranial measurements see Table,
p. 125.
Specimen examined. — The type.
Remarks. — If not an abnormal specimen of E. europseus italicus
the type of consolei represents a very distinct local race.
1. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 1.
(Type of subspecies.)
ERINACEUS ROUMANICUS Barrett-Hamilton.
1900. Erinaceus europseus roumanicus Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., v, p. 365, April, 1900. (Gageni, Boumania.)
Type in British Museum.
1901. Erinaceus danubicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.
Freunde, Berlin, p. 229, December, 1901. (Prundu, Boumania.)
1910. Erinaceus europssus roumanicus and E. europseus danubicus Troues-
sart, Faune Mamm. d'Europe, pp. 40-41.
Type locality. — Gageni, Prahova, Roumania.
Geographical distribution. — -From eastern Germany (Konigs-
berg) and northern Bohemia south through Hungary, into
Greece.
Diagnosis. — In general like Erinaceus europseus, but colour of
furred parts usually much darker, and chest often with a large,
conspicuously contrasted whitish area ; skull with upper length of
maxillary greater than depth of rostrum at middle ; teeth
essentially as in E. europseus, but first upper premolar with
postero-basal cusp usually more distinct.
Colour. — Furred area dark hair-brown interspersed with buffy
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. europseus, the
general effect of the spiny area darker and less speckled.
Skull and teeth. — The skull differs from that of E. europseus
in the relatively 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
128 INSBCTIVOEA
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
157. europseus, and less distinctly marked off from general contour
Fig. 24.
Erinaceus roumanicus. Nat. size.
of skull, its degree of development somewhat as in E. algirus.
Teeth not certainly distinguishable from those of Erinaeeus
europseus, 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.M.).
ERINACEUS 129
Austria-Hungary: Haida, Arva, Bohemia, 1; Vasoar, Eisenburg,
Hungary, 1 (U.S.N.M.).
Roumania : Gageni, Prahova, 2.
Greece: Corfu, 2; Cephalonia, 1; Tatoi, near Athens, 1.
Bemarles. — Though at first sight very similar to Erinaceus
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 pattern which appears to be rarely if ever well
developed in the related species.
9. Haida, Bohemia. Lord Lilford (p). 97. 8. 14. 1.
<5, 9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 15-16.
(TV. Dodson.) (Type of species 4. 4. 6. 16.)
6. Potamos, Corfu, Greece. J. I. S. Whitaker (p). 8. 10. 1. 6.
(C. Mottaz.)
9. Argostoli, Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 7.
(C. Mottaz.)
A. Tatoi, Athens. C. Mottaz (c). 8. 11. 3. 8.
ERINACEUS NESIOTES Bate.
1906. Erinaceus europseus nesiotes Bate, Proc. Zool. Soc. London, 1905, n,
p. 316, April 5, 1906. Type in British Museum.
1910. Erinaceus europmus nesiotes Trouessart, Faune Mamm. d'Europe,
p. 40.
Type locality. — Near Gonia, western Crete.
Geographical distribution. — Island of Crete.
Diagnosis. — Similar to Erinaceus roumanicus but smaller (hind
foot, 35 to 38 ; 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.
Colour. — The colour is like that of E. roumanicus. 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
dark wash on posterior half of underparts.
Skull. — As in E. roumanicus, but not attaining as large size.
Measurements. — Adult 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
Table, p. 132.
Specimens examined. — Three, all from Crete.
Bemarles. — The Cretan hedgehog is nearly related to Erinaceus
nnimanicus, though well differentiated by its smaller size and by
the relatively large fourth unicuspid tooth.
■i, ?. Gonia, Crete. Miss D. Bate (c). 5. 12. 2. 11-12.
(5. 12. 2. 11 Type of species.)
9. Mesoghia. Miss D. Bate (c). 5. 12. 2. 13.
* Type.
130
INSECTIVORA
ERINACEUS alcirus Duvernoy and Lereboullet.
(Synonymy under subspecies.)
Geographical distribution. — Northern Africa ; also in southern
Spain, southern France, and the Balearic Islands.
Diagnosis. — Smaller than Erinaceus europseus (hind foot less
than 40 mm.) ; spines 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 portion of posterior lower
premolar with only two cusps (fig. 25 b).
External characters. — Externally Erinaceus alyirus is dis-
tinguishable from E. europseus 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. europseus.
Skull. — The form of the skull is essentially as in E. europseus
except that rostrum is less elevated posteriorly, so that the dorsal
profile tends to become slightly concave. Sagittal crest when"
fully developed extending forward to middle of frontal, while in
E. europseus it is usually confined to parietals, rarely encroaching
on posterior edge of frontal. Ridge at margin of orbit very short,
scarcely more than a process above lachrymal foramen. Bony
palate extending behind transverse ridge as a well-defined flat area
divided along median suture by a longitudinal ridge representing
the median spine of E. europseus. Basisphenoid pit narrower,
relatively deeper, and with more overhanging edges than in
E. europseus.
Teeth. — In general the teeth show no departure from those
of E. europseus. The posterior lower pre-
molar, however, lacks all trace of the meta-
conid, so that the resemblance of the elevated
portion of the tooth to the first triangle lof
)»! and m 2 is completely destroyed. In the
upper jaw the third incisor, canine, and
a b first premolar are two-rooted, apparently
I'ig. 25. without exception.
Large lower premolar of Measurements. — Head and body about
fndT^i>«Tb) M xl 200 to 250 ; tail, 25 to 40 ; hind foot, 32 to
37 ; condylobasal length of skull, 54 to 59 mm.
Remarks. — This species is readily distinguishable from Erinaceus
europseus by the bare spot among spines of forehead, the perfectly
two-cusped large lower premolar, and the two-rooted third upper
ERINACEUS
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.
Erinaceus algirus algirus Duvernoy and Lereboullet.
1840. Erinaceus algirus Duvernoy and Lereboullet, Mem. Soc. Mus.
d'Hist. Nat. Strasbourg, in, fasc. 2, p. 4.
1898. Erinaceus algirus de Winton, Proo. Zool. Soc. London, 1897, p. 955.
Type locality. — Oran, Algeria.
Geographical distribution. — Northern Africa ; also in southern
Spain and south-eastern Prance.
Diagnosis. — 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.
Colour. — Furred area bufly white to base of hairs, except on
muzzle, cheeks, interramial region, a narrow band along 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
dark. Throughout the spiny area the whitish strongly pre-
dominates, especially when animal is viewed from in front.
Claws light yellowish horn-colour.
Measurements. — Adult male from Schaf-el-Kab, Morocco
(teeth much worn) : head and body, 206 ; tail, 26 ; hind foot, 32.
For cranial measurements see Table, p. 132.
Specimens examined. — Numerous specimens from Northern Africa ; also-
an adult from "Andalucia," Spain; a young, less than half grown, from
Elche, Alicante, Spain ; and a still younger specimen from Lecques, Var,
France.
Remarks. — Owing to the unsatisfactory nature of the Spanish
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 the North African form
it would seem probable that they have been introduced within
historic times. The specimen from Elche, though undoubtedly a.
wild-bred animal, is too young to be positively determined as to
geographical race. The same is even more true of that from Var.
The adult from " Andalucia " lacks detailed history. Externally
it resembles the African form as compared with E. algirus vagans,
but the skull is small,* essentially as in the Balearic race.
* Condylobasal length, 53 - 8; zygomatic breadth, 32 - 0; least inter-
orbital breadth, 14 ■ 8 ; mandible, 41 • 2 ; upper tooth-row, 27 • ; lower tooth-
row, 22-0.
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ERINACEUS
133
1. Andaluoia, Spain. Lord Lilford (p). 94. 6. 11. 5.
S juv. Blohe, Alicante. G. S. Miller (c). 8. 8. 4. 16.
juv. al. Lecques, Var, France. Dr. P. Siepi (p). 98. 8. 25. 1.
Eeinacbus algirus vagans Thomas.
1901. Erinuceus algirus vagans Thomas, Proe. Zool. Soc. London, p. 38.
Type in British Museum.
1910. Erinaceus algirus vagans Trouessart, Paune Mamm. d'Europe, p. 39.
Type locality. — San Cristobal, Minorca, Balearic Islands.
Geographical distribution. — Balearic Islands.
Diagnosis. — 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 examined. — One from Majorca and five from Minorca,
Balearic Islands.
Remarks. — 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.
6. Inca, Majorca ; 300 m. O. Thomas & R. I. Pocock 0. 7. 1. 6.
Balearic Islands. (c & p).
4 <5, 9. San Cristobal, Minorca. O. Thomas & B. I. Pocock 0. 7. 1. 35-39.
(c & p).
(0. 7. 1. 36. Type of subspecies.)
134
CHIROPTERA
Order CHIROPTERA.
1779. Ghiroptera Blumenbach, Handbuch der Naturgeschichte, p. 74.
Geographical distribution. — Practically cosmopolitan ; only
absent from the treeless arctic and antarctic regions, 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 backward.
Remarks. — The order Ghiroptera, 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, equally
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 Insectivora, is
shown by numerous peculiarities of structure, among others the
relatively simple character of the brain. Two sub-orders are
recognized among the recent members of the order, the
Megachiroptera, not represented in Europe, with less highly
modified skeleton of fore limb, and more modified teeth, and the
Microchiroptera with more highly developed wing and in most
instances more primitive teeth.
Sub-Order MICROCHIROPTERA.
1821. Insectivora Gray, London Medical Repository, xv, p. 299, April 1,
1821.
1872. Animalivora Gill, Arrangement of the Families of Mammals, p. 16,
November, 1872.
1875. Microchiroptera Dobson, Ann. and Mag. Nat. Hist., 4th ser., xvi,
p. 346, November, 1872.
1878. Microchiroptera Dobson, Catal. Chiropt. Brit. Mus., p. 2.
1907. Microchiroptera Miller, Families and Genera of Bats, p. 78, June 29,
1907.
Geographical distribution. — The same as that of the order.
In Europe north to the limits of tree growth, west to Ireland
and the Azores.
MICROCHIEOPTEEA
135
Characters. — Anterior limb very highly modified, the second
finger 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 different from each other
(except when excessively reduced, as in the South American
Desmodontidse) ; margin of ear not forming a ring ; tragus usually
present. Mammse in all European genera, p 1-1 = 2.
Bemarlcs. — The sub-order Microchiroptera is essentially cos-
mopolitan in distribution. Though more highly modified than
the Megachiroptera in wing structure, the members of this group
for the most part retain 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 present 17 families and nearly 150
genera are recognized ; the species are 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 MICBOCHIBOPTEBA.
Tragus absent ; muzzle with leaf -like outgrowths ;
premaxillaries represented by palatal branch
only, not fused with surrounding parts (often
lost in prepared specimens) Bhinolophidai, 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 Molossidas, 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 Vespertilionidss, p. 165.
Presternum with median lobe much smaller
than body of bone ; coracoid curved out-
ward ; second phalanx of third ringer not
specially elongated Vespertilioninse, 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 Miniopterinx, p. 26S.
136 CHIEOPTERA
KEY TO THE GENERA OP EUROPEAN BATS.
(A wholly artificial key based primarily on external characters.)
Muzzle with leaf-like outgrowths Rhinolophus, p. 137.
Muzzle without leaf-like outgrowths.
Tail projecting conspicuously beyond membrane ... Nyctinomus, p. 276.
Tail not projecting conspicuously beyond membrane.
Ears joined.
Ear longer than head Plecotus, p. 256.
Ear shorter than head Barbastella, p. 263.
Ears separate.
Second phalanx of third finger nearly three
times as long as first Miniopicrus, p. 268.
Second phalanx of third finger less than twice
as long as first.
Fifth finger about as long as metacarpal of
fourth or third Nyctalus, 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 Vespcrtilio, p. 238.
Ear higher than wide, its lower margin
not forming pocket near angle of
mouth.
Upper cheek-teeth 6-6 Myotis, p. 166.
Upper cheek-teeth less than 6-6.
Upper cheek-teeth 5-5 Pipistrellus, p. 202.
Upper cheek-teeth 4-4 Epteshus, p. 224.
Family RHINOLOPHIDiE.
1827. Rhinolophina Lesson, Man. de Mammalogie, p. 81 (part).
1857. Phyllostomata Blasius, Saugethiere Deutschlands, p. 26.
1866. RMnolophidai Gray, Proc. Zool. Soc. London, p. 81 (part).
1878. Rhinolophidai Dobson, Catal. Ghiropt. Brit. Mus., p. 100 (part).
1907. Rhinolophidse Miller, Families and Genera of Bats, p. 106, June 29,
1907.
Geographical distribution. — 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 (fig. 26) consisting of a horizontal
anterior horseshoe, a perpendicular median sella, and a posterior
erect lancet; skull with premaxillaries represented 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 but distinct ; fibula thread-like ; foot normal, the hallux
with two phalanges, the other toes with three.
Remarks. — The ffliinolophidse are the most widely distributed
EHINOLOPHUS
"137
of the Old World leaf-nosed bats, and the only family 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. Bhinolophus Lacepede, Tabl. des div. sousdiv. ordres et genres des
Mammiferes, p. 15 (femcm-equinum).
1836. Rhinocrepis Gervais, Diet. Pittoresque d'Hist. Nat. IV, pt. 2, p. 617
(attributed to Geoflroy and Ouvier, Mag. Encyclopedique, 1795,
tut the name does not occur in the paper alluded to).
1847. Aquias Gray, Proc. Zool. Soc. London, p. 15 (luctus and trifoliatus) .
1857. Bhinolophus Blasius, Saugethiere Deutschlands, p. 26.
1866. Phyllotis Gray, Proc. Zool. Soc. London, p. 81 (philippcnsis) not
Phyllotis Waterhouse, 1837.
1866. Caelophyllus Gray, Proc. Zool. Soc. London, p. 427 (ccelophyllus).
1878. Bhinolophus Dobson, Gatal. Chiropt. Brit. Mus., p. 100.
1901. Euryalus Matschie, Sitz.-Ber. Gesellsch. naturforsch. Preunde,
Berlin, p. 225 (mehelyi).
1904. Euryalus Matschie and Andersen, Sitz.-Ber. Gesellsch. naturforsch.
Freunde, Berlin, p. 71 (euryala group).
1907. Bhinolophus Miller, Families and Genera of Bats, p. 108, June 29,
1907.
Type species. — Vespertilio ferrum-equinum Schreber.
Geographical distribution. — Same as that of family (p. 136).
Characters. — Dental formula: i^ ^, c^J, pi» |~? , m |^f = 32.
2, 2, XI-* 1 oo o <j
Upper incisor very small, but usually well formed and with
distinct rounded crown with slight cusp on inner side. Lower
incisors trifid, 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 (pm 3 ) and
middle lower premolar (pm 3 ) small, functionless, usually crowded
quite out of tooth-row. Other teeth showing no special peculiari-
ties ; m 1 and m 2 without hypocone, m 3 with five cusps and
three commissures (in many species a rudimentary fourth com-
missure), the crown area much more than half that of m 1 or
m 2 . Skull with large brain-case and much shortened, globular ly
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,
extending 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
Bhinolophus are at once recognizable by the presence of the nose-
138
CHIEOPTERA
leaf and absence of tragus. The skull differs from that of all
other members of the fauna in the short, globularly inflated
rostrum and long, projecting maxillaries, between which lie the
FlS. 26.
Noseleaf of Rhiaolophvx ferrum-equinum (a), R. hipposideros (b), R. euryale (c),
and R. blasii (d). Nat. size.
horizontal free premaxillaries (often lost in prepared specimens),
About 100 forms have been described,* eight of which occur in
Europe.
KEY TO THE EUROPEAN FOBMS OP BHINOLOPHUS.
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) R. ferrum-eqitimmi, p. 139.
Wing relatively long ; forearm 54-58 mm.,
longest finger 84 to 92 mm. (Southern
and central Continental Europe) R. f. ferrum-egvimim, p. 142.
Wing relatively shorter ; forearm 52 to
55 mm., longest finger 83 to 88 mm.
(England) R. /• imidanus, p. 147.
* See Andersen, Ann. and Mag. Nat. Hist., 7th ser., svi, pp. 648-662.
RHIN0L0PHUS 139
Forearm under 43 mm. ; oondylobasal length
of skull under 16 mm. ; sella ouneate ;
large upper premolar not in contact
with canine (Lesser Horseshoe) B. hipposideros, p. 147.
Greatest length of skull 14-5 to 15-5 mm.
(Mediterranean region) r. ;,. minimus, p. 151.
Greatest length of skull more than
15 '5 mm.
Forearm 36-3 to 39 mm. (England and
Ireland) r. u. minutus, p. 154.
Forearm 39 to 41 -7mm. (Central Europe) r. /,. hipposideros, 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) R. 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) ; point of lancet gradually
narrowed, never linear R. euryale, 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 R. mehelyi, p. 159.
RHINOLOPHUS FERRUM-EQUINUM Schreber.
(Synonymy under subspecies.)
Geographical distribution. — From southern Japan and China,
through the Himalayas, the Mediterranean sub-region (exclusive
of Egypt), and central Europe to southern England (Andersen).
Diagnosis. — Size largest of the European species (forearm
more than 50 mm., condylobasal length of skull about 21 mm.,
mandible, 15 to 16 mm.); noseleaf with horseshoe less than
10 mm. wide, the sella pandurate, the connecting process low,
abruptly rounded ; fourth finger with first phalanx considerably
more than half as long as second ; large upper premolar broadly
in contact with canine, the small premolar minute (sometimes
absent), completely external to tooth-row.
External characters. — Size large and form rather heavy
(among the European members of the genus). General outline
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 tip of lancet. Sella broadly rounded at tip, noticeably con-
stricted somewhat above middle, the resulting outline pandurate.
Connecting process rising slightly but evidently above level of
sella, its upper border straight anteriorly, its tip rather abruptly
rounded off. Ear large, extending when laid forward slightly
beyond extremity of muzzle, abruptly narrowed to a rather acute
recurved tip ; antitragal lobe less than half as high as conch,
its width about equal to its height, its upper border nearly
horizontal. Wings broad, the membrane attached to ankle.
First phalanx of fourth 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
produced 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 paler 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 occur the extremes are more often met with,
evidently representing two dichromatic phases. Underparts
usually rather lighter than back but never approaching whitish.
SJcull. — General outline of skull long and narrow, the breadth
of brain-case much less than twice that across canines, and con-
tained about 2 }j times in greatest length ; zygomata projecting
slightly beyond general outline ; interorbital region greatly con-
stricted and conspicuously hour-glass shaped ; occipital portion of
brain-case noticeably overhanging foramen
magnum and marked off from main portion
by a slight transverse depression corre-
sponding to suture between parietals and
unusually large interparietal ; sagittal
crest well developed, extending forward to
narrowest portion of interorbital region :
lambdoid crest low but evident ; auditory
bullte 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 wider
anteriorly than posteriorly, about 1£
times as long as wide, its rounded
anterior margin at level of postero-
internal angle of to 2 ; palate with an
evident emargination on each side ex-
tending between m 3 and mesopterygoid space ; anterior palatal
emargination extending back to line joining protocones of
anterior molars ; premaxillary ligulate, somewhat wider pos-
teriorly than anteriorly, the inner 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^&sg&A
Rhinolophusferrum-equinum .
Nat. size.
RHINOLOPHUS
141
bottom of palatal emargination 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 premolars 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 anterior border of premaxillary, the space between the
two nearly double diameter of crown. Lower incisors strongly
imbricated, forming a short, very convex row between canines,
their crowns longer than high, compressed (the outer tooth less
than the inner), deeply and equally trifid. 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 cingulum about three-
fourths height ; cross section of shaft triangular, the inner surface
flattened, slightly concave near cingulum, the posterior edge
trenchant, the anterior edge narrowly sub-trenchant, 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
oppose front of upper canine in mastication. Anterior upper
premolar minute, resembling upper incisor in both size and form,
entirely external to tooth-row, occasionally absent. Posterior
upper 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-row. Upper molar with large though not unusually high
protocone, behind which in m 1 and m 2 there is a low-lying
heel (best developed in m 1 ), but no indication of a true hypocone ;
paracone and metacone well developed, the latter slightly the
higher in m 1 and m 2 , this reversed in m 3 ; styles and commis-
sures in m 1 and m 2 well developed, forming a normal VV-pattern ;
in m 3 the metastyle and fourth commissure are absent, and the
third commissure is much reduced in length ; crown area of m 3
142
CHIEOPTBEA
about two-thirds that of wr ; lower molars with protoconid
noticeably higher than the other cusps, the posterior triangle
slightly the wider of the two in m 1 and ro 2 , a little the narrower
in m 3 ; behind entoconid the cingulum forms a minute though
evident accessory cusp.
Remarks. — 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 It. mehelyi, which approaches it in size, the connecting process
is sharply pointed above. Two geographical races are known, one
occupying the Continental range of the species, the other confined
to Great Britain.
Rhinolophus ferrum-equinum ferrum-equinum Schreber.
1774. Vespertilio ferrum-equinum Schreber, Saugthiere, i, pi. lxii, upper
figures ; description, i, p. 174 under name : Die Hufeisennase
(part). Prance ; based primarily on Daubenton.
1776. Vespertilio equinus P. L. S. Miiller. Natursyst. Suppl. u. Regist.-
Band, p. 20 (part), Prance.
1779. [Vespertilio] perspicillatus ~Blumenba,ch, Handb. d. Naturgesch., p. 75
(part : included the leaf -nosed bats of Europe and South America).
1785. [Vespertilio'] ungula Boddaert, Elenchus Animalium, i, p. 71
(Burgundy).
1792. Vesp[ertilio] fer[rum]-equ[i'ium] major Kerr, Anim. Kingd., p. 99
(not V. molossus major Kerr, I.e., p. 97), Prance.
1798. Vespertilio hippocrepis Schrank, Pauna Boica, I, p. 64 (Renaming
of ferrum-equinum, Schreber) . '
1803. Bhinolophus major Geoflroy, Catal. Mamm. Mus. Nat. d'Hist. Nat.,
Paris, p. 56 (Burgundy).
1813. Bhinolophus unihastatus Geoffrey, Ann. Mus. d'Hist. Nat., Paris,
xx, p. 257 (Prance).
1829. ? Bhinolophus unifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ.
Thierwelt, I, p. 104 (nomen nudum).
1857. Bhinolophus ferrum-equinum Blasius, Saugethiere Deutschlands
p. 31.
1863. [Rhinolophus ferrum-equinum] a var. germanicus Koch, Jahrb. des
Vereins fur Naturkunde im Herzogthum Nassau, xvin, p. 522
(Wiesbaden, Hessen-Nassau, Germany).
1863. [Bhinolophus ferrum-equinum] B var. italicus Koch, Jahrb. des
Vereins fur Naturkunde im Herzogthum Nassau, xvni, p. 523
(Italy).
1878. Bhinolophus ferrum-equinum Dobson, Catal. Ohiropt. Brit. Mus.,
p. 119 (part).
1885. Bhinolophus unihastatus, var. homorodalmasiensis Daday, Orvos-
Termeszettudomanyi ErtesetiS, Kolozsvar, x, p. 274 (Homorod-
Almas cave, Hungary).
1886. Bhinolophus unihastatus var. homorodalmasiensis Daday, Verhandl.
u. Mittheilungen des Siebenbiirgischen Vereins f iir Naturwissensch.
in Hermannstadt, xxxvi, p. 79.
1887. Bhinolophus ferrum-equinum var. homorodensis Daday, Brtekezesek a
Termeszettudomanyok Korebi'il, Budapest, xvi, pt. 7, p. 13
(Renaming of homorodalmasiensis) .
RHINOLOPHUS 143
1904. Rhinolophus ferrum-equinum obscurus Cabrera, Mem. Soc. Espafi.
Hist. Nat. Madrid, n, p. 257 (Valencia, Spain).
1905. Rhinolophus ferrum-equinum typicus Andersen, Proo. Zool. Soc.
London, 1905, n, p. 113, October 17, 1905.
1905. Rhinolophus ferrum-equinum obscurus Andersen, Proc. Zool. Soc.
London, 1905, n, p. 116, October 17, 1905.
1910. Rhinolophus ferrum-equinum, R. ferrum-equinum obscurus and R.
ferrum-equinum homorodensis Trouessart, Paune Mamm. d'Europe,
pp. 4-5.
Type locality. — Burgundy, France.
Geographical distribution. — Central and southern Continental
Europe.
Diagnosis. — Wing relatively long ; forearm, 54 to 58 mm. ;
third finger, 84 to 92 mm.
Measurements. — Average and extremes of six males from
Silos, Burgos, Spain : head and body, 62 ■ 1 (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, Prance : tibia,
24-1 (23-4-24-6); foot, 12-] (11-8-12-8); forearm, 554
(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
Rimini, Italy : 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, Wurtemberg, 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 examined. — One hundred and forty-one, from the following
localities : —
Portugal : Cintra, 1.
Spain : Silos, Burgos, 22 ; Granada, 4 ; Elche, Alicante, 3 ; San
Cristobal, Minorca, Balearic Islands, 4.
144
CHIEOPTEEA
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CHIEOPTEEA
France : Troubate, Hautes-Pyrenees, 6 ; St. Genies, Gard, 43 (Mottaz) ;
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.M. and Genoa); no exact locality, 1 ;
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 : Tubingen, Wiirtemberg, 1.
Switzerland : Geneva, 6 (U.S.N.M. and Mottaz) ; Boudry, Neuchatel,
2 (Mottaz) ; Tremona, Tioino, 1 ; Mendrisio, Ticino, 2 ; Lugano, Ticino, 3.
Austria-Hungary : Herkulesbad, Hungary, 6 ; Ofener Mts., Hungary, 1.
Remarks. — Spanish specimens of JRhinoloplius ferrum-equinwu
have been regarded by both Cabrera and Andersen at represent-
ing a peculiar race, B. f. obscurus, 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. Portugal.
0. Thomas (c&p).
98. 2. 2. 1.
6al.
Silos, Burgos, Spain.
N. & S. Gonzalez
(c).
G. S. Miller (c).
G. S. MiUer (c).
G. S. MiUer (a).
8. 7. 7. 38-43.
2 6, 2 9.
Silos, Burgos.
8. 8. 4. 1-4.
6,9.
Granada, 2250 ft.
8. 8. 4. 8-9.
2 6, 9.
Elche, Alicante, 20 m.
8. 8. 4. 5-7.
3,2 9.
San Cristobal, Minorca,
learic Islands.
Ba-
0. Thomas and
R. I. Pocock
(c & P).
0. 7. 1. 24-26.
<5al.
San Cristobal, Minorca.
0. Thomas and
R. I. Pocock
(c & P).
0. 7. 1. 68.
6 6.
Troubate, Hautes-Pyrenees,
0. Thomas (p).
6. 4. 1. 1-6.
France. (A. Robert.)
lal.
Meounes, Var.
Dr. K. Jordan
(c & p).
8. 3. 15. 1.
9al.
Turin, Italy.
Prof. BoneUi (p).
?al.
Isoverde, Genoa.
O.Thomas (c&p).
88. 12. 7. 1.
Skeleton.
Italy. (Prince Bonapan
ie.)
Tomes CoUection.
7. 1. 1. 727.
2<Jal.
Sicily.
cJal.
Barbaggio, Corsica.
Dr. C. I. Forsyth
6. 4. 14. 1.
6 al. Sassari, Sardinia.
<5, 2 ?. Mt. Goria Monastery, Crete.
9 al. Tiibingen, Wiirtemberg, Ger-
many.
9. Tremona, Ticino, Switzerland.
(E. H. Zollihofer.)
2 9. Mendrisio, Ticino. (E. H.
Zollihofer.)
3. Lugano, Ticino. (E.H. Zolli-
hofer.)
2 ci, 4 9. Herkulesbad, Hungary.
9 al. Ofener Mts., Budapest.
Major (c&p).
Marquis G. Doria 6. 12. 1. 11.
Miss D. Bate (c). 5. 12. 2. 1-3.
Dr. A. Giinther 66. 2. 1. 1.
(*)■
0. Thomas (p).
0. Thomas (p).
0. Thomas (p).
2. 8. 4. 1.
2. 8. 4. 2-3.
4. 4. 5. 1-3.
7. 9. 16. 1-6.
Hon. N. C. Roths
child (p).
Budapest Museum 94. 7. 18. 5.
(■)•
RHINOLOPHUS
147
Rhixolophus pbeeum-equinum insulands Barrett-Hamilton.
1910. Rhinolophus ferrum-equimminsulanus Barrett-Hamilton, Ann. and
Mag. Nat. Hist., 8th ser., v, p. 292, March, 1910. Type in British
Museum.
1910. Rhinolophus ferrum-equinum insulanus Trouessart, Faune Mamm.
d'Europe, p. 273.
Type locality. — Cheddar, Somersetshire, England.
Geographical distribution. — Central and southern England.
Diagnosis. — Wing relatively short ; forearm, 52 to 55 ; third
finger, 83 to 88.
• . Measurements. — Type (adult male) : head and body, 67 ; tail,
37 ; tibia, 23-4 ; foot, 11 ; forearm, 54 ; third finger, 86 ; 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,
69-9 (69-71) ; ear from meatus, 24-2 (23-25). Three females
from the type locality : head and body, 64, 66 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, p. 144.
Specimens examined. — Twenty-four, from the following localities in
England : — Cheddar, Somersetshire, 10 ; Wells, Somersetshire, 10 ; Dorset-
shire, 3 ; Bonchurch, Isle of Wight, 1.
6, 9. Cheddar, Somerset, J. A. Coward (c & p). 7. 1. 10. 1-2.
England.
6 al. Cheddar. J. A. Coward (o & p). 7. 1. 10. 3-10.
(J, 2 Sal. Wells. Hon. N.C.Rothschild 5.1.23.1-3.
(*)■
3 6, 6 juv., 9 st. Wells. Hon.N. C.Rothschild /l. 9. 3. 1-4.
(p). \2. 9. 6. 1.
6,9. Wells. Hon. N.C.Rothschild 11.1.3.3-4.
(*)■
■1 9. Dorset. W. M. Hardy (c & p). 11. 1. 3. 1-2.
1. Bonchurch, Isle of Rev.C. A.Bury (c&p). 11.1.3.388.
Wight.
RHINOLOPHUS hipposideros Bechstein.
(Synonymy under subspecies.)
Geographical distribution. — -From Gilgit 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.
Diagnosis. — Size small, the forearm less than 43 mm. in
length, condylobasal length of skull about 14 to 15 mm., mandible
about 10 mm.; noseleaf (fig. 26 6) with bluntly cuneate sella
And low, broadly rounded connecting process ; fourth finger with
l 2
148
CHIROPTBEA
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 Bhinolophus ferrum-equinum. General outline of
noseleaf narrower than in B. ferrum-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 species 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 5 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 oblique.
Wings and feet essentially as in B. ferrum-equinum.
Colour. — The colour does not differ appreciably from that of
Bhinoloplius ferrum-equinum.
Skull. — The skull differs from that of Bhinoloplius ferrum-
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|- times in greatest length it is probable that the difference
between the two animals is due rather to enlarged brain-case in
the smaller than to widened palate in the larger. Sagittal crest
slightly developed ; lambdoid crest obsolete. Nasal region
relatively more inflated than in B. 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 genera] resembling those of Bhinoloplius
ferrum-equinum, apart 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, and
slightly below that of posterior premolar ; lower canine corre-
spondingly short ; anterior upper premolar a well developed
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 premolar ; anterior and posterior lower pre-
RHINOLOPHUS
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 S. ferrum-
equinum. Large upper premolar
and upper molars essentially as in
B. ferrum-equinum except that in
tn 3 there is a distinct meta-
style and fourth commissure,
and area of tooth is nearly
equal to that of m 2 . Lower
molars as in Bhinolophus
ferrum-equinum, 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
30 mm., forearm, 34-5 to
41 • 7 mm., condylobasal length
of skull, 13 • 8 to 15 mm.
(greatest length 14-5tol6'2).
Details under subspecies.
Bemarhs. — 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. 28.
Ithinolophus hipposideros. Teeth X 10_
Rhinolophus hipposideros hipposideros Bechstein
1789. VespertiUo ferrum-equinum B. , Die kleine Huf eisennase, Bechstein.
Gemeinn. Naturgesch. Deutsehlands, i, 1st ed., p. 186.
1792. Vcsp[ertilio~] fer\rum\-eqit,\inum\ minor Kerr, Anim. Kingd., p. 99
(not V. molossus minor Kerr, I.e., p. 97) (France).
1800. VespertiUo hipposideros Bechstein, Thomas Pennant's Allgemeine
Uebersicht der vierfiissigen Thiere, n, p. 629.
1803. Bhinolophus minor Geoffroy, Oatal. Mamm. Mus. Nat. d'Hist. Nat.,
Paris, p. 57 (Neighbourhood of Paris).
1813. Bhinolophus bihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xx,
p. 259 (Neighbourhood of Paris).
1816. ? Phyllorhina minuta Leach, Syst. Catal. Spec. Indig. Mamm. and
Birds Brit. Mus., p. 5 (nomen nudum : " Small Leafnose").
* For further discussion of the subject see Mottaz, Mem. Soc. Zool. de
France, Paris, xx, pp. 21-22, September, 1907 ; Andersen, Ann. and Mag.
Nat. Hist., 7th ser., xx, pp. 384-389, November, 1907.
1 50 CHIEOPTBEA
1829. ? Bhinolophus bifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Burop.
Thierwelt, I, p. 104 (nomen nudum).
1857. ? Bhinolophus hipposideros Blasius.'Saugethiere Deutschlands, p. 29.
1863. [Bhinolophus hipposideros'] a var. typus Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xvm, p. 530 (Wiesbaden).
1863. [Bhinolophus hipposideros] var. alpinus Koch, Jahrb. des Vereins
fiir Naturkunde im Herzogthum Nassau, xvm, p. 530 (Alps).
1870. Bhinolophus eggentioffner Pitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lxi, Abth. i, p. 151
(MS. synonym of bihastatus).
1878. Bhinolophus hipposideros Dobson, Oatal. Chiropt. Brit. Mus., p. 117.
1885. Bhinolophus bihastatus var. kisnyiresiensis Daday, Orvos-Termesz-
ettudomanyi Ertesito, KojLozsvar, x, p. 274 (Kis-Nyires, Szolnok-
Dobaka, Hungary).
1886. Bhinolophus bihastatus var. kisnyiresiensis Daday, Verhandl. u.
Mittheilungen des Siebenburgischen Vereins fiir Naturwissensch.
in Hermannstadt, xxxvi, p. 80.
1887. Bhinolophus hipposideros var. troglophilus Daday, Ertekezesek a
Termeszettudomanyok Korebol, Budapest, xvi, pt. 7, p. 8 (Re-
naming of kisnyiresiensis).
1904. Bhinolophus euryale helvetica Bretscher, Vierteljahrsschrift der
Naturforsch. Gesellsch. in Zurich, xlix, p. 256. See Mottaz,
Bull. Soc. Zool., Geneve, ±., p. 172, 1908 (Baar, Zug, Switzerland).
1905. Bhinolophus hipposiderus typicus Andersen, Proc. Zool. Soc. London,
1905, n, p. 141, October 17, 1905.
1910. Bhinolophus hiijposiderus Trouessart, Faune Mamm. d'Europe, p. 9.
Type locality. — France.
Geographical distribution. — Central Europe, north of the Alps,
east through Armenia and north-west Persia to the Himalayas.
Diagnosis. — Forearm* usually 39 to 41 • 7 mm.; greatest
length of skullf about 16 mm.
Measurements. — Two males from Strass 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 European specimens are
given by Andersen (I.e. p. 142) : 1ST. Bulgaria (1), 39 ; Roumania
(13), 39 to 41-2 ; Transsylvania (2), 40 to 41 ; S. Carpathians
(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),! 37 ' 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 Table, p. 152.
* Bases of metacarpals included,
t Prom back of occiput to front of canine.
X Measured by Chas. Mottaz (base of metacarpals included) and verified
by Gerrit S. Miller.
RHINOLOPHUS 151
Specimens examined. — One hundred and seventeen, from the following
localities: —
France : Dions, Gard, 1 (Mottaz) ; St. Gergues, Haute-Savoie, 1.
Germany : Strass, near Burgheim, Bavaria, 3 ; Mainz, 1 (Strassburg) ;
Bitsoh, Alsace, 1 (Strassburg).
Austria-Hungary : Hatszeg, Hunyad, Hungary, 1.
Switzerland: Near Geneva, 94 (B.M., U.S.N.M. and Mottaz); St.
Moritz, 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.).
£. St. Cergues, Haute-Savoie, A.Robert (c &p). 5.4.9.1.
France.
2 9. Burgheim, Bavaria, 375 m. Lord Lilford (p). 11. 1. 1. 117-
Germany. 118.
<J. Hatszeg, Hunyad, Tran- C.G.Danford(c). 8.11.8.1.
sylvania.
3 6. Grand Pre, Geneva, Swit- C. Mottaz (c & i>). 6. 2. 6. 1-3.
zerland.
1. St. Moritz, Grisons. Leon 0. Galliard 75. 9. 20. 3.
(o & p).
3 6, 2 9, imm. Roggwil, Thurgau. (E. H. 0. Thomas (p). 2. 8. 4. 4-9.
ZolUkofer.)
?al. Thurgau. (E.H. ZolUkofer.) 0. Thomas (p). 2.8.4.55.
Rhinolophus hipposideros minimus Heuglin.
1861. Rhinolophus minimus Heuglin, Nov. Act. Acad. Caes. Leop.-Carol.,
xxix, Abhandl. vin, p. 6 (articles separately paged) (Keren,
Abyssinia).
1863. [Rhinolophus hipposideros'] y var. pallidus Koch, Jahrb. des Vereins
fiir Naturkunde im Herzogthum Nassau, xviii, p. 531 (Mediter-
ranean region).
1904. Rhinolophus phasma Cabrera, Mem. Soc. Espan. Hist. Nat., Madrid,
ii, p. 252 (Madrid, Spain).
1904. Rh[inolophus] h[ipposiderus] minimus Andersen, Ann. and Mag. Nat.
Hist., 7th ser., xiv, p. 456, December, 1904.
1905. Rhinolophus hipposiderus minimus Andersen, Proc. Zool. Soc.
London, 1905, n, p. 140, October 17, 1905.
1910. Rhinolophus hipposiderus minimus Trouessart, Faune Mamm.
d'Europe, p. 10.
Type locality. — Keren, Abyssinia.
Geographical distribution. — Mediterranean region.
Diagnosis. — Forearm usually 34 ■ 7 to 38 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), 36 • 3 ; Cyprus (6),
34-7 to 37-7; Malta (8), 36 to 37; Ostia, Italy (2), 35*7' to
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 ( <J ),
36-4; Siena (3 $ ), 36 '8 to 38; Marseilles, France (?), 38;
Dions, Gard, France ( <J ), 34 ■ 8, ( ° ), 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 -
(36 '2-38). Mottaz collection: St. Genies, Gard, France (15,
152
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both sexes), 36 • 4 to 38 ■ 2) ; Sardinia (2 ? ), 37 ■ 4 to 37 ■ 8. For
cranial measurements see Table, p. 153.
Specimens examined. — Eighty-one, from the following localities : —
Spain: Inoa, Majorca, Balearic Islands, 1; San Cristobal, Minorca,
Balearic Islands, 3 ; Elche, Alicante, 2 ; Silos, Burgos, 5 ; Seville, 1.
Portugal : Cintra, 1.
France: Ax-les-Thermes, Ariege, 5; St. Genies, 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.); Paenza, 1
(U.S.N.M.) ; Borne, 3 ; Ostia, Rome, 2 ; Picuzza, Sicily, 6.
Sardinia : No exact locality, 2 (Mottaz) ; Zimmigas, Siliqua, 2.
Corsica : Dintomi di Patrimonio, 3 ; no exact locality, 1.
Malta : 7.
9. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 1-2.
Islands. Pocock (c & p).
6, 9. San Cristobal, Minorca. 0. Thomas and R. I. 0. 7. 1. 27-28.
Pocock (c & p).
1 al. San Cristobal, Minorca. 0. Thomas and R. I. 0. 7. 1. 69.
Pocock (c & pi
9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 11.
6. Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 12.
£ al. Cintra, 500 m. Portugal. O. Thomas (c & p). 98. 2. 2. 57.
6, 9. Ax-les-Thermes, Ariege, G. S. Miller (o). 8.8.4.124-
2400 ft. Prance. 126.
S. Borne, 37 m. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 42.
if)-
2. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 5-6.
6 9. Picuzza, Sicily. (A.Robert.) 0. Thomas (p). jg ®" * - ]^ 16 -
(5, 9 al. Zimmigas, Siliqua, Sardinia. Marquis G. Doria (p). 6.12.1.20-21.
(R. Meloni.)
3 <$, 2 9. Malta. (J. Micalleff.) Lord Lilford (p). 11. 1. 1. 119-
123
2. Malta. Lord Lilford (p). 95. 3.' 2. 1-2.
Rhinolophus hipposideeos minutus Montagu.
1808. Vespertilio minutus Montagu, Trans. Linn. Soc. London, ix, p. 163.
1905. Rhinolophus hipposiderus minutus Andersen, Proc. Zool. Soc.
London, 1905, u, p. 142, October 17, 1905.
1910. Rhinolophus hipposiderus minutus Trouessart, Faune Mamm.
d'Europe, p. 10.
Type locality. — Wiltshire, England.
Geographical distribution. — England and Ireland.
Diagnosis. — Forearm, 36 '3 to 39 mm . • greatest length of
skull about 16 mm.
Measurements. — Forearm in 30 English 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; Bowdon, 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.
5.
1. 24. 1.
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'. 2. 21. 1.
4.
11. 6. 1-2.
11
. 1. 3. 8-9.
7.
1. 1. 289-
291.
RHIN0L0PHUS 155
3 i. St. Asaph, Denbighshire, Charles Oldham (c & p). 11. 1. 3. 5-7.
Wales.
1. Conway, Carnarvonshire. Sir W. Jardine (c & p). 60. 9. 17. 1.
?. Great Grimsby, Lincoln- G. Barrett-Hamilton (p). 11. 1. 2. 98.
shire, England. (Caton
Haigh.)
6. Bowdon, Cheshire. (T.A. G. Barrett-Hamilton (p). 11.1.2.99.
Coward.)
i al. Hope End, Herefordshire. N. C. Hewitt (c & p).
6 al. Wells, Somerset. S. Lewis (o & p).
1. Devizes, Wiltshire. J. E. Harting (c & p).
6, 9 al. Zeals, Wiltshire. F. Norgate (c & p).
2 ?. Devonshire. Oxley Grabham (c & p)
3. Ragley House, Warwick- Tomes Collection,
shire.
RHINOLOPHUS EURYALE Basius.
1853. Bhinolophus euryale Blasius, Wiegmann's Arehiv fur Naturgesch.,
1853, i, p. 49 (Milan, Italy).
1857. Bhinolophus euryale Blasius, Saugethiere Deutschlands, p. 35.
1878. Bhinolophus euryale Dobson, Catal. Chiropt. Brit. Mus., p. 116.
1904. E[uryalus] toscanus Andersen and Matschie, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 77 (Caverna di Parignano, Mt.
Pisani, Italy).
1904. E[uryalus'] atlanticus Andersen and Matschie, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 77 (St. Paterne, Indre-et-Loire,
France).
1904. E\uryalus~\ cabrerse Andersen and Matschie, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 78 (Alcala de Henares, Madrid,
Spain).
1910. Bhinolophus euryale, B. euryale atlanticus, B. euryale toscanus,
B. euryale cabrerai Trouessart, Faune Mamm. d'Europe, pp. 5-7.
Type locality. — Milan, Italy.
Geographical distribution. — Southern Europe from Portugal
to 1 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 13, 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
phalanx 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 characters. — Size intermediate between that of the
Greater and Lesser Horseshoes. General outline of noseleaf
(fig. 26 c) about as in Bhinolophus ferrum-equinum ; sella parallel
sided, rounded off above, connecting process sharply linear-
pointed, rising conspicuously above sella ; lancet with slight con-
striction above middle, beyond which the tip narrows gradually to
a bluntly cuneate point. Ear when laid forward extending about
■5 mm. beyond extremity of muzzle, its tip less attenuate and less
noticeably curved backward than in Bhinolophus ferrum-equinum
156
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and B. hipposideros ; antitragal lobe about balf as bigb as conch,
its width about equal to its height. Wing peculiar in the
shortening of the first phalanx of fourth finger to a little more
than one-third that of second (average of 10 specimens from Gard,
France: first phalanx, 6 - 6 mm.; second phalanx, 17 '4 mm.;
ratio of first to second, 38 +)• Foot averaging slightly more
than half as long as tibia.
Colour. — While essentially as in BMnolophus ferrum-equinum
and B. 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
B. mehelyi.
Skull. — In both size (greatest length about 19 mm.) and form
the skull is somewhat intermediate between that of BMnolophus
ferrum-equinum and B. hipposideros. It resembles or surpasses
the latter in the breadth of brain-case relatively to narrow
maxillary region, but more nearly agrees with the former in the,
nearly parallel-sided mesopterygoid space, the anterior border of
which is separated from posterior molars by well defined palatal
emarginations. The mesopterygoid space is, however, shorter in
proportion to its width than in B. ferrum-equinum. Floor of
brain-case between cochleee less narrowed than in the preceding
species. Nasal region slightly less inflated than in B. hipposideros,
and inflated area relatively shorter, its posterior border rising
abruptly above interorbital level.
Teeth. — In all respects the teeth closely resemble those of
BMnolophus ferrum-equinum (apart from their smaller size), except
that the upper canine is relatively less robust, the anterior upper
premolar is less reduced (its crown area about double that of
upper 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 examined. — About 130, from the following localities : —
Portugal : Cintra, 6.
Spain: Villalba, Lugo, 1; Madrid, 2; Silos, Burgos, 1.
France : St. Paterae, Indre-et-Loire, 3 (B.M. and U.S.N.M.) ; St. Genies,
Gard, about 50 (Mottaz) ; Gapeau Eiver, 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 ; VeUetri, Borne, 5 (U.S.N.M.) ;
Nieotera, Calabria, 1 ; Marsala, Sicily, 2.
Sardinia : Mount Gennargentu, 3 (U.S.N.M.).
Austria-Hungary : Ofener Mountains, 2 ; Orsova, 1.
Dalmatia : Zara, 1.
Greece : Missolungi, Acarnania, 2 (U.S.N.M.).
Bemarks. — BMnolophus euryale is so readily distinguishable
from all the other European members of the genus, except
B. mehelyi, as to require no special comparisons. From B. mehelyi
it is most easily distinguished by the form of the lancet and
EHINOLOPHUS
159
antitragus, together with its rather smaller size and usually
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 to be within the
range of normal individual variation.
2 9. Cintra,500m. Portugal. O. Thomas (c & p). 98. 2. 2. 2-3.
2 <?, 2 ? al. Cintra, 500 m. 0. Thomas (c & p). 98. 2. 2. 53-56.
9 al. Villalba, Lugo, N.W. Dr. V. L. Seoane (p). 94. 1. 1. 1.
Spain.
i, 9 al. Madrid. A. Cabrera (p). 5. 2. 3. 1-2.
1 al. St. Paterne, Indre-et- Boyal Army Medical 9. 1. 4. 9.
Loire, France. College (p).
9 al. St. Paterne, Indre-et- G. E. Dobson (p). 80. 12. 14. 3.
Loire.
9 al. Gapeau Eiver, Var. Dr. K. Jordan (c & p). 8. 3. 15. 2-10.
3 <5, 9 al. Finalborgo, Liguria, Marquis G. Doria (p). 6.12.1.14-17.
Italy. (A. Gagero.)
2 al. Monte Pisanino, Liguria. Lord Lilford (p). 73.1.8.6.
■2 6. Eome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 40-41.
(*)■
6 al. Nicotera, Calabria. Florence Museum (e). 85.7.6.1.
2 6. Marsala, Sicily. (A. 0. Thomas (p). 6. 8. 4. 10-11.
Robert.)
S, 9 al. Ofener Mts., Budapest. Budapest Museum (e). 94. 7. 18. 2-3.
9. Orsova, Hungary. Hon. W. Bothschild 7. 9. 16. 7.
(*)■
9. Zara, Dalmatia, 50 m. Lord Lilford (p). 11. 1. 1. 128.
[Kolombatovic.)
1 al. S. Europe. Purchased (Parreys). 47. 5. 27. 44.
RHINOLOPHUS MEHELYI Matschie.
1901. Bhinolophus mehelyi Matschie, Sitz.-Ber. Gesellsch. Naturforsch.
Freunde, Berlin, p. 225 (Bucharest, Boumania).
1904. Bhinolophus carpetanus Cabrera, Mem. Soc. Espafi. Hist. Nat., n,
p. 254 (Madrid, Spain).
1910. Bhinolophus euryale mehelyi and B. carpetanus Trouessart, Faune
Mamm. d'Europe, pp. 7-8.
Type locality. — Bucharest, Roumania.
Geographical distribution. — Roumania, southern France
(Gard), Sardinia, central Spain. Details of distribution not
known.
Diagnosis. — Like Bhinolophus euryale but larger (forearm,
48 • 6 to 51 • 4 ; upper tooth-row about 7 mm.); noseleaf with
lancet abruptly narrowed to a linear tip ; ear with antitragal
lobe relatively broad and low ; fourth finger with first phalanx
decidedly more than one-third as long as second (ratio about 44) ;
colour usually paler than in the related animal.
<j External characters. — Slightly larger and more robust than
Bhinolophus euryale, a difference especially noticeable in freshly
killed individuals. Noseleaf as in B. euryale, except that the
lancet is very abruptly narrowed above middle to a distinctly
linear tip. Ear as in B. euryale but broader, the antitragal lobe
160
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162 CHIROPTEEA
scarcely half as high as conch, its width slightly greater than
height. Wing differing from that of the related animal in the
less degree of shortening of the first phalanx of fourth finger as
compared with second (average of ten specimens from Gard,
Prance, 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 Rhinolophus 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-grey.*
Skull and teeth. — Except for its slightly greater average size
(greatest length about 20 mm.) the skull agrees with that of
Rhinolophus 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 carpetanus).
Pbance : Near St. Gerries, Gard, about 50, skins and in flesh (Mottaz) .
Sardinia: Sassari, 3 (B.M. and U.S.N.M.).
Boumania : Bucharest, 1 (U.S.N.M.) ; Dobrudscha, 1 (Mottaz).
Remarks. — At first sight this species appears very similar to
Rhinolophus 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 Rhinolophus mehelyi may prove to be more
strictly confined to the Mediterranean region than the smaller
form.
9. Madrid. A. Cabrera (p.) 5. 2. 3. 1.
(Paratype of R. carpetanus Cabrera.)
2 <5 al. Sassari, Sardinia. Marquis G. Doria(p). 6.12.1.18-19.
rhinolophus blasii Peters.
1857. Rhinolophus clivosus Blasius, Siiugethiere Deutschlands, p. 33. Not
of Riippell, 1824 (Italy, Sicily, Istria and Dalmatia).
1866. Rhinolophus blasii Peters, Monatsber. k. Akad. Wissensch. Berlin,
p. 17 (Renaming of clivosus Blasius).
1878. Rhinolophus blasii Dobson, Catal. Chiropt. Brit. Mus., p. 117.
1910. Rhinolophus blasiusi Trouessart, Paune Mamm. d'Europe, p. 9.
Type locality. — South-eastern Europe.
Geographical distribution. — Eastern portion of the Mediter-
ranean region : Cyprus, Greece, Italy?
Diagnosis. — Size essentially as in Rhinolophus euryale ; nose-
* When seen by candle-light flying in caverns these bats are said to
appear entirely white.
KHINOLOPHUS
163
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Locality.
R. mehelyi.
Roumania: Dobrudscha .
Bucharest
Sardinia : Sassari .
France: near Nimes, Gard
if j>
Spain : near Madrid
R. blasii.
Cyprus ....
Greece: Nauplia
M 2
164 CHIROPTERA
leaf with cuneate sella and very high, sharply pointed connecting
process ; fourth finger with first 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 Rhinolophus euryale.
Skull. — In general the skull resembles that of Rhinolophus
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 Rhinolophus euryale.
Small upper premolar slightly less reduced, perfectly in the
tooth-row, but showing no tendency to develop 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 members
of the genus in the approximately equal crown areas of the two
larger teeth, the anterior subterete, the posterior with trapezi-
form section ; shaft of posterior tooth with diameter in axis of
tooth-row much less than transverse diameter, the cusp when
viewed from the side appearing to rise from middle of crown with
noticeable flat area before and behind it.
Measurements. — 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, 45-3 (44-6-
47) ; thumb, 7 '5 (7-8) ; third finger, 70-3 (69-72) ; fifth finger,
57'7 (56-60); ear from meatus, 19 -7 (19-20) ; ear from c'rown,
15-5 (15-16-4); width of ear, 14-7 (14-15). Adult from
Nauplia, Greece : tibia, 18 • 8 ; foot, 9 ■ 6 ; forearm, 44 • 6 ; thumb,
8 ; third finger, 69 ; fifth finger, 57. For cranial measurements
see Table, p. 163.
Specimens examined. — Five, from the following localities : —
Cyprus : No exact locality, 4.
Gbeecb : Nauplia, 1 (U.S.N.M.).
lieimirkt. — 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 require no special comparisons.
Its range appears to be strictly confined to the eastern portion
of the Mediterranean region, not extending west of Italy.
VESPEKTILIONINJE 165
Family VESPERTILIONIDSE.
1821. Vespertilionidm Gray, London lied. Repos., xv, p. 299, April 1, 1821
(part).
1857. Vespertiliones Blasius, Saugethiere Deutschlands, p. 37.
1878. Vespertilionidm Dobson, Catal. Chiropt. Brit. Mus., p. 167 (except
the genera NataVus and Thyroptera).
1907. Vespertilionidse Miller, Families and Genera of Bats, p. 195, June 29,
1907.
Geographical distribution. — Eastern and western hemispheres
to the limits of tree growth ; in the Atlantic to the Azores, and
in the Pacific 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 by nasal
branches only, the two bones very early 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 Bhinolophidse ; fibula very slender, not adding appreciably to
strength of leg ; foot normal, the toes slender ; tail not project-
ing conspicuously beyond membrane.
Remarks. — 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 VESPERTILIONIJST^E.
1878. Vespertiliones Dobson, Catal. Chiropt. Brit. Mus., p. 168 (except
genera Kerivoula and Harpiocephalus).
1907. Vespertilioninse Miller, Families and Genera of Bats, p. 197, June 29,
1907.
Geographical distribution. — 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 presternum ; six ribs connected
with sternum ; seventh cervical vertebra not fused with first
* The American Nycteris cinerea has been recorded (under the name
Vespertilio pruinosus) from South Bonaldshay, Orkney Islands, but the
occurrence seems open to question (see Wolley, The Zoologist, vn, p. 2343,
1849; viii, pp. 2695-96, 2813-14, 1850; Barrett-Hamilton, Hist. Brit,
llamm., i, pp. 222-224, March, 1911).
A specimen of another North American member of this genus
IN. borealis), bearing the label: "Villevgque (Maine & Loire), 8. 1. 89," is
figured 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 CHIEOPTEEA
dorsal ; scapula with coracoid curved outward ; nostrils simple ;
lower incisors in all known genera, 3-3.
Remarks. — The sub-family Vespertilioninse contains all but
eight of the known genera of Vespertilionidse, and all but one,
Miniopterus, of the eight found in Europe. It is the central,
least specialized portion of the family.
KEY TO THE EUROPEAN GENERA OP VESPEBTILION1D&].
Cheek teeth jtjj Myotis, p. 166.
Cheek teeth less than ||.
Upper premolars 1-1.
Rostrum noticeahly concave on each side of middle
line ; nares extending about halfway to inter-
orbital constriction ; palatal emargination
broader than deep Vespertilio, 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 Pipistrellus, p. 202.
Upper surface of rostrum concave ; ears
joined Barbastella, p. 263.
Genus MYOTIS Kaup.
1829. Myotis Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, I,
p. 106 (myotis).
1829. Nystactes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, I,
p. 108 (bechsteimi).
1830. Leuconoe Boie, Isis, p. 256 (daubentonii).
1839. Vespertilio Keyserling and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, I, p. 306 (Not Vespertilio Linnaeus, 1758).
1841. Selysius Bonaparte, Iconogr. Faun. Ital. i., Introd. alia Classe
Mamm., p. 3 (mystacinus).
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 (hasseltU, macellus = adversus and blepotis).
MYOTIS 167
1849. Tralatitus Gervais, Diet. Univ. d'Hist. Nat., xiii, p. 213 (Modifica-
tion of Trilatitus).
1856. Brachyotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden,
neue Folge, n, p. 131 (mystacinus, daubentonii, and dasyeneme).
Not Brachyotus Gould, 1837.
1856. Isotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, neue
Folge, ii, p. 131 (nattereri and emarginatus).
1857. Vespertilio Blasius, Saugethiere Deutschlands, p. 78. Not Vespertilio
Linnaeus, 1758.
1866. Tralatitius Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvn, p. 90,
February 1866 (Modification of Trilatitus).
1867. Ptemopterus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin,
p. 706 (sub-genus of Vespertilio = Myotis, type lobipes = muricota) .
1870. Exochurus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 75 (macrodactylus, horsfleldii =
adversus and macrotarsus).
1870. Aeorestes Pitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 427 (villosissimus, albescens, and
nigricans).
1870. Comastes Pitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.-
Naturwiss. Classe, lxii, p. 565 (capaccinii, megapodius, dasyeneme,
and limnophilus).
1878. Vespertilio Dobson, Catal. Chiropt. Brit. Mus., p. 284. Not Vespertilio
Linnaeus, 1758.
1897. Myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 382,
October, 1897.
1899. Euvespertilio Acloque, Paune de Prance, Mammiferes, p. 38 (emargin-
atus, mystacinus, murinus = myotis, nattereri, and bechsteinii).
1907. Myotis Miller, Families and Genera of Bats, p. 201, June 29, 1907.
Type species. — Vespertilio myotis Borkhausen.
Geographical 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 g, c J--1, pm jg, m g = 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.
Remarks. — The genus Myotis 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 CHIHOPTEKA
KEY TO THE EUBOPEAN SPECIES OP MYOTIS.
Size large (forearm 53 to 64 mm., oondylobasal
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.
Oondylobasal 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. myotis, p. 192.
Oondylobasal 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. oxygnathus, p. 199.
Size small or medium (forearm 34 to 47 mm., oondy-
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-oase more
than half greatest length; crown area of
molars relatively small ; upper molars with
evident protoconule.
Forearm about 47 mm. ; oondylobasal length
of skull about 16 mm. (Central and
southern Europe) M. dasycneme, p. 189.
Forearm less than 45 mm. ; oondylobasal length
of skull never more than 15 mm.
Tibia and adjacent membrane densely furred;
forearm about 42 mm. ; oondylobasal
length of skull 14-0 to 14-8 mm. (Medi-
terranean region) 71/. capaccinii, p. 187.
Tibia and adjacent membrane not furred ;
forearm about 35 mm. ; oondylobasal
length of skull 13-2 to 13 -8 mm.
(Distribution general) M. daubentonii, 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. ; oondylobasal length
of skull about 15 mm. ; posterior border
of ear conch with deep, almost angular
emargination slightly above middle M. emarginatus, p. 177.
Forearm about 34 mm. ; oondylobasal 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 169
Combined length of tibia and foot less than
25 mm. ; oondylobasal length of skull
14 • to 14 • 6 mm. ; ear narrow, its width
about 10 mm. ; tragus conspicuously
more than half as high as conch 21. nattercri, p. 174.
Combined length of tibia and foot about
30 mm. ; oondylobasal length of skull
16 to 17 mm. ; ear broad, its width about
14 mm. ; tragus scarcely half as high as
conch 31. bechsteinii, p. 179.
MYOTIS MYSTACINUS Kuhl.
1819. Vespertilio 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, Prance).
1834. Vespertilio humeralis Baillon, Mem. Soc. Royale d'Emulation
d'Abbeville, 1833, p. 50 (Abbeville, Somme, Prance).
1837. Vespertilio schinsii Brehm, Ornis, Heft in, p. 27 (Renthendorf,
Thiiringen, Germany).
1843. V[espertilio] schranhii Wagner, Wiegmann's Archiv fur Naturgesch.,
tx, Bd. n, p. 25 (Munich, Germany ? See Pitzinger, Sitzungsber.
kais, Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 219, 1871).
1857. Vespertilio mystacinus Blasius, Saugethiere Deutschlands, p. 96.
1863. [Brachyotus mystacinus] var. nigricans Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xvin, p. 444 (Wiesbaden,
Nassau, Germany).
1863. [Brachyotus mystacinus] var. rufofuscus Koch, Jahrb. des Vereins
fur Naturkunde im Herzogthum Nassau, xvin, p. 444 (Wiesbaden,
Nassau, Germany).
1863. [Brachyotus mystacinus] var. aureus Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xvin, p. 445 (Breisgau,
Germany).
1869. [Vespertilio mystacinus] var. nigricans Patio, Faune Vert. Suisse,
i, p. 92 (Switzerland). Not of Koch, 1863.
1869. [Vespertilio] lugubris Patio, Paune Vert. Suisse, i, p. 93 (Alternative
for nigricans Patio).
1871. Vespertilio mystacinus, nigro-fuscus Pitzinger, Sitzungsber. kais.
Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 217 (Renaming of V. schinzii Brehm).
1878. Vespertilio mystacinus Dobson, Catal. Chiropt. Brit. Mus., p. 314.
1900. Myotis mystacinus Mehely, Monogr. Chiropt. Hungariae, p. 200.
1910. Myotis mystacinus Trouessart, Paune Mamm. d'Europe, p. 33.
Geographical distribution. — Entire Continent of Europe north
to about the limits of tree growth ; west to Ireland ; east into
Asia.
Diagnosis. — Smallest species of European Myotis (forearm
about 34 mm., longest finger about 60 mm., oondylobasal length
of skull 12-6 to 13 "2 mm.) ; ear moderately long, extending 1 to
2 mm. beyond tip of muzzle when laid forward, its posterior
border with shallow inconspicuous emargination ; foot about half
as long as tibia ; wing membrane extending to base 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 anterior base contained about 2J 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 vertebra free from membrane.
Fur and colour. — The fur is soft and loose, the hairs on middle
of back about 10 mm. in length, those 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 ufopatagium not fringed.
Colour of upper parts 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 blackish-
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 whitish, especially on
6 "-^— ^ chest ; muzzle and cheeks dusky ; ears and
V, W s Sirf« membrane blackish.
£5Ud<oa^ 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 over anterior portion of
Fig. 29. brain-case ; occipital region distinctly pro-
mote gw duced 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-
able transverse constriction following anterior margin of inter-
171
parietal. Ventral profile rising slightly but evidently in region
of floor of brain-case. Greatest depth of brain-case about three-
quarters 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 bullse.
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 secondary 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 3 sub-
terete, its area more than
half that of canine and
about equal to that of pm v
its blunt main cusp, the
highest in the incisor series,
situated at outer side, the
three smaller cusps (third
obsolete) on inner margin ;
i x and i 2 sub-equal, slightly
lower than i 3 , their crowns
compressed, longer than
high, trifid, 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
FIS. 80.
Myotis mystacinus. Teeth X 10.
CHIROPTERA
alike in form, crowded between canine and large premolar but
perfectly in the tooth-row, the first about the same size as the
upper 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 equal to that of largest molar cusps ;
anterior and posterior 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 slight 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
but 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 2 , 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
uninterruptedly 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 m l , the protocone as in the
other teeth but smaller, paracone larger than metacone, the first
outer commissure longer than in m 1 and m 2 , 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 V slightly larger than first in cross section, this
condition reversed in m 3 ; protoconid decidedly higher than
hypoconid in all three teeth ; inner cusps of about equal height
throughout (slightly more than half as high as protoconid) ; a
distinct cingulum cusp behind entoconid.
Measurements. — Adult female from Skane, Sweden : tibia,
1 5 ; foot, 7 ' 6 ; forearm, 32 ; thumb, 6 • 2 ; third finger, 49 ; fifth
finger, 38; ear from meatus, 12; 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 ; Strass, near
Burgheim, Bavaria, 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.).
Austria-Hungary : Haida, Arva, Bohemia, 1 ; Carpathian Mts., 1
(U.S.N.M.) ; Csall6k6z-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. Gallen, 1 (U.S.N.M.) ; no
exact locality, 1 (Geneva: type of lugubris Patio).
Bemarkts. — Myotis 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 toe. From the small species of Pipistrellus it is at once
distinguished by the narrow muzzle and slender tragus.
2 ?. Aberia, Merionethshire,
Wales.
9 st. Colwyn, Denbighshire.
1 al. Newby Bridge, Cumber-
land, England.
9. Cheadle, Staffordshire.
6. Manchester, Lancashire.
<J. Ragley House, Warwick-
shire.
2 9. Welford, Herefordshire.
6. P e w s e v, Wiltshire.
(P. S.'Hembly.)
9. Dover, Kent. (B. Hesse.)
1. Hastings, Sussex.
6 st. Bath, Somerset.
1. Skullno, Sweden.
9. Haida, Bohemia. (Wol-
terstorff.)
1. Csall6kbz - Somorja,
Pressburg, Hungary.
1 al. Tatra Mountains,
Hungary.
G. H. Caton Haigh
(c & p).
Hon. N. C. Roths-
child (p).
T. Paul (c & p).
E. Blagg (c & p).
C. Oldham (p).
Tomes Collection.
Tomes Collection.
C. H. B. Grant (p).
C. H. B. Grant (p).
Miss I. Roods (c & p).
G. Dalgleish (c & p).
Stockholm Museum
(»■)
Lord Lilford (p).
Budapest Museum
(e)
Dr. R. Collett (p).
11. 1. 3. 13-14.
6. 2. 4. 1.
94. 9. 3. 1.
11. 1. 3. 15.
11. 1. 3. 10.
7. 1. 1. 498.
7.1.1.496-497.
11. 1. 3. 11.
11. 1. 3. 12.
49. 1. 16. 1.
4. 10. 13. 3.
46. 1. 2. 22.
11. 1. 1. 6.
94. 3. 1. 20-21.
91. 1. 21. 1.
174 CHIROPTERA
MYOTIS NATTERERI Kuhl.
1818. Vespertilio nattereri Kuhl, Ann. Wetterau. Gesellsch. Naturk., it
(= Neue Ann., i), pt. 1, p. 33.
1857. Vespertilio nattereri Blasius, Saugethiere Deutschlands, p. 88.
1863. [Isotus nattereri] var. typus Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 430 (Wiesbaden, Nassau ,
Germany).
1863. [Isotus nattereri] var. spelseus Koch, Jahrb. des Vereins fur Natur-
kunde im Herzogthum Nassau, xviii, p. 430 (Erdbach, Nassau,
Germany).
1878. Vespertilio nattereri Dobson, Cat'al. Chiropt. Brit. Mus., p. 88.
1900. Myotis nattereri MShely, Monogr. Chiropt. Hungariae, p. 179.
1904. Myotis escalerai Cabrera, Mem. Soo. Espafi. Hist. Nat., n, p. 279
(Bellver, Lerida, Spain).
1910. Myotis nattereri and M. escalerai Trouessart, Paune Mamrc. d'Europe,
pp. 29-30.
Type locality. — Hanau, Hessen, Germany.
Geographical distribution. — Central and southern Europe,
west to Ireland, north to southern Sweden.
Diagnosis. — Size medium among the European species (forearm
about 38 mm., longest finger about 70 mm., condylobasal length
of skull, 14" 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 M. 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
greatest width contained about 3£ times in length of anterior
border, its terminal third almost linear, sometimes faintly
recurved, the basal lobe usually small and ill defined ; insertion
of wing membrane as in M. 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
better defined than in M. 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. mystacinus, but free border of interfemoral membrane
distinctly though not very densely fringed with hairs about
I mm. long. Colour of upper parts a lighter and less yellowish
brown than in M. mystacinus, 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. mystacinus ; ears and membranes dark
brown.
Skull. — In all its dimensions the skull is appreciably larger
than that of Myotis mystacinus. In general form it is slightly
less slender. Forehead rising more 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 3 , with narrower
crowns.
Measurements. — 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,
I I • 0, 1 1 • 2. Two adult females from Spain (No. 94. 1 . 1. 8, Seville,
and No. 8. 7. 23. 4 from Bellver, Lerida, paratype of M. escalerai) :
head and body, 46, 46 ; tail, 41, 43 ; tibia, 15, 17 ; foot, 7 ■ 6, 9 • 2 ;
176
CHIROPTERA
forearm, 36-4, 40-6; thumb, 7 -2, 8-4; third finger, 69, 72;
fifth finger, 52, ,16; 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, 6 ; 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. Gallon, 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, Lerida, 2 (B.M. and Genoa; paratypes of escalerai
Cabrera) ; Seville, 1.
Remarks. — 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 the 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.
2 9, 9 juv. Harlech, Merioneth-
shire, Wales.
6. Arrow Church, Warwick-
shire, England.
8 al. Alcester, Warwickshire.
5 9 al. Lilford Hall, Northamp-
tonshire.
9. Bradfield, Berkshire.
9 st. Henley, Oxfordshire.
<5 st. Henley, Oxfordshire.
6 al. Queen Camel, Somerset.
1 al. England.
9. Oberlausitz, Silesia,
340 m. Germany.
(W. Baer.)
i al. Colpin, Brandenburg,
Prussia.
<$ al. Magdeburg, Saxony.
i. Magdeburg, Saxony.
(Wolterstcrff.)
9. Haida, Bohemia. (Wol-
terstorff.)
3<5,9. St. Gallen,600m. Swit-
zerland. (E. H. Zolli-
kofer.)
Duke of Argyll (p). 58. 8. 16. 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.
Tomes Collection.
Tomes Collection.
Lord Lilford (o & p).
N. H. Joy (c & p).
J. G. Millais (c & p).
Heatley Noble (c & p) .
R. H. Read (p).
Dr. J. E. Gray (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.
1. 11. 2. 1.
0. 3. 23. 1.
11.
51.
99.
3. 21.
29. 12.
9.6.
Dr. H. Gadow(c&p). 82.7.31.2.
Dr. W. Wolterstorff 92. 12. 1. 1.
Lord Lilford (p).
Lord Lilford (p).
O. Thomas (p).
11. 1. 1. 8-9.
11. 1. 1. 7.
4. 4. 5. 4-8.
MYOTIS 177
2 al. Arezzo, Italy. Florence Museum (e). 85.7.6.4-5.
9 al. Bellver, Lerida, Spain. A. Cabrera (p). 8. 7. 23. 4.
(Paratype of M. escalerai, Cabr.)
1 al. Seville. Dr. V. L. Seoane 94. 1. 1. 8.
(o & p).
MYOTIS EMARGINATUS Geoffroy.
1806. Vesp[ertilid] cmarginatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris,
viii, p. 198 (Charlemont, Givet, Ardennes, Prance).
1844. Vesp[ertilio] rufescens Crespon, Faithe Meridionale, i, p. 20 (near
Nimes, Gard, Prance). Type in Nimes Museum.
1853. V[espertilio] ciliatus Blasius, Wiegmann's Archiv fur Naturgesch.,
1853, i, p. 287 (near Cologne, Germany).
1856. V[espertilio] schrankii Kolenati, Allgem. deutsche Naturhist. Zeitung,
Dresden, neue Polge, n, p. 178. A nomen nudum (ex Koch and
Giebel) cited as synonym of cmarginatus. Not of Wagner, 1843.
1858. Vespertilio ciliatus Blasius, Saugethiere Deutschlands, p. 91.
1878. Vespertilio emarginatus Dobson, Catal. Chiropt. Brit. Mus., p. 303.
1880. Myotis ciliata var. budapestiensis Marg6, " Magyar orv. es termes-
zetvizsg. xx,nagygydl. munk, p. 255" (Budapest, Austria-Hungary).
See Mehely, Monogr. Chiropt. Hungariae, 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. Myotis emarginatus Mehely, Monogr. Chiropt. Hungariae, p. 170.
1910. Myotis emarginatus Trouessart, Paune Mamm. d'Europe, p. 28.
Type locality. — Charlemont, Givet, Ardennes, France.
Geographical distribution. — Central and southern Continental
Europe.
Diagnosis. — Size essentially as in Myotis nattereri (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 interfemora'l 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. — Whole animal larger and more robust
than Myotis mystacinus, 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 surface of conch marked by 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
178
CHIEOPTERA
to rather acute tip. Wing as in the related species ; metacarpals
falling 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 M.
mystacinus, extending to between ears when laid forward, only
the minute cartilaginous tip free from membrane.
Fur and colour. — Fur shorter and more dense than in M.
mystacinus and M. nattereri, and of a slightly woolly texture
unique among the European species, the hairs at middle of back
about 8 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.
Skull. — In form the skull does not differ appreciably from
that of M. nattereri, except that the rostral portion and palate are
relatively more elongate and interorbital concavity is less strongly
pronounced ; occipital region similarly low as compared with that
of 31. mystacinus. Mandible slightly more robust than in M.
nattereri, but of similar form.
Teeth. — The teeth are larger than those of Myotis mystacinus,
in this respect agreeing with those of M. nattereri. 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 evenly convex
(U-shaped) row without anterior angle, the crowns of i 1 and «,
alike in form and distinctly 4-cusped, that of i x not thickened
posteriorly and with no trace of postero-internal cusp. Canines
about equal to those of M. 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 M. mystacinus, the cusp of second only a httle
exceeding cingulum of first ; the two teeth less crowded between
canine and large premolar than in M. mystacinus and M. nattereri ;
lower premolars essentially as in 31. 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.
3Ieasurements. — Two adult females from Florence, Italy
head and body, 46' 6 and 50 ; tail, 40 and 42 ; tibia, 19 and 19
foot, 8 - 4 and 86; 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
France: Near Ntmes, Gard, 1 (Nimes : type of rufescens Crespon).
Austria-Hungary : Herkulesbad, 2.
Switzerland : Valavran, near Geneva, 1 (Geneva : type of neglectus
Patio).
Italy : Florence, 2 (U.S.N.M.) ; no exact locality, 1.
Bemarhs. — The peculiar form of the ear, the short, somewhat
woolly fur, and the yellowish colour are highly characteristic of
this well-defined species.
2 9. Herkulesbad, Hungary. Hon. N. C. Roths- 7. 9. 16. 9-10
child (p).
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 733.
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, Saugethiere Deutschlands, p. 85.
1878. Vespertilio bechsteinii Dobson, Oatal. Chiropt. Brit. Mus., p. 308.
1900. Myotis bechsteinii Mehely, Monogr. Chiropt. Hungarise, p. 184.
1905. Vesp\ertilw\ bechst[emii] ghidinii vel Vesplertilio] ghidinii Patio,
Arch. Sci. Phys. et Nat. ; Geneve, 4th ser., xix, p. 511, May 15,
1905 (Lugano, Ticino, Switzerland). Type in Geneva Museum.
1906. Myotis bechsteinii 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. bechsteini favonicus Trouessart, Faune
Mamm. d'Europe, pp. 30-31.
Type locality. — Hanau, Hessen-Nassau, Germany.
Geographical distribution. — Central and southern Europe,
west to England, north to southern Sweden.
Diagnosis. — Size slightly greater than that of M. nattereri
(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 30 mm. instead of
less than 25). Muzzle with moderately developed glandular
swellings (these less evident than in M. mystacinus). Ear extend-
ing considerably 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 mm.) that the ear is relatively larger
than in any other European bat except Plecotus auritus ; 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-defined transverse ridges near posterior border ; anti-
tragus about 2-5 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 fifth 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 membrane as in M. mystacinus. Tail shorter
than head and body (laid forward it extends to middle of crown),
the terminal vertebra free.
Fur and colour. — Quality and distribution of fur as in Myotis
mystacinus and M. nattereri. Upper parts uniform wood-brown
with a slight tinge of umber, the exact shade almost impossible
to describe exactly ; underparts a buffy 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 Myotis
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 bi - ain-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 portion 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 developed at sides, obsolete
at middle ; greatest depth 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 oblique posterior border than in the other small species.
Teeth. — The teeth are rather small relatively to size of skulh
MYOTIS 181
Upper incisors essentially as in M. mystacinus 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 equal 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-quarters that of
first molar, its antero-internal cusp well developed. Lower
premolars essentially as in M. nattereri. Upper molars with no
special peculiarities except that m 3 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 3 a little more reduced
than in the other small species, though much less so than in
M. myotis and M. oxygnathus.
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 favonicus 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.
Specimens examined. — Seventeen, from the following localities : —
England : Henley-on-Thames, Oxfordshire, 1 ; New Forest, Hamp-
shire, 6.
Sweden: Skane, 1 (U.S.N.M.).
182
CHIROPTERA
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184
CHIROPTERA
Fbance : Ktupes, Doubs, 1 (Mottaz) ; Ste. Baume, Var, 1 (Genoa) ;
Cadillac, Gironde, 1 (Lataste).
Switzeeland : Lugano, Ticino, 1 (Geneva: type of ghidinii Fatio).
Spain: La Granja, Segovia, 1 (type of favonicus Thomas).
Atjstbia-Hongaey : Zay-Ugr6cz, Trenesen, 4 (B.M. and U.S.N.M.).
Remarks. — 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.
<5. 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.
i al. La Granja, Segovia, M. de la Escalera (c). 6.11.4.1.
Spain. (Type of M. favonicus Thos.)
c5, ? al. Zay-Ugrocz, Trenesen, Budapest Museum 0. 6. 20. 1-2.
Hungary. (e).
1. Europe. Leyden Museum. 37. 4. 28. 23.
MYOTIS DAUBENTONII Kuhl.
1619. Vespertilio daubentonii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 195 (Hanau, Hessen, Germany).
1839. Vespertilio sedilis 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. Vespertilio daubentonii Blasius, Saugethiere Deutschlands, p. 98.
1871. Vespertilio capucinellus • ' Koch, Bayr. Fauna," Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 206 (Bavaria ?).
1871. Vespertilio minutellus "Koch, Bayr. Fauna," Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,
p. 206 (Bavaria?).
1871. Vespertilio daubentonii, albus Fitzinger, Sitzungsber. kais. Akad.
Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, p. 210
(Renaming of V. sedilis Jenyns).
1878. Vespertilio daubentonii Dobson, Catal. Chiropt. Brit. Mus., p. 297.
1890. Vespertilio staufferi Fatio, Faune Vert. Suisse, v, 3 mo suppi. aux
Mamm., p. 6 (Lucerne, Switzerland). MS. synonym; repudiated.
1900. Myotis daubentonii Mehely, Monogr. Chiropt. Hungarise, p. 164.
1910. 2Ii/otis daubentoni Trouessart, Faune Mamm. d'Europe, p. 27.
Type locality. — Hanau, Hessen-Nassau, Germany.
Geographical distribution. — Europe from the Mediterranean
north to central Norway, west to Ireland, east into Asia.
Diagnosis. — Size nearly as small as in Myotis mystacinua
(forearm about 35 mm., longest finger about 60 mm., condylo-
basal length of skull, 13-4 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.
mystacinus, 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 denned. Wing slightly broader than in M. mysta-
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 M. mystacinus, but with no peculiarities of distribution
except that it tends to spread farther outward along dorsal
surface of interfemoral membrane ; free border of uropatagium
not fringed, but a slight fringe is usually present along basal
half of calcar. Colour above essentially as in M. na/lereri,
though usually inclining more definitely toward wood-brown ;
underparts buffy grey usually less contrasted than in M. nattered,
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 daubentonii is smaller than that
of any other European species except M. mystacinus. 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 Myotis 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, the form of the row vacillating between (J -shaped
and broadly V-shaped, the cusps as in M. mystacinux but less
186
CHIKOPTERA
FIG. 31.
Mi/otis daubentonii. Teeth x 10.
developed. Canines both above and below weaker and less
trenchant than in M. myslacinus, their form essentially as in
M. nattereri. Premolars
and lower molars not
obviously different from
those of M. mystacinus.
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 1 and
m 2 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-Avon, Warwickshire, 2 ; Hillingdon, Middlesex 1, ; Epping, Essex, 2 ;
Northamptonshire, 1 ; Henley-on-Thames, Oxfordshire, 2 (B.M. and
U.S.N.M.) ; Christchnrch, Hampshire, 1 ; Devonshire, 1.
Sweden: Upsala, 3 (B.M. and U.S.N.M.); upland, 1; no exact locality, 2.
Switzerland : Geneva, 1 (Mottaz).
Pkance : 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).
Remarks. — 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).
6. Bowdon, Cheshire, Eng- T. A. Coward (c & p). 11. 1. 3. 60.
land.
MYOTIS 187
?. Knutsford, Cheshire. T. A. Coward (c & p). 11. 1. 3. 22.
2. Stratford-on- Avon, War- Tomes Collection. 7.1.1.486-487.
wickshire.
6 al. Hillingdon, Middlesex. 0. 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 (o & p). 1. 11. 2. 2.
1 al. Christohurch, Hamp- Lord Lilford (p). 87. 9. 1. 1.
shire.
S. Upland, Sweden. Lord Lilford (p). 11. 1. 1. 25.
(G. Kolthoff.)
2. Sweden. Stockholm Museum 46. 6. 2. 15.
(e). 48. 6. 28. 3.
MYOTIS CAPACCINII Bonaparte.
1837. VespertiUo capaccinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx
(Sicily). Type in British Museum.
1839. VespertiUo megapodius Temminck, Monogr. de Mamm., n, p. 189
(Sardinia).
1841. Vesp[ertilio] dasypus de Selys-Longchamps, Atti della seconda
Riunione degli Scienziati Italiani, Torino, 1840, p. 249 (Published
as synonym of capaccinii).
1844. Vesp[ertilio] pellucens Crespon, Faune Meridionale, i, p. 16 (Cave
near Pont-du-Gard, Gard, Erance).
1857. VespertiUo capaccinii Blasius, Saugethiere Deutschlands, p. 101.
1877. VespertiUo blasii Major, Atti Soc. Tosc. Sci. Nat., Pisa, in, p. 108
(New name for the capaccinii of Blasius should it prove to be
different from that of Bonaparte) .
1878. VespertiUo majori Ninni, Atti Beale Instit. Veneto, 5th ser., iv,
pt. 1, p. 721 (Substitute for blasii Major).
1878. VespertiUo capaccinii Dobson, Catal. Chiropt. Brit. Mus., p. 293.
1901. Myotis capaccinii Thomas, Proc. Zool. Soc. London, p. 37.
1910. Myotis (Leuconoe) capaccinii Trouessart, Paune Mamm. d'Europe, p. 26.
Type locality. — Sicily.
Geographical distribution. — Mediterranean region, north to
Italian Switzerland, east into Asia.
Diagnosis. — Not so small as Myotis daubentonii (forearm about
42 mm., longest finger, 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 characters. — In all essential features the external
form is as in M. daubentonii. 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 distinct 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
CHIEOPTERA
above a light drab tinged with grey or with ecru-drab, the
general effect paler and more greyish than in any of the other
European species ; underparts pale buffy 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
light brown.
Skull. — In all respects the skull so closely resembles that of
Myotis daubentonii that it is only distinguishable by its larger
size. Erom 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 post-palatal region.
Teeth. — Except for their larger size the teeth resemble those
of Myotis daubentonii. The crown area of upper molars is
relatively less than in M. nattereri and M. emarginatus, though
m 3 retains the broader outline characteristic of M. daubentonii
and M. mystacinus. Upper molars with protoconule and its
accessory small commissure, and m 1 and m 2 with commissure
between hypocone and metacone as in M. daubentonii.
Measurements. — 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 ; third
finger, 56 ■ 2 ; fifth finger, 50. Adult male from Oorleone, 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,
1 5 ; 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 examined. — 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); iPavia, 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.
Hemarlts. — In general appearance this species resembles
Myotis 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, ?. Near Lugano, Ticino, O. Thomas (p). 2. 8. 4. 10-14.
Switzerland.
(E. H. Zollikofer.)
2. Ostia, Rome. Dr. L. Sambon(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. Marsala, Sicily. 0. Thomas (p). 6. 8. 4. 21.
(A. Robert.)
2 al. Cagliari, Sardinia. FlorenceMuseum(E). 85.7.6.6-7.
2 al. Grotte de Sardale, Sar- 0. Thomas (p). 0. 12. 3. 1-2.
dinia.
c5. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 3.
Islands. Pocock (c & p).
6. Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 13.
MYOTIS DASYCNEME Boie.
1823. VespertiUo mystacinus Boie, Isis, p. 965. Not V. mystacinus Kuhl
(Jutland, Denmark).
1825. VespertiUo dasycneme Boie, Isis, p. 1200 (Renaming of mystacinus
Boie).
1839. VespertiUo limnophilus Temminck, Monogr. de Mamm., n, p. 176
(Holland).
1857. VespertiUo dasycneme Blasius, Saugethiere Deutschlands, p. 103.
1878. VespertiUo dasycneme Dobson, Catal. Chiropt. Brit. Mus., p. 295.
1904. Myotis dasycneme Trouessart, Catal. Mamm. Tam viv. quam foss.,
suppl., p. 88.
1910. Myotis dasycneme Trouessart, Paune 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 essentially as in Myotis
daubentonii and M. capaccinii, but size much larger (forearm
about 47 mm., longest finger 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 Myotis, 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- off tip, the two margins
essentially parallel through lower half. Free border of uro-
patagium without fringe.
Fur and colour. — Distribution of fur as in Myotis daubentonii ;
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 buff, the line of demarcation along sides of neck well
defined. Muzzle and cheeks scarcely contrasted dusky. Under-
fur slaty black. Ears and membranes an indefinite dark brown.
* The supposed British record is probably erroneous (see Barrett-
Hamilton, Hist. Brit. Mamm., I, pp. 157-158. December, 1910).
190
CHIROPTERA
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CHIEOPTERA
Skull. — The skull is large, slightly exceeding that of Myotis
bechsteinii 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 daubentonii, but the rostrum is relatively
shorter and the brain-case broader and more depressed. Sagittal
crest barely indicated ; auditory bullae proportionately as in the
smaller animal, less inflated than in M. bechsteinii ; 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
daubentonii ; molars decidedly more robust than those of M.
bechsteinii, 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. daubentonii and M. capaccinii ; mandibular teeth
with no special peculiarities, the premolars and outer cusps of
molars more slender than in M. bechsteinii and M. daubentonii.
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 linger, 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 examined. — Six, from the 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. Schlegel.)
6 al. Namur, Belgium. Bev. D. B. Lebbe 9. 1. 11. 1.
(c & p).
MYOTIS MYOTIS Borkhausen.
1775. Vespertilio murinus Schreber, Saugthiere, I, p. 165 (Not of Linnaeus,
1758).
1797. Vespertilio myotis Borkhausen, Deutsche Fauna, i, p. 80 (Germany).
1797. Vespertilio myosotis B[orkhause]n, Der Zoologe (Compendiose
Bibliothek gemeinnutzigsten Kenntnisse fur alle Stande, pt. xxi),
Heft v-vin, p. 46 (Germany).
1800. Vespertilio myosotis Bechstein, Pennant's Allgemeine Uebersicht der
Vierfiissigen Thiere, p. 632 (Germany).
MYOTIS 193
1801. Vespertilio myotis Bechstein, Gemeinn. Naturgesch. Deutschl., i,
2nd ed., p. 1154 (Described but not named in 1st ed., 1789, p. 164)
Thiiringen, Germany.
1827. Vespertilio submurinus Brehm, Ornis, Heft in, p. 24 (Renthendorf,
Thiiringen, Germany).
1844. V[espertilio] latipinnis Crespon, Paune Meridionale, I, p. 17 (Near
Nimes, Gard, France).
1857. Vespertilio murinus Blasius, Saugethiere Deutschlands, p. 82 (Not of
Linnaeus, 1758).
1863. [Myotus murinus'] var. typus Kocb, Jahrb. des Vereins fur Natur-
kunde im Herzogthum Nassau, xvin, p. 415 (Wiesbaden, Nassau,
Germany).
1863. [Myotus murinus] var. alpinus Koch, Jahrb. des Vereins fur Natur-
kunde im Herzogthum Nassau, xvm, p. 415 (St. Gothard, Uri,
Switzerland).
1878. Vespertilio murinus Dobson, Catal. Chiropt. Brit. Mus., p. 309 (Not
of Linnaeus, 1758).
1886. Myotis murma var. spelsea Bielz, Verhandl. u. Mittheilungen des
Siebenbiirgischen Vereins fur Naturwissensch. in Hermannstadt,
xxxvi, p. 83 (Homorod- Almas cave, Hungary).
1897. Myotis myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 383,
October, 1897.
1909. Myotis myosotis Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S., in,
No. 3, p. 1, April 26, 1909.
1910. Myotis myotis and M. m. spelsea Trouessart, Faune Mamm. d'Burope,
p. 32.
Type locality. — Thiiringen, 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 finger, 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 remarkably 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 margin moderately convex from base
nearly to rather narrowly rounded-off tip ; posterior border with
shallow ill-defined concavity above ; antitragus low and long,
marked off posteriorly by a well-defined notch and not continuous
with posterior border of conch ; tragus about half as high as
* Bell, Hist. British Quadrupeds, p. 38, 1836 : "But in England it . . .
has hitherto only been taken in the gardens of the British Museum."
O
194
CHIEOPTBEA
conch, its greatest width (slightly above level of anterior basey
contained about 2J 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 phalanges as compared with the smaller
species, the third, 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
interfemoral 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 Myotis
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 buffy 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 Myotis, M.
oxygnathus excepted, the skull of
Myotis 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
no. 32.
Myotis myotis. Nat. size.
195
about on level with main portion of brain-case, and very slightly
overhanging foramen magnum ; ventral profile scarcely elevated
posteriorly ; palate rather narrow (essentially as in M. mystacinus) ;
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 relatively 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 Myotis. 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
ij and i 2 trifid, but decidedly
oblique owing to reduction
in size of outer cusp ; a
minute cingulum cusp usu-
ally present at extreme outer
edge ; i 2 with small postero-
FIG. 33.
Myotis myotis. Teeth X 10.
internal tubercle ; i 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. natter eri
and JIT. emarginatus. Small
upper premolars much
crowded, the second usually
forced inward from line of tooth-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 secondary 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 m 1 and m 2 barely
indicated ; m 3 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
o 2
196
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CHIROPTEEA
in the small species, except that cingulum does not form a postero-
internal cusp behind entoconid ; m 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 different 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, 18-1 (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.
Specimens examined. — Ninety-five, from the following localities : —
France : Cadillac, Gironde, 1 (U.S.N.M.) ; Nimes, Gard, 2 (B.M. and
Nimes ; representing latipinnis Crespon, but not type).
Germany : Hamburg, 1 ; Niesky, Silesia, 11 ; Strass, near Burgheim,
Bavaria, 8 ; Heidelberg, Baden, 1.
Aostria-Hungary : Herkulesbad, 1 ; Fiinfkirchen, S.W. Hungary, 2.
Roumania: Bustenari, 3; Sinaia, 1 (U.S.N.M.).
Switzerland : Geneva, 7 (Mottaz) ; Grotte de Vallorbe, Vaud, 1
(Mottaz) ; Boudry, Neuchatel, 2 (Mottaz) ; St. Moritz, 1 ; Thayngen, Schaff-
hausen, 2 (U.S.N.M.); Canton Thurgau, 5; Tagerwilen, Thurgau, 10
(U.S.N.M.); Andermatt, Uri, 1 (U.S.N.M.); St. Gothard, Uri, 3 (B.M.
and U.S.N.M.) ; Monte Generoso, Ticino, 4 (U.S.N.M.).
Italy : Domodossola, 2 (U.S.N.M.) ; Finalborgo, Liguria, 1 (Genoa) ;
Florence, 1 (U.S.N.M.); Rome, 2; Ostia, Borne, 2; Marsala, Sicily, 6.
Sardinia : Oristano, Cagliari, 10.
Spa™ : Seville, 2.
Portu&al : Cintra, 2.
Remarks. — Myotis myotis differs strikingly from the other
European members of the genus, M. oxygnathus excepted, in its
much larger size. From the large Vespertilionid.ee of other genera
it is immediately recognizable by its long ears, extending notice-
ably beyond nostril when laid forward, and by the greyish white
colour of the underparts. In the Mediterranean region Myotis
myotis is associated with M. oxygnathus ; but it is the only large
species known to occur north of the Alps.
lal.
Nimes, Gard, Prance.
G. E. Dobson (e).
80. 12. 14. 2.
1.
Hamburg, Germany.
Dr. J S E. Gray (p).
2 6,8 9,
Niesky, Silesia. (Dr.
Dr. E. Hamilton (p).
97. 12. 4. 7-17.
? juv.
W. Baer.)
6 9.
Strass, Burgheim, Ba-
varia. (Korbitz.)
Lord Lilford (p).
11. 1. 1. 10-15.
?.
Strass, Burgheim, Ba-
Lord Lilford (p).
11. 1. 1. 124.
varia. (Korbitz.)
11. 1. 1. 131.
MYOTIS 199
9. Heidelberg, Baden. Hon. N. C. Botha- 10. 5. 29. 1.
child (p).
■i juv. Herkulesbad, Hungary. Hon. W. Rothschild 7. 9. 16. 8.
(F. J. Cox.) (p).
2 al. Piintkirchen. Budapest Museum (e). 94.7. 18. 11-12.
2 (J, 9. Bustenari, Prahova, Lord Lilford (p). 4. 4. 6. 8-10.
840 m. Boumania.
(W. Dodson.)
1. St. Moritz,Grisons,Swit- Leon 0. Galliard (p). 75.9.20.4.
zerland.
5 9. Thurgau. (E. H. Zolli- 0. Thomas (p). 4. 4. 5. 9-13.
kofer.)
2 9. Borne. (C. Goli.) G. Barrett-Hamilton 11. 1. 2. 26-27.
2. Ostia, Borne. Dr.L. Sanation (c&p). 1.1.2.1-2.
6 9. Marsala, Sicily. (A. O. Thomas (p). 6. 8. 4. 17-20.
Robert.) 8. 9. 1. 3-4.
10 al. Oristano, Cagliari, Sar- Hon. N. C. Boths- 7. 5. 24. 1-10.
dinia. child (p).
2 6 al. Seville, Spain. (Dr. A. Lord Lilford (p). 73. 1. 8. 1-2.
Ruiz.)
2 1. Cintra, Portugal. 0. Thomas (o & p). 98. 2. 2. 4-5.
MYOTIS OXYGNATHUS Monfcicelli.
1885. Vespertilio oxygnathus Monticelli, Ann. Accad. 0. Costa de Aspir.
Nat., I, p. 82. Type in Naples Museum.
1909. Myotis oxygnathus Miller, Ann. Mus. Zool. B. Univ. Napoli, N.S.,
in, No. 3, p. 1, April 26, 1909.
1910. Myotis myotis oxygnathus Trouessart, Faune Mamm. d'Burope, p. 32.
Type locality. — Matera, Basilicata, Italy.
Geographical distribution. — Mediterranean region from Spain
to Greece, north to Italian Switzerland ; 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 body, 63 ; tail,
54; tibia, 24-4; foot, 13; forearm, 57; thumb, 11-4; third
finger, 98 ; fifth finger, 76 ; ear from meatus, 23 ; width of ear,
13-6; tragus, 1 ■ 8. Two adult males from Velletri, Rome,
Italy : head and body, 60 and 62 ; tail, 58 and 58 ; tibia, 23
and 24 • 6 ; foot, 1 2 • 8 and 1 3 ; forearm, 53 • 6 and 57 ; thumb, 1 2
and 11 ; third finger, 89 and 86 ; fifth finger, 73 and 76 ; ear
from meatus, 23 and 24 ; width of ear, 14 and 15. Two adult
females from the same locality : head and body, 62 and 66 ;
tail, 58 and 57 ; tibia, 24 and 24- 6 ; foot, 13 and 12 ; forearm,
56 and 58 ; thumb, 11 and 11 -8 ; third finger, 93 and 97 ; fifth
finger, 74 and 78 ; ear from meatus, 23 and 22 ; width of ear,
13 ■ 6 and 13-6. Adult male and female from Bozen, Tirol : head
and body, 68 and 71 ; tail, 53 and 53 ; tibia, 25-4 and 24 ; foot,
200
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202
CHIROPTEEA
14 and 14 ; forearm, 59 and 60 ; thumb, 11 • 4 and 11 • 6 ; third
finger, 97 and 98 ; fifth finger, 76 and 78 ; ear from meatus, 26
and 25 ; width of ear, 17 and 15. For cranial measurements see
Table, p. 200.
Specimens examined. — Seventy-four, from the following localities : —
Spain : Near Burgos, 2.
Switzebland : Lugano, Tioino, 1 (U.S.N.M.).
Austbia-Hungaby : Bozen, Tirol, 3 (U.S.N.M.).
Italy : Finalborgo, Liguria, 2 (Genoa) ; Isoverde, 1 (Genoa) : Vallom-
brosa, Florence, 1 (U.S.N.M.); Borne, 4; Velletri, Borne, 15 (U.S.N.M.);
Matera, Basilicata, 1 (Naples, type).
Sabdinia: Gagliari, 5 (U.S.N.M.); Monte Gennargentu, 3 (U.S.N.M.) ;
no exact locality, 3 (U.S.N.M.).
Malta : El Ghain, 5 ; Bubato, 3 ; no exact locality, 4.
Montenegbo : Beri, 4.
Gbeece: Patras, 2; Corinth, 7 (U.S.N.M.); Nauplia, 1 (U.S.N.M.);
Lamia, Thessaly, 1 (U.S.N.M.)
Cbete : Labyrinth, 4.
Tunis : No exact locality, 3 (U.S.N.M.).
Remarks. — In general appearance Myotis oxygnaihus resembles
M. myotis, though the colour perhaps averages somewhat darker.
It is readily distinguishable, however, by its smaller skull
(distinctly smaller head in spirit specimens), and shorter, narrower
ears. In form the skull is like that of M. myotis ; and the
teeth are not peculiar except for their small size, a character
readily appreciable on comparison of the canines or of the crown
area of upper molars. The range of Myotis oxygnaihus is, so far
as known, strictly Mediterranean, probably coincident with that
of M. capaccinii and Pipistrellus kuhlii. North of the Alps
ilf. myotis occurs alone.
9. Burgos, Spain. G. S. Miller (c). 8. 8. 4. 10.
4 <$. Borne, Italy. (Goli.) G. Barrett-Hamilton 11. 1. 2. 22-25.
(*)■
3 i, 2 9. El Ghain, Malta. Lord Lilford (p.) 11. 1. 1. 16-20.
(Micallef.)
3 6, 9. Bubato, Malta. Lord Lilford (p). 11. 1. 1. 21-24.
(Micallef.)
4. Malta. (Micallef.) Lord Lilford (p). 95. 3. 2. 3-6.
2 (J, 2 9. Beri, 50 m. Montenegro. O. Thomas (p). 5.8.4.1-4.
(L. Filhrer.)
2 9. Patras, Greece. Hon. N. C. Boths- 8. 10. 2. 15-16.
(C. Mottaz.) child (p).
3 S, 9. Labyrinth, Crete. Miss T>. Bate (c). 5. 12. 2. 5-8.
Genus PIPISTRELLUS Kaup.
1829. Pipistrellus Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt,
I, p. 98 (pipistrellus).
1837. Pipistrellus Bonaparte, Iconogr. Faun. Ital., I, fasc. sx.
1838. Bomicia Gray, Mag. Zool. and Bot., ii, p. 495, February, 1838
(calcarata = kuhlii).
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur-
gesch., 1839, i, p. 312 (part).
1856. Hypsugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, II, p. 131 (maurus and krascheninikowii).
PIPISTiJELLUS
203
1856. Nannugo Kolenati, Allgom. deutsohe Naturhist. Zeitung, Dresden,
neue Folge, n, p. 131 (nathusU, pipistrellus, and kuMU).
1857. Vesperugo Blasius, Saugethiere TJeutschlands, p. 49 (part).
1878. Vesperugo Dobson, Oatal. Chiropt. Brit. Mus., p. 183 (part).
1897. Pipistrellus Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,
October, 1897.
1899. Euvesperugo Acloque, Faune de France, Mammiferes, p. 35 (part,
included noctula, leisleri, maurus, kuMii, pipistrellus, and
abramus).
1907. PipistrelVus Miller, Families and Genera of Bats, p. 204, June 29,
1907.
Type species. — Vespertilio pipistrellus Schreber.
Geographical distribution. — Entire mainland of Eastern Hemi-
sphere to limits of tree growth, also Malay Archipelago, New-
Guinea, Solomon Islands, and northern Australia ; in America
from northern United States (except in boreal zone) to southern
Mexico.
Characters. — Like Eptesicus (p. 224), but with 2 — 2 upper
premolars ; dental formula : i ||, c t|, pm ^|, « jt| = 34.
Bemarhs. — The genus Pipistrellus is widely distributed in the
warmer portions of the Old World and of North America.
About forty species are now known, four of which occur in
Europe. Externally these may often be confused with the smaller
Myotis, though they may usually be recognized by a certain
heaviness of form, and more especially by the shorter ear and
less slender tragus. Though often regarded as nearly related to
Nyctalus, on account of the similarity of dental formula, this
genus is much the more primitive of the two, its members
showing no tendency to modify the ordinary vespertilionine wing
structure. It is in reality not much more than a sub-genus of
Eptesicus (see remarks under the latter), though for the sake of
convenience the two groups are best treated as distinct.
KEY TO THE EUROPEAN SPECIES OF PIPISTRELLUS.
Anterior upper premolar excessively minute, some-
times hidden by the gum, its crown area much
less than that of outer incisor ; anterior lower
premolar with crown area less than half that
of posterior premolar ; greatest width of tragus
nearly equal to length of anterior border ; hairs
of back usually with noticeably contrasted light
tips P. savii, p. 219.
Anterior upper premolar not excessively minute,
never hidden by the gum, its crown area about
equal to that of outer incisor ; anterior lower
premolar with crown area more than half that
of posterior premolar ; greatest width of tragus
much less than length of anterior border ; hairs
of back without noticeably contrasted light tips.
Outer upper incisor less than half as high as
inner ; large upper premolar almost or quite
in contact with canine, the small premolar
forced inward from tooth-row and scarcely or
not visible from outer side P. kuhlii, p. 215.
204
CHIROPTERA
Outer upper incisor more than half as high as
inner; large upper premolar separated from
canine by distinct space in which the small
premolar is clearly visible from outer side.
Lower canine robust, the length of base along
cingulum about equal to length of anterior
border of shaft; oondylobasal length of
skull 11 to 12 mm. ; thumb short, its length
about equal to width of wrist ; fifth finger
about 40 mm P. pipistrellus, t>. 204.
Lower canine slender, the length of base along
cingulum slightly more than half length
of anterior border of shaft ; oondylobasal
length of skull 12-6 to 13-4 mm. ; thumb
long, its length much greater than width
of wrist; fifth finger about 46 mm P. nathusii, p. 213.
PIPISTRELLUS pipistrellus Schreber.
1774. Vespertilio pipistrellus Schreber, Saugthiere, I, pi. liv. Described, i,
p. 167, 1775, under name Die Zwergfledermaus. (Prance, based
primarily on Daubenton.)
1776. VespertiUo pipistrelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 16 (Prance, based on Schreber).
1825. Vespertilio pygmseus Leach, Zool. Journ., i, p. 560, January, 1825
(Dartmoor, Devonshire, England).
1834. VespertiUo brachyotos Baillon, Mem. Soc. Royale d'Emulation
d' Abbeville, 1833, p. 50 (Abbeville, Somme, Prance).
1839. ? [VespertiUo pipistrellus] var. nigra de Selys-Longchamps, Etudes
de Micromamm., p. 140 (nomen nudum).
1839. ? [Vespertilio pipistrellus'] var. rufescens de Selys-Longchamps, Etudes
de Micromamm., p. 140 (nomen nudum).
1840. V[espertilio] pusillus Schinz, Europ. Pauna, I, p. 9 (Synonym of
pipistrellus ; Brehm cited as authority).
1840. V[espertilio] melanopterus Schinz, Europ. Pauna, I, p. 9. Brehm
cited as authority, but name apparently published here for first
time (Rhentendorf, Thiiringen, Germany).
1840. V[espertilio] stenotus Schinz, Europ. Pauna, ±, p. 9. Brehm cited as
authority, but name apparently published here for first time
(Rhentendorf, Thiiringen, Germany).
1840. Vespertilio minutissimus Schinz, Europ. Pauna, i, p. 9 (Zurich,
Switzerland).
1845. P[ipistrellus] nigricans Bonaparte, Atti della 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 genei Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, Milano, 1844, p. 340 (Alternative name for nigricans).
1845. P[ipistrellus] typus Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, Milano, 1844, p. 340 (Substitute for pipistrellus
Schreber).
1857. Vesperugo pipistrellus Blasius, Saugethiere Deutschlands, p. 61.
1862. Vesperugo pipistrellus var. macrcpterus Jeitteles, Verhandl. der k. k.
Zool. Bot. Gesellsch., Wien, hi, p. 250 (Kaschau, Hungary).
1863. [Nannugo pipistrellus] var. typus Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xvin, p. 490. Not of
Bonaparte, 1845 (Wiesbaden, Hessen-Nassau, Germany).
PIPISTRELLUS 205
1863. [Nannugo pipistrellus] var. flavescens Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xviii, p. 491 (Nassau,
Germany).
1863. [Nannugo pipistrellus] var. nigricans Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xviii, p. 491. Not of
Bonaparte, 1845 (Nassau, Germany).
1863. [Nannugo pipistrellus'] var. Umbatus Koch, Jahrb. des Vereins fur
Naturkunde im Herzogthum Nassau, xviii, p. 491 (Siegen, Nassau,
Germany).
1878. Vesperugo pipistrellus Dobson, Catal. Ghiropt. Brit. Mus., p. 223.
1897. Pipistrellus pipistrellus Miller, Ann. and Mag. Nat. Hist., 6th ser.,
xx, p. 384, October, 1897.
1904. Pipistrellus pipistrellus mediterraneus Cabrera, Mem. Soc. Espafi.
Hist. Nat., II, p. 273 (Valencia, Spain).
1910. Pipistrellus pipistrellus and P. pipistrellus mediterraneus Trouessart,
Paune 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 membrane 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 nearly straight or very
slightly convex ; posterior basal lobe small but well defined.
Wing rather narrow, with no special peculiarities of form, the third,
fourth and fifth metacarpals sub-equal (fifth slightly shorter than
206
CHIEOPTBRA
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 tapering rapidly and terminating without lobe, its keel well
developed, with evenly convex margin. Tail about as long as
body without head and 2^ times as long as tibia, the snort
terminal vertebra free from membrane.
Fur and colour. — The fur is closely confined to the body,
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-brown and
cinnamon of Ridway but sometimes darker, with a strong tinge
of prouts-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 behind
middle, its surface smooth or with faintly indi-
cated sagittal crest and lateral portion of
lambdoid crest ; greatest breadth of brain-case
"fig 34 noticeably exceeding that of rostrum and slightly
Pipistreiius pipi*- though evidently more than half greatest length
trains. Nat. size, of skull ; floor of brain-case 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
broadly 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
PIPISTEELLUS 207
inflated, the inflated region with a median angle suggesting
a rudimentary postorbital process ; rostrum short and broad,
narrowing gradually in front, a slight concavity at each side
bordering lachrymal inflation, and an evident median longi-
tudinal groove, most noticeable posteriorly ; nasal emargina-
tion 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
emargination 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 symphysis 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 irregularly 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 doveloped. 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 U -shaped
row between canines, their crowns very slightly imbricated ;
crowns much longer than high, trifid, that of i x narrowest,
longest and lowest, that of i 2 and i 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
208 CHIEOPTBRA
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,
forming 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-fifth 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
/?V^ ^ narrow and flattened-concave, with evident
-^?T\l elevated rim, the posterior border strongly
Cx 3 . „ f\ concave, the anterior border usually convex
30 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
*K£Mh x ""*■ tooth along outer cingulum, posterior cutting
edge well 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 ; metacone higher than paracone ; styles
well developed ; VV-p a ttern normal ; m 3 with crown area about
two-thirds that of m 1 , the hypocone absent, the metacone smaller
than paracone ; no trace of metastyle or fourth commissure.
Lower molars with no special peculiarities ; protoconid higher
than hypoconid in all three teeth ; hypoconid with greater basal
area than protoconid in m. 1 and m 2 , but with less in m 3 ; cingulum
well developed, forming a slight postero-internal cusp behind
entoconid.
Measurements. — Adult male from Henley-on-Thames, Oxford-
PIPISTRELLUS
209
shire, England : head and body, 44 ; tail, 32 ; tibia, 10- 4; foot,
6 ; forearm, 30 • 4 ; thumb, 4 • 4 ; third finger, 52 ; fifth finger,
38. Two adult males from Sorrento, 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, ll - 4 and
11-4; 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, 12 and 11 - 2 ; width of ear, 8-4 and 8. Adult
female from Burgos, 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.
Specimens 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 Nlmes, the last wrongly marked
type of migrans Crespon) ; St. Genies, Gard, 1 (Mottaz) ; Marseilles, 1
(U.S.N.M.)
Germany : Bonn, 4 ; Ingelheim, Bheinhessen, 1 ; Magdeburg, Saxony, 1 ;
Berlin, 1 ; Budolstadt, Bavaria, 3 ; Niesky, Silesia, 1.
Austria-Hungary : Transylvania, 1 ; Zara, Dalmatia, 1.
Switzerland : Geneva, 15 (Mottaz) ; Buchillon, Vaud, 4 (Mottaz) ;
Morat, Pribourg, 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 Genoa, 10 (U.S.N.M. and Genoa) ;
Isola Giglio, 2 (Genoa) ; 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 locality, 1 ; Ustica Island, 1
(U.S.N.M.). K ' .
Sardinia : Cagliari, 17 (B.M. 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 Seal, Madrid, 2 ; Granada, 1 ; Seville, 1 ; Alcala, near
Seville, 1. ,
Remarks. — Pipistrellus pipistrellus is the smallest as well as
one of the commonest and most generally distributed of European
210
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212
OHIEOPTBKA
bats. Superficially it may be distinguished from the almost
equally small Myotis mystacinus by its smaller ears and shorter
legs ; but for positive discrimination from the members of the
genus Pipistrellus 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.
Go. 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 6, 2 9.
Tring, Hertfordshire.
Hon. N. C. Rothschild (r).
9. 2. 19. 1-4.
9.
Lilf ord, Northampton-
shire.
■ Lord Lilf ord (p).
11. 1. 1. 125.
2 9.
Chelmsford, Essex.
M. Christy and E. L.
Thompson (p).
11. 1. 3. 23-24.
9 juv. al.
London.
Dr. A. Giinther (p).
74. 7. 6. 1.
S.
Wimbledon, Surrey.
C. H. B. Grant (c & p).
11. 1. 3. 27.
juv.
New Forest, Hamp-
shire.
Col. J. W. Yerbury (c & p).
11. 1. 3. 389.
Sal.
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.
Upsala, Sweden.
(Kolthoff.)
Lord Lilf ord (p).
11. 1. 1. 26-28.
S.
Hillerod, Zealand,
Denmark.
0. Thomas (c & p).
96. 6. 7. 1.
,5,9.
Boulogne, Pas - de -
Calais, France.
0. Thomas (c & p).
98. 1. 9. 1-2.
S, 9 al.
, Bonn, Rhineland,
Dr. A. Giinther (p).
Germany.
9.
Ingelheim, Rhein-
hessen.
C. H. Hilgert (c).
8. 11. 2. 3-4.
Sal.
Magdeburg, Saxony.
Dr. W. Wolterstorff (p).
92. 12. 1. 2.
dal.
Berlin.
Dr. A. Giinther (p).
C. G. Danford and J. A.
66. 2. 1. 22.
Sal.
Transylvania, Hun-
74. 7. 4. 6.
gary-
Brown (c & p).
i.
Zara, Dalmatia.
Lord Lilf ord (p).
11. 1. 1. 29.
2al.
Ciudad Real, Spain.
A. Cabrera (p).
8. 7. 23. 2-3.
$.
Burgos, Prov. Burgos
. G. S. Miller (c).
8. 8. 4. 14.
S.
Silos, Burgos.
G. S. Miller (c).
8. 8. 4. 15.
lal.
Campiglio, Tirol,
Italy.
G. C. Champion (c & p).
96. 8. 7. 1.
6 juv. al
. Sicily.
46. 6. 15. 6.
Ual.
Aristano, Cagliari,
Sardinia. (C.
Krausse.)
Hon. N. C. Rothschild (p)
. 7. 5. 24. 11-24.
2 juv. al
. Sardinia. (P.Bonomi.
) E. N. Buxton (p).
95. 4. 16. 4-5.
3$.
Tatoi, Athens, Greece.
(C. Mottcus.)
, Hon. N.C. Rothschild (p).
8. 10. 2. 19-21.
$, 8 9.
Kephissia, Athens.
C. Mottaz (c).
8. 11. 3. 4-7.
PIPISTRELLUS 213
PIPISTRELLUS NATHUSH Keyserling and Blasius.
1839. V[espertilio~] nathusii Keyserling and Blasius, Wiegmann's Archiv
fur Naturgesch., 1839, I, p. 320 (Berlin, Germany).
1857. Vesperugo nathusii Blasius, Saugethiere Deutschlands, p. 58.
1878. Vesperugo abramus Dobson, Catal. Ohiropt. Brit. Mus., p. 226 (Part :
not of Temminck).
1900. Pipistrellus nathusii Mehely, Monogr. Chiropt. Hungarise, p. 276.
1905. Vesp[erugo] nathusii var. unicolor Fatio, Arch. Soi. Phys. et Nat.,
Geneve, 4th ser., xrx, p. 510, May, 1905 (Geneva, Switzerland).
Type in Geneva Museum.
1910. Pipistrellus abramus Trouessart, Paune 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 Pipistrellus pipistrellus (forearm,
32 to 35 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.
External characters. — In general and apart from the animal's
less diminutive size, the external characters are essentially as in
Pipistrellus pipistrellus. Ear larger and broader with more
obtuse apex, more evidently 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 less well defined than in P. pipistrellus. 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. pipistrellus, 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
CHIROPTEEA
tendency away from the cinnamon and raw-umber tints toward
a clearer brown more resembling Ridgway's mars-brown. Mem-
branes less blackish than in P. pipistrellus, the wing from foot
nearly to fifth finger with a noticeable pale border about 1 mm.
in width, similar to that present in P. huhlii, but less sharply
denned and less nearly white.
Skull. — The skull is less diminutive than that of Pipistrellus
pipistrellus, its general size about as Myotis mystacinus. General
form less robust than in P. pipistrellus, the width of brain-case
barely one-half greatest length, but more contrasted with that of
rostrum. Dorsal profile as in the smaller species, but with more
evident anterior concavity and posterior convexity, the anterior
edge of interparietal indicated by a slight transverse constriction.
Other details of form essentially as in P. pipistrellus.
Teeth. — As compared with those of Pipistrellus pipistrellus the
teeth throughout show a tendency toward slenderness and height.
Inner upper incisor noticeably more slender than that of P.
pipistrellus, and with less well developed secondary cuspj 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. pipis-
trellus, a slight space usually present in
median line, another between i 2 and i s , and
another between i s and canine ; outer edge
of i x slightly overlapping « 2 ; in form the
teeth are not peculiar. Upper canine like
that of P. pipistrellus, except that the
greatest diameter of its crown is only about
half length of anterior border of shaft.
Fig. 36. Mandibular canine high and slender, its apex
Pipistrellus 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 border. Upper
premolars as in P. pipistrellus, 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. pipistrellus.
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, 1 2 ; width of ear, 1 1 . Average and extremes of six
adults from Buchillon, Vaud, Switzerland : tibia, 1 2 • 9 ( 1 2 • 6-1 4) ;
foot, 7-3 (6-8-7-6); forearm, 33 (32-4-34-6); 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, 11 and 11-4. For cranial measurements see
Table, p. 222.
Specimens examined. — Thirty-three, from the following localities : —
Fbance : St. Gilles, Gard, 1.
Gebmany: Berlin, 1; Bavaria, 1 (U.S.N.M.); Ingelheim, Bhein-
hessen, 1.
Switzerland: Geneva, 6 (Mottaz and Geneva, including type of
unicolor Patio) ; Montreux, Vaud, 1 (Mottaz) ; Buchillon, Vaud, 6 (Mottaz) ;
Neuchatel, 1 (U.S.N.M.) ; Canton TJri, 1 ; St. Gothard, Uri, 1 (U.S.N.M.).
Aostbia-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.).
Remarks. — Though readily distinguishable from the other
European members of the genus by its cranial and dental
characters, Pipistrellus nathusii is superficially much like P. pipis-
trellus. 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.
9. Ingelheim, Bheinhessen. C. Hilgert (c). 8. 11. 2. 3.
2 al. Palics, Bacser, Hungary. Budapest Museum (b). 0. 4. 9. 1-2.
<5. Canton Uri, Switzerland. Tomes Collection. 7. 1. 1. 398.
i, 9. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 20-21.
9. Borzoli, Liguria, Italy. Marquis G. Doria (c & p). 5. 12. 15. 7.
PIPISTRELLUS KOHLII Kuhl.
1819. Vespertilio kuhlii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 199 (Triest).
1835. Vesp[ertilio~] alboUmbatus Kiister, Isis, p. 75 (Cagliari, Sardinia).
1837. Vespertilio vispistrelhis Bonaparte, Iconogr. Faun. Ital., ±, fasc. xx
(near Borne, Italy). Type in British Museum.
1837. Vespertilio alcythoe Bonaparte, Iconogr. Faun. Ital., I, fasc. xxi
(Sicily.) Type in British Museum.
1838. Bomicia calcarata Gray, Mag. Zool. and Bot., n, p. 495 (locality
unknown).
1841. Pipistrellus marginatus Bonaparte, Iconogr. Fauna. Ital., Indie.
distrib., nomencl. mod. (Substitute for alboUmbatus).
1844. Vespertilio marginatus Wagner, Schreber's Saugthiere, 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
CHIROPTEKA
1840. V[espertilio] Ursula Wagner, Schreber's Saugthiere, Suppl., i, p. 505
(Morea, Greece).
1857. Vesperugo kuhtti Blasius, Saugethiere Deutschlands, p. 63.
1878. Vesperugo kuhlii Dobson, Catal. Chiropt. Brit. Mus., p. 230.
1886. [Vesperugo kuhlii] var. albicans Montioelli, Atti Soo. Ital. Soi. Nat.,
Milano, xxvu, p. 200, March, 1886 (Caivano, Naples, Italy).
1886. [Vesperugo kuhlii] var. pullatus Monticelli, Atti Soc. Ital. Sci. Nat.,
Milano, xxvn, p. 200, March, 1886 (Bella Vista, near Portici,
Naples, Italy).
1900. Pipistrellus kuhlii M6hely, Monogr. Chiropt. Hungariae, 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.
Diagnosis. — Size about as in Pipistrellus nathusii (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 quite 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. pipistrellus, less
slender than in P. nathusii ; 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
denned whitish border.
External characters. — General form very similar to that of
Pipistrellus pipistrellus, the wing similarly narrow as compared
with that of P. nathusii. 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. nathusii, the posterior border noticeably
convex. Wing, foot, calcar and tail as in P. pipistrellus.
Fur and colour. — Quality and distribution of fur essentially
as in Pipistrellus pipistrellus, 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.
Shull. — The skull resembles that of Pipistrellus nathusii in
size, but its form is even more robust than that of P. pipistrellus.
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. nathusii, but not so much flattened as in P. savii. Narial
emargination more abruptly narrowed posteriorly than in the
PIPISTRELLUS
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 parallel with alveolar line.
Teeth. — Inner upper incisor essentially as in Pipistrellus
■pipistrellus 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. pipistrellus, 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 above
and below intermediate in form between those
of P. pipistrellus and P. naihusii. Anterior , . ... ,
upper premolar crowded inward from tooth-
row and closely wedged between canine and H , |( ,
large premolar which are nearly or quite in Pivistre uZkubZn.
contact ; the small tooth is usually though not Anterior teeth x 5.
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. pipistrellus and P. naihusii, but somewhat more robust.
Measurements. — 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, 31 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
CHIROPTBRA
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PIPISTRELLUS
219
Specimens examined. — One hundred and eight, from the following
localities : —
France : St. Gilles, Gard, 2 ; Nimes, Gard, 3 (Mottaz) ; St. Genies,
Gard, 4 (Mottaz) ; Marseilles, 1 (U.S.N.M.).
Switzerland : Coremmo, Tioino, 1 (Mottaz) ; Lugano, Ticino, 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
Corleone, Sicily, 1 (U.S.N.M.) ; Ustica Island, Sicily, 4
Sicily, 1 (type of alcythoe Bonaparte). '
Sardinia : Gagliari, 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.
IS:
S.N.M.)
S.N.M.)
Bemarhs. — Pipistrellus Ttuhlii is easily recognizable by tbe
form and relative size of tbe upper incisors. Externally it may
usually be known by tbe sharply denned wbitisb border to tbe
wing membrane, tbough too mucb reliance sbould not be placed
on this character alone. Many specimens from Sardinia are
lighter in colour than those from the mainland. These represent
the albolimbatus 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. Gilles, Gard,
France.
i. Siena, Italy.
(S. Brogi.)
3 6, 9. Rome. (C. Coli.)
skeleton with- Sicily. (Prince Bona-
out skull. parte.)
skeleton with- Near Rome, Italy,
out skull. (Prince Bonaparte.)
S, ?. Corfu, Greece.
(G. Mottaz.)
3 <5. Argostoli, Cephalonia.
(C. Mottaz.)
Patras. (C. Mottaz.)
<5, ?.
2 S.
San Cristobal,
Minorca, Balearic
Islands.
G. S. Miller (c). 8. 8. 4. 127.
Dr. E. Hamilton (p). 98. 10. 2. 2.
G. Barrett-Hamilton 11. 1. 2. 16-1P.
Tomes Collection. 7. 1. 1. 730.
(Type of V. alcythoe Bonaparte.)
Tomes Collection. 7. 1. 1. 729.
(Type of V. vispistellus Bonaparte.)
J. I. S. Whitaker (p). 8. 10. 1. 4-5.
J. I. S. Whitaker (p). 8. 10. 1. 1-3!
Hon. N. C. Rothschild 8. 10. 2. 17-18.
O. Thomas and R. I. 0. 7. 1. 29-30.
Pocock (c & p).
PIPISTRELLUS SAVH Bonaparte.
1837. Vespertilio savii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx (Pisa).
Type in British Museum.
1837. VespertiUo aristippe Bonaparte, Iconogr. Faun. Ital., I, fasc. xxi
(Sicily).
1837. Vespertilio leucippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Sicily). Type in British Museum.
1838. Vespertilio bonapartii Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvn,
p. 226 (Tuscany).
220
CHIEOPTBBA
1844. Vesp[ertilio] nigrcms Greapon, Paune Meridionale, i, p. 24 (Nimes,
Gard, Prance).
1853. Vesperugo maurus Blasius, Wiegmann's Arohiv fur Naturgesch.,
1853, 1, p. 35 (Central chain of the Alps).
1857. Vesperugo maurus Blasius, Saugethiere Deutschlands, p. 67.
1872. V[espertilio] agilis Patio, Paune Vert. Suisse, Append, au vol. I,
p. iii (Alternative name for V. savii Bonaparte, ex Savi MS.).
1878. Vesperugo maurus Dobson, Catal. Chiropt. Brit. Mus., p. 218.
1904. Vespertilio ochromixtus Cabrera, Mem. Soc. Espafi. Hist. Nat., n,
p. 267 (Sierra de Guadarrama, Madrid, Spain).
1910. Pipistrellus savii and P. savii ochromixtus Trouessart, Paune Mamm.
d'Europe, pp. 13-14.
Type 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
Pipistrellus 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 and 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 Pipistrellus kuhlii
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
Pipistrellus sari
Nat. size.
conspicuous, a peculiar dull brownish ochraceous-buff; adult female :
light tips as conspicuous as in last, dull brownish cream-buff.
Skull. — The skull is slightly larger than that of Pipistrellus
nathusii and P. huhlii, 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
Verpertilio murinus. Dorsal profile of skull as
in P. pipistrellus and P. lcuhlii, 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
general outline, aside from the notch caused
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. Jnihlii,
but with angular process less curved and relatively longer than
in the other European species.
Teeth. — Incisors both above and below essentially as in
Pipistrellus pipistrellus, except that inner upper tooth has the
secondary cusp somewhat better developed.
Canines with no special 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
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
any 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, 5 and 5-6;
third finger, 54 and 56 ; fifth finger, 42 and 41 ; ear from meatus,
12 - 4 and 12 • 6 ; width of ear, 12 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.
C2»
Flfl. 39.
Pipistrellus savii.
Anterior teeth, x 5.
222
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224
CHIROPTERA
Adult male from Escorial, Spain (paratype of ochromixtus Cabrera) :
head and body, 48 • 6 ; 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.N.M.) ; no exact locality, 1 (type) ; Palermo, Sicily, 8
(U.S.N.M.) ; Sicily, no exact locality, 1 (type of leucippe Bonaparte) ;
Ustica Island, Sicily, 1 (U.S.N.M.).
Prance : 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, Madrid, 1 (paratype of ochromixtus Cabrera).
Remarks. — This species is readily distinguishable among the
European members of the genus Pipistrellus by the peculiar form
of the tragus, apart from its very pronounced cranial and dental
characters. Its colour gives it a superficial resemblance to
Ejptesicus 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 perhaps twice described as a member of the
genus Eptesicus*
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.
<5. Borzoli, Liguria. Marquis G. Doria 5. 12. 15. 6.
(c & p).
skeleton with- Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 732.
out skull. parte.) (Type of species.)
skeleton with- Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 731.
out skull. parte.) (Type of V. leucippe Bonaparte.)
9 al. Prance. Purchased (Lefebre). 46.1.2.12.
S. El Escorial, Madrid, A. Cabrera (p). 8. 7. 23. 1.
Spain. (Paratype of V. ochromixtus Cabrera.)
Genus EPTESICUS Rafinesque.
1820. Eptesicus Rafinesque, Annals of Nature, p. 2 (melanops = fuscus).
1829. Cnephseus Kaup, Entw.-Gesch. u. Natiirl Syst. Europ., Thierwelt, i,
p. 103 (serotinus).
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur-
geseh., 1839, i, p. 312 (part).
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fur Natur-
gesch., 1839, i, p. 313 (Sub-genus of Vesperugo, part). Not Vesperus
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. When 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 Vesperugo
caucasicus (Zool. Anzeiger, xxiv, p. 462, August 5, 1901) was based on
similar specimens.
EPTESICUS
225
1841. Noctula Bonaparte, Iconogr. Faun. Ital., I, faso. xxi, in account of
Vespertilio alcythoe (serotinus).
1856. Cateorus Kolenati, AUgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, n, p. 131 (serotinus).
1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, n, p. 131 (part).
1857. Vesperus Blasius, Saugethiere Deutschlands, p. 51 (Sub-genus of
Vesperugo), part.
1858. Amblyotus Kolenati, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwissenseh. Classe, xxix, p. 252 (atratus = nilssoni).
1863. Aristippe " Kolenati, Beitriige zur Kenntniss der Phthiriomyiarien,
Petersburg, 1863 " (part, included both discolor = murinus and
nilssoni).
1866. Pachyomus Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvn, p. 90,
February, 1863 (pachyomus).
1870. Nyctiptenus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 424 (smithii).
1878. Vesperus Dobson, Oatal. Chiropt. Brit. Mus., p. 184 (Sub-genus of
Vesperugo), part.
1892. Adelonycteris H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1891,
p. 466, January 19, 1892 (part; substitute for Vesperus, pre-
occupied).
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,
October, 1897 (part).
1900. Eptesicus Mehely, Monogr. Chiropt. Hungariee, p. 219 (part).
1907. Eptesicus Miller, Families and Genera of Bats, p. 207, June 29, 1907.
Type species. — Eptesicus melanops Rafinesque = Verpertilio
fuscus Beauvois.
Geographical distribution. — Europe, Asia (except Malay region),
Australia, Africa, Madagascar ; America from southern Canada
southward (except Lesser Antilles).
Characters. — Dental formula : i £?. c J4, P m iS; m S = 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 3 variable in form, usually with well developed
metacone and three commissures in the smaller species, but with
metacone and third commissure obsolete in larger forms. Skull
without special peculiarities of form or structure, the rostrum
flattish or more usually rounded off 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).
Bemarlis. — Among European bats the members of the genus
Eptesicus may be distinguished by their dental formula combined
with a simple Pipistrellus-like ear and not specially modified
skull. The group is nearly related to Pipistrellus through
P. savii, in which the small premolar is occasionally absent and
not infrequently so minute as to be concealed by the gum. About
forty-five species are known, three of which occur in Europe.
Q
226 CHIEOPTEKA
KEY TO THE EUROPEAN SPECIES OP EPTESICUS.
Porearm less than 40 mm. ; condylobasal length of
skull less than 16 mm. ; a distinct line of demar-
cation between colours of upper and lower surfaces
of neck E. nilssonU, p. 234.
Porearm more than 45 mm. ; condylobasal length of
skull more than 17 mm. ; no line of demarcation
between colours of upper and lower surfaces of
neck.
Condylobasal 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. Vespertilio serotinus Schreber, Saugthiere, i, pi. liii (Description, i,
p. 167, 1775, under name: Die Blasse Pledermaus). Prance,
based primarily on "La Se>otine" of Daubenton, Hist. Acad.
Royale des Sci., 1759, p. 377. 1765.
1776. Vespertilio serotine P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 16 (Based on "die Blasse Pledermaus " of Schreber).
1827. Vespertilio wiedii Brehm, Ornis, Heft in, p. 24 (Renthendorf,
Thviringen, Germany).
1827. Vespertilio okenii Brehm, Ornis, Heft in, p. 25 (Renthendorf,
Thiiringen, Germany).
1844. Vesp\eartilio\ incisivus Crespon, Paune Meridionale, I, p. 26 (Nimes,
Gard, Prance).
1857. Vesperugo serotinus Blasius, Saugethiere Deutschlands, p. 76.
1863. [Cateorus serotinus'] var. typus Koch, Jahrb. des Vereins fur Natur-
kunde im Herzogthum Nassau, xvni, p. 466 (Wiesbaden, Nassau,
Germany).
1863. [Cateorus serotinus] var. rufescens Koch, Jahrb. des Vereins fiir
Naturkande im Herzogthum Nassau, xvm, p. 466 (Preiburg,
Breisgau, Germany).
1878. Vesperugo serotinus Dobson, Catal. Chiropt. Brit. Mus., p. 191.
1885. Vesperus serotinus var. transylvanus Daday, Orvos-Termeszettudo-
manyi Erteseto', Kolozsvar, x, p. 275 (Also-Szoes, Szolnok-Doboka,
Hungary).
1886. Vesperus serotinus var. transsylvanus Daday, Verhandl. u. Mittheil-
ungen des Siebenbiirgischen Vereins fur Naturwissensch. in
Hermannstadt, xxxvi, p. 81.
1900. Eptesicus serotinus M6hely, Monogr. Chiropt. Hungarise, p. 209.
1904. Vespertilio serotinus insularis Cabrera, Mem. Soc. Espafl. Hist. Nat.,
n, p. 263 (Minorca, Balearic Islands).
1904. Vespertilio isabellinus Cabrera, Mem. Soc. Espafl. Hist. Nat., n,
p. 264 (southern Spain). Not of Temminck.
1904. Vespertilio boscai Cabrera, Mem. Soc. Espafl. Hist. Nat., n, p. 265
(Muchamiel, Alicante, Spain).
1910. Eptesicus serotinus, E. serotinus transsylvanus, and E. boscai Troues-
sart, Paune Mamm. d'Europe, pp. 20-22.
Type locality. — France.
Geographical distribution. — Central and southern Europe
from England and Denmark to the Mediterranean ; eastward
into Asia.
Diagnosis. — Size rather large (forearm more than 45 mm.,
EPTBSICUS 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 characters. — 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
swellings, 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, e±tending 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 off 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 Nyctalus 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-defined, its terminal lobe well
developed though small. Tail extending to between shoulders
when laid forward, the last vertebra and distal third of penulti-
mate vertebra free from membrane.
Fur and colour. — Pur 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.
Ground 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 with
inconspicuous lighter (buffy) tips ; underparts slightly paler,
sometimes approaching ochraceous-buff, 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
Skull. — General aspect of skull robust and flattened, with
widely spreading zygomata, but rather narrow brain-case and
rostrum. Dorsal profile 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 Nyctalus 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 bullae small, the transverse diameter con-
siderably less than distance between bullse. 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 lachrymal fora-
men ; rostral depth at front of orbit
less than distance from orbit to outer
incisor ; anteorbital foramen less re-
duced than in Nyctalus 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 posterior 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 Nyctalus noctula. Inner upper
incisor about half as high as canine, its crown area about one-
FIG. 40.
Epte&ieus serotinus. Nat. size.
BPTBSICUS
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 ; cingulum well developed, but
without cusps. 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, tbe 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 1 and i 2 overlapping 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 2 ) ; cross section of crown somewhat triangular, the
posterior angle occupied by a low tubercle in i 2 and i 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 Nyctalus 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 i 3 , where it becomes abruptly
obsolete, terminating in a slight tubercle corresponding to the
cusp present in Nyctalus and Pipistrellus. Upper premolar with
crown area about equal 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, flat-
tened-concave on inner side, a well developed external and antero-
internal longitudinal groove ; crown with slight anterior and
more marked 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
constricted at middle ; protocone robust, not very high ; no true
hypocone, but region which it would occupy indicated by slight
230
CHIEOPTEEA
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 3 with crown area less than half that of m 1 , 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. m 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 finger, 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, 1 6. 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;
Wingham, Dover, Kent, 1 ; Yalding, Kent, 5 (B.M. and U.S.N.M.) ; Isle
of Wight, 5.
France: Barsao, Gironde, 1 (U.S.N.M.); near Nlmes, Gard, 2 (Nlmes;
type of incisivus Crespon, and a specimen wrongly marked type of paluslris
Crespon).
Germany : Ingelheim, Bheinhessen, 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.M.) ; Tubingen, Silesia, 2.
Austria-Hungary : Moravia, 1 ; Csall6kbz-Somorja, Pressburg, Hun-
gary, 1 ; Budapest, Hungary, 2 ; Transylvania, 5 ; Zara, Dalmatia, 2.
Eoumania : 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, 1 (U.S.N.M.) ; Volterra,l (U.S.N.M.) ; Eome,3; SanMartino
al Cimino, Borne, 1 (Genoa) ; Ustica Island, Sicily, 1 (U.S.N.M.).
Sardinia : Cagliari, 2 (Genoa).
Spain: Pa] ares, Leon, 1; Seville, 3; Muchamiel, Alicante, 1 (Madrid;
type of boscai Cabrera). <
Bernarks. — 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
EPTESICHS
231
distinguish any local geographical forms. Specimens from Seville
representing the isabellinus of Cabrera I am unable to separate
from true serotinus ; the type of boscai 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.*
i. Keuley, Surrey, England.
2 9.
Sal.
1st.
3?.
Sal.
2.
1.
1.
6,9.
2<5, S.
2al.
lal.
2al.
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IS.
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lal.
t.
S.
Sal.
lal.
1.
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, Rheinhessen,
Germany.
Magdeburg, Saxony.
(Wolterstorff.)
Strass, Burgheim, Bavaria.
(KOrbitz.)
Tubingen, Silesia.
Moravia, Hungary.
Csall6k6z-Somorja, Press-
burg.
Budapest.
Transylvania.
Zara,Dalmatia. (E.Blos.)
Bustenari, Prahova, 840 m.
Roumania. (W.Dodson.)
Patras, Greece. (C.Mottaz.)
W. R. Ogilvie-Grant
(c&p).
G. H. B. Grant (c & p).
G. Donker (c & p).
H. Reid (c & p).
W. R. Ogilvie-Grant
(c & p).
Rev. C. Bury (c & p).
P. Bond (c&p).
Tomes Collection.
Tomes Collection.
C. Hilgert (c).
Patras.
Athens.
(0. Mottaz.)
Florence, Italy.
Rome. (C. Coli.)
Pajares, Leon,
(N. Gonzalez.)
Seville.
Seville.
Europe.
Spain.
Lord Lilford (p).
Lord Lilford (p).
Dr. A. Giinther (p).
Purchased (Parreys).
Budapest Museum (e).
Budapest Museum (e).
0. G. Danford and J. A.
Brown (o & p).
Lord Lilford (p).
Lord Lilford (p).
Hon. N. C. Rothschild
ro-
cs. Mottaz (c).
Hon. N. C. Rothschild
Florence Museum (e).
G.Barrett-Hamilton (p).
0. 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. 15. 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. 3. 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 sodalis 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.
Type locality. — Bustenari, Prahova, Roumania (in Carpa-
thians, alt. 840 m.).
Geographical distribution. — 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
CHIEOPTEEA
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234
CHIBOPTEEA
Diagnosis. — Similar to Eptesicus 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
measurements see Table, p. 233.
Specimens examined. — Two, from the following localities : —
Switzerland : St. Gothard, 1 (U.S.N.M.).
Roumania : Bustenari, Prahova, 1 (type).
Remarks. — The two specimens on which this species is based
indicate the existence of an animal bearing much the same
relationship to Eptesicus serotinus as Nyctalus noctula to N.
maximus.
6. Bustenari, Prahova, 840 m. Lord Lilford (p). 4. 4. 6. 1.
Boumania. (W. Dodson.) (Type of species.)
eptesicus nilssonii Keyserling and Blasius.
1836. Vespertilio kuhlii Nilsson, Ilium. Pig. Skand. Fauna, pt. 17, pi. 34
upper figure. Not of Kuhl, 1819.
1838. Vespert[ilio] borealis Nilsson, Tllum. Pig. Skand. Fauna, pt. 19,
pi. 34 (renumbered 36) upper figure. Not of P. L. S. Miiller, 1776
(Scandinavia).
1839. V[espertilw] nilssonii Keyserling and Blasius, Wiegmann's Archiv
fur Naturgesch., 1839, p. 315 (Mountains of Scandinavia. Based
on the V. kuhlii of Nilsson, 1836).
1857. Vesperugo nilssonii Blasius, Saugethiere Deutschlands, p. 70.
1858. Amblyotus atratus Kolenati, Sitzungsber. kais. Akad. Wissensch.
Wien, Math.-Naturwissensch. Classe, xxix, p. 252 (Altvater,
Austrian Silesia, alt. 2400-4600 ft.).
1878. Vesperugo borealis Dobson, Catal. Chiropt. Brit. Mus., p. 203.
1894. Vesperugo nillsoni (sic) Bhoads, Beprint Ord's N. Amer. Zoology,
Append., p. 3.
1907. E[ptesicus~\ nilssoni Miller, Families and Genera of Bats, p. 209,
June 29, 1907.
1910. Eptesicus nilssoni Trouessart, Faune Mamm. d'Europe, p. 23.
Type locality. — 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 longer,
EPTESICUS
235
extending to nostril when laid forward, though of the same
general form as in E. serotinus ; tip less narrowly rounded off,
and flattened or concave portion of posterior border less con-
spicuous ; tragus relatively shorter and wider, though not
essentially different in form, its greatest width nearly equal to
length of anterior border ; transverse striations on inner surface
of conch obsolete. 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 fully 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 dark 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-buff in evident contrast
with ground colour, the light tips most numerous behind shoulders,
and forming a noticeable mantle over middle of back ; under-
pays rather strongly contrasted light yellowish brown, between
the wood-brown and ochraceous-buff 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 differs 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 posterior
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 bullae slightly
larger than in E. serotinus. Interorbital fig. 41.
region relatively less constricted than in Epteswm nOnomi.
E. serotinus, but of essentially the same form ; ^«*- size -
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
CHIEOPTBRA
noteworthy peculiarities, its general outline less narrow than in
the related species.
Teeth. — Inner upper incisor as in Eptesicus 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 portion of palate, instead
of at right angles to main axis. Lower incisors less crowded
than those of E. serotinus, their imbrication distinct but not
unusual, the general outline of the row broadly V-shaped ; in form
the individual teeth show no special peculiarities ; i 2 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 and
metacone less evident ; m 3 with crown area about two-thirds that
of m l , its longitudinal diameter through metacone slightly more
than half transverse 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 in ™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 finger 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.).
Germany : Wernigerode, Saxony, 2.
Austeia-Hungaby : Csall6k6z-Somorja, Pressburg, 1.
Switzerland : Grotte de Vallorbe, Vaud, 1 (Mottaz).
Remarks. — This species is readily distinguishable from Eptesicus
serotinus by its smaller size and by the conspicuous pale tips to
the hairs of back. From Vesperiilio murinus, which resembles it
in colour pattern, the narrow ear, more yellowish (less whitish)
hair-tips on back, and slightly smaller size distinguish it super-
ficially. Pipistrellus savii, which often has almost exactly similar
coloration, is a much smaller animal (forearm, 31 to 33 instead
of 38 to 40).
1. Norway. (Collett.) E. B. Alston (p.) 81. 6. 9. 2.
4?. Upsala, Sweden. (Eolthoff.) Lord Lilford (p). 11.1.1.39-42.
6, 9. Wernigerode, Saxony, Ger- Dr. W. Wolterstorfl 0. 2. 8. 3-4.
many. (c & p).
1, Csall6koz-Somorja, Press- Budapest Museum (e.) 94.3.1.14.
burg, Hungary.
BPTESICUS
237
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CHIROPTEKA
Genus VESPERTILIO Linnaeus.
1758. Vespertilio Linnaeus, Syst. Nat., i, 10th ed., p. 31 (murinus by
tautonymy).
1839. Yesperugo Keyserling and Blasiua, Wiegmann's Archiv fur Natur-
gesoh., 1839, 1, p. 312 (part).
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fur Naturgesch.,
1839, i, p. 313. Sub-genus of Yesperugo (part).
1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, n, p. 131 (part).
1857. Yesperus, misprinted Yesperugo Blasius, Saugethiere Deutschlands,
p. 69 (Sub-genus of Yesperugo).
1863. " Aristippe Kolenati, Beitrage zur Kenntniss der Phthiriomyiarien,
Petersburg, 1863" (part).
1872. Marsipoleemus Peters, Monatsber. k. preuss. Akad. Wissenseh.,
Berlin, p. 260 (Sub-genus of Yesperugo for albigularis = murinus).
1878. Yesperus Dobson, Catal. Chiropt. Brit. Mus., p. 183. Sub-genus of
Yesperugo (part).
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,
October, 1897 (part).
1900. Yespertilio Mehely, Monogr. Chiropt. Hungarise, p. 219 (part).
1907. Vespertilio Miller, Families and Genera of Bats, p. 209, June 29, 1907.
Type species. — Vespertilio murinus Linnaeus.
Geographical distribution. — Forested northern portion of the
Palsearctic 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.
Remarks. — 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 Lasionycteris.
VESPERTILIO MURINUS Linnasus.
1758. [Vespertilio] murinus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden).
1819. Yespertilio discolor Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 187 (Vienna, Austria).
1857. Yesperugo discolor Blasius, Saugethiere Deutschlands, p. 73.
1872. Vesperus (Marsipoleemus) albigularis Peters, Monatsber. k. preuss.
Akad. Wissenseh., Berlin, p. 260 (Type supposed to have been
taken in Mexico ; for its reference to this species see Mehely,
Monogr. Chiropt. Hungarise, pp. 229, 341, 1900).
1878. Yesperugo discolor Dobson, Catal. Chiropt. Brit. Mus., p. 204.
1885. Vesperus siculus Daday, Orvos-Termeszettudomanyi Erteseto, Kolozs-
var, x, p. 275 (Homorod-ALmas cave, Hungary). See Mehely,
Monogr. Chiropt. Hungarise, pp. 229, 346, 1900.
VESPERTILIO
239
1886. Vesperus siculus Daday, Verhandl. u. Mittheilungen des Siebenbiirg-
ischeniVereins fiir Naturwissensch. in Hermannstadt, xxxvi, p. 82.
1897. Vesjoertilio murinus Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,
p. 382, October, 1897.
1910. Eptesicus siculus and Vespertilio murinus Trouessart, Faune Mamm.
d'Burope, pp. 23 and 25.
Type locality.- — Upsala, Sweden.
Geographical 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 1£ 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 shallowly 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-like,
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 nilssonii, but light tips to
hairs very pale, almost whitish cream-buff, producing a decidedly
" frosted " appearance ; underparts cream-buff, 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
CHIROPTERA
Skull. — The skull combines the short, smooth brain-case of
Eptesicus nils8onii with the broad, flat rostrum of E. serotinus, but
differs conspicuously in aspect from both in the very deep,
Nyctalus-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
Ves P e,-tiKo Zurinm. to f ront of inner incisor J . anteorbital foramen
Nat. size. moderately large, over point of 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
Vespertilio murinus show certain notable peculiarities. Inner
upper incisor similar to that of Eptesicus 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,
producing 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. nilssonii. Upper canine with transverse
diameter slightly greater than longitudinal diameter, the reverse
of the condition in the two species of Eptesicus ; lower canine
with same peculiarity in outline of crown though to a less degree.
Upper premolar with longitudinal diameter of crown relatively
VESPEBTILIO
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CHIKOPTERA
less than in the European species of Eptesicus, and antero-internal
basal cusp better developed ; lower premolars more crowded and
compressed than in E. nilssonii, and crown area of first relatively-
smaller. Molars as in E. nilssonii except that columnar hypocone
of m 1 and m 2 stands out more prominently from outline of
protocone.
Measurements. — 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 finger, 73, 76 and 74; fifth finger, 52, 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.
Specimens 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.).
Austria-Hungary : Csallokoz-Somorja, Pressburg, 1.
Switzerland : Morat, Fribourg, 1 (Mottaz).
Locality unknown : One ; type of albigularis Peters (Berlin).
'Remarks. — Superficially this bat somewhat resembles Eptesicus
nilssoni. It is readily distinguishable, however, by its slightly
larger size (forearm, 43 to 45 instead of 38 to 40), low, rounded
ear, and by 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 Museum (e). 94. 3. 1. 15.
burg, Hungary.
Genus NYCTALUS Bowdich.
1825. Nyctalus Bowdich, Excursions in Madeira and Porto Santo, p. 36
(verrucosus).
1829. Pterygistes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt,
i, p. 99.
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur-
gesch., 1839, i, p. 312 (part).
1842. Noctulinia Gray, Ann. and Mag. Nat. Hist., x, p. 258 (part ; contained
proterus = noctula anAfulvus = Scotophilus kuhUi).
1856. Panugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,
neue Polge, ii, p. 131 (noctula and leisleri).
1857. Vesperugo Blasius, Saugethiere Deutschlands, p. 49 (part).
NYCTALUS 243
1878. Vesperugo Dobson, Oatal. Ohiropt. Brit. Mus., p. 183 (part).
1898. Noctulinia H. Allen, Proo. U.S. National Museum, xvi, p. 30,
June 13, 1893.
1897. Pterygistes Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,
October, 1897.
1899. Euvcsperugo Acloque, Faune de France, Mammiferes, p. 32 (part,
inoluded noctula, 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 Bowdich.
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 bed
FIG. 43.
Nyctalus Trmxvmus (a), N. noctula, (b), N. leisleri (c), and AT. azoreum (d). Nat. size.
striction, and with large anterior palatal emargination ; fifth finger
much shortened, scarcely exceeding metacarpal of fourth or third.
Remarks.- — This genus is well differentiated 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 EUBOPEAN 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,^. 252.
Condylobasal length of skull 13 to 14-2 mm. ; fore-
arm 35 to 42 mm. (Azores) N. azoreum, p. 254.
R 2
244 CHIROPTEHA
NYCTALUS MAXIMUS Fatio.
1781. ? VespertiHo lasiopterus Schreber, Saugethiere, pi. lvii b. See
Thomas, Ann. and Mag. Nat Hist., 8th ser., vin, pp. 379-380,
September, 1911.
1827. ?? Vespertilio ferrugineus Brehm, Ornis, Heft III, p. 26 (Benthendorf,
Thiiringen, Germany).
1869. [Vesperugo noctula'] var. maxima Patio, Paune Vert. Suisse, i, p. 57.
Co-type in Geneva Museum.
1900. Pterygistes maximus Miller, Proo. Biol. Soc. Washington, xiir,
p. 156, June 13, 1900.
1910. Nyctalus maximus Trouessart, Faune Mamm. d'Europe, p. 19.
Type 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 characters. — 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 rising 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
finger, 119 and 123 ; fifth finger, 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. — Pive, from the following localities : —
Switzeeland: Amsteg, Uri, 1 (Geneva; co-type).
Itam : Pisa, 3 (B.M. and U.S.N.M.) ; Ravenna, 1.
Ninni, Atti Soc. Ital. Sci, Nat., Milano, xxvi, p. 109, 1883.
NYCTALUS 245
Bemarks. — Nijctalus maximus is readily distinguished among
the bats of Europe by its large size. It is at present a rare and
little known animal.
9 al. Pisa, Italy. Dr. A. Senna (b). 94. 6. 3. 3.
<5 al. Ravenna. Florence Museum (b). 85. 7. 6. 9.
6 al. Lidth de Jeude Coll. 67. 4. 12. 337.
NYCTALUS NOCTULA Schreber.
1774. Vespertilio noctula Schreber, Saugthiere, I, pi. lii ; description, i,
p. 166, under name : Die Speckmaus (Prance ; based primarily
on "La Noctule" of Daubenton, Hist. Acad. Boyale des Sci.,
Paris, 1759, p. 376. 1765).
1776. Vespertilio lardarius P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 15 (Prance; 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, 1892), p. 5 Nomen
nudum : " Great Bat."
1818. Vespertilio proterus Kuhl, Ann. Wetterau, Gesellsch. Naturk, rv
(= Neue Ann., i), pt. 1, p. 41 (Substitute for noctula).
1829. Vespertilio rufescens Brehm, Isis, p. 643 (Jena, Thiiringen, Germany).
1841. Vespertilio noctula Bonaparte, Iconogr. Faun. Ital., i, Ind. Distrib.
(sp. illustr.).
1841. Noctula serotina Bonaparte, Iconogr. Faun. Ital., I, Ind. Distrib.
(nomencl. moderna).
1844. Vesp[ertMo] palustris Orespon, Faune Meridionale, i, p. 22 (marshes
near Nlmes, 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. Pterygistes noctula Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,
p. 384, October, 1897.
1910. Nyctalus noctula Trouessart, Faune Mamm. d'Europe, p. 18.
Type locality. — France.
Geographical distribution. — 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 yellowish 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
CHIEOPTBRA
way from eye to nostril, its breadth when flattened decidedly
greater than height above crown ; anterior border of conch
abruptly 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 conih 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 fifth
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
\vith 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 mm.) free.
Fur and colour. — Fur dense and velvety, the hairs at middle
of back only about 5 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 upper parts a rather
dark yellowish brown, near the wood-brown and cinnamon of
Ridgway, the hairs showing a faint clouding of prouts-brown in
certain lights, their basal portion a light dull isabella-colour ;
underparts scarcely different 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 truncate 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,
NYCTALUS
247
the region on each side of it flat or slightly 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 denned anteriorly but communicating posteriorly with
vacuity between cochlea and basioccipital ; auditory bullae well
developed but of moderate size, the transverse diameter about
equal to distance between bullae. Interorbital region deeply
constricted, short hour-glass shaped, the lachrymal 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 premolar and first
molar, lachrymal foramen slightly above
and behind it, on orbital rim. Palate
rather short, owing to the large size of
the anterior emargination, the posterior , """ "'. ,
, ,, .... ° -. , ... ■ i ii j? Nyctalus noctvZa.
edge ot which is on level with middle ot Nat. size,
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 process ; angular process
moderately long, about on level with alveolar line, its main
axis directed rather abruptly outward, its distal extremity
obliquely widened.
Teeth. — 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-.iondary
cusp relatively much larger than that of inner incisor, cingulum
well developed, bearing a small but distinct antero-external cusp
on opposite side of concavity from secondary cusp. The two
248
CHIROPTEEA
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 (J shaped
row between canines, the crowns of i l and i 2 compressed, trifid,
that of i 3 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
cusp 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 reniforni 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 crowns 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 special peculiarities,
the styles low but well developed, the W -pattern normal, m 3 with
crown area distinctly more than half that of m 1 , 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
compared 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 m 2 an£ l
approximate equality in m 3 ; cingulum well developed but not
forming basal cusp 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 locality : 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 the following
localities : —
England : Bowdon, Cheshire, 1 ; Stoke, Staffordshire, 1 ; Oundle,
Northamptonshire, 4 ; Bedfordshire, 1 ; Trumpington, Cambridgeshire, 1 ;
Cambridgeshire, no exact locality, 1 ; Henley-on-Thames, Oxfordshire, 1 ;
Sandringham, Norfolk, 1 ; Kingsbury, Middlesex, 1 ; Chelmsford, Essex, 1 ;
Wellyn, Hertfordshire, 1 ; Tring, Hertfordshire, 3 ; Wandsworth Common,
Surrey, 1 ; Shalford, Surrey, 2 ; Earlsfield, Surrey, 5 ; Kew Gardens,
Surrey, 1 (U.S.N.M.) ; Fulham Park, London, 1 ; Eastwell, Kent, 2 ; St.
Leonards, Sussex, 1 ; Farnborough, Hampshire, 1 ; Wareham, Dorset-
shire, 1.
Holland : Oosterbeek, Guelderland, 1.
Germany : 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.
Austeia-Hungary : Csall6k6z-Somorja, Pressburg, Hungary, 2.
Roumania : Gageni, 1 ; Bustenari, 1 ; no exact locality, 1 (U.S.N.M.).
Switzerland : Geneva, i (Geneva and Mottaz) ; St. Margarethen,
Appenzell, 8 (B.M. and U.S.N.M.).
Spain : Seville, i ; no exact locality, 2.
Italy : Pisa, 18 (U.S.N.M.) ; Florence, 4 (Mottaz) ; Vallombrosa, near
Florence, 1 (U.S.N.M.) ; Siena, 1 ; Rome, 2.
Greece : Corinth, 12 (U.S.N.M.).
Remarks. — This is one of the most widely distributed and
abundant of the larger European bats. From Eptesicus 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.
$. Aberia, Merionethshire, Col. Yerbury (c & p).
"Wales.
?. Bowdon, Cheshire, Eng- G. Barrett-Hamilton
land. (T. A. Coward.) (p).
2 <$, 2 9. Oundle, Northampton- Lord Lilford (p).
shire.
9.' Bedfordshire. J. S. Elliot (c & p).
1. Trumpington, Cambridge- G. Barrett-Hamilton
shire. (p).
9 al. Cambridgeshire. Rev. L. Jenyns (c & p).
11.
1.3.
26.
11.
1.2.
100.
11.
1.1.
126-
127.
11.
1.1.
L30.
129-
U.
1. 3.
391.
11.
1.2.
101.
250
CHIEOPTERA
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„ not worn.
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„ moderately worn.
„ slightly worn.
„ moderately worn.
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,, not worn.
„ slightly worn.
„ not worn.
„ moderately worn.
„ slightly worn.
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NYCTALUS
251
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CHIROPTEEA
<5. Henley-on-Thames, Ox-
fordshire.
£ st. Sandringham, Norfolk.
9. Chelmsford, Essex.
?. Wellyn, Hertfordshire.
2 i, 9 st. Tring, Hertfordshire.
2 S. Shalford, Surrey.
3 <5, 2 9. Barlsfield, Surrey.
(J st.
Fulham Park, London.
Eastwell, Kent.
9. St. Leonards, Sussex.
6. Farnborough, Hampshire.
9. Wareham, Dorset.
6. Oosterbeek, Guelderland,
Holland.
9. Niesky, Silesia, Germany.
(W. Boer.)
6. Schwartzburg, Thiiringen,
200 m.
3 c?. Strass, Burgheim,
Bavaria. (Kiirbitz.)
1 al. Stuttgart, Wurtemburg.
2. Csall6k6z-Somorja, Press-
burg, Hungary.
S. Gageni, Prahova, Bou-
mania. (W. Dodson.)
9. Bustenari, Prahova. (W.
Dodson.)
2<5,39. St. Margarethen,
Appenzell, Switzer
land. (E.H.Zollikofer.
4 al. Seville, Spain. (A. Ruiz.
2 al. Spain.
S juv. al. Borne.
Heatley Noble (c & p).
H.M. King Edward VII.
M. Christy and L. E.
Thompson (o & p).
C. H. B. Grant (c & p).
Hon. N. C. Rothschild
(o & p).
W. B. Ogilvie - Grant
(o & p).
C. H. B. Grant (c & p).
J. Saunders (o & p).
C. H. Caton - Haigh
(c & p).
B. Butterfield (c & pi.
H. S. Soott (c & p).
W. T. Blanford (c & p).
0. Thomas (o & 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. 1. 3. 40.
96. 7. 27. 1.
11. 1. 3. 41.
11. 1. 3. 33.
I. 5. 22. 1-3.
II. 1. 3. 34-
35.
11. 1. 3. 28-
32
87. 7. 22. 1.
11. 1. 3. 36-
37.
11. 1. 3. 392.
11. 1. 3. 39.
11. 1. 3. 38.
98. 2. 1. 5.
99. 1. 9. 7.
95. 4. 18. 1.
11. 1. 1. 30-
32.
66. 2. 1. 6.
94. 3. 1. 16-
17.
4. 4. 6. 4.
4. 4. 6. 5.
4. 4. 5. 15-19.
94. 5. 8. 1-4.
72. 8. 21. 1-2.
85. 7. 6. 8.
NYCTALUS LEISLERI Kuhl.
1818. Vespertilio leisleri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., I), pt. 1, p. 46.
1818. Vespertilio dasycarpos Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 49 (Alternative name for leisleri).
1839. Vespertilio pachygnathus Michahelles in Wagner, Schreber's Saug-
thiere, Suppl., i, pi. lv b (Dalmatia. See Pitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii,
Abth. I, p. 222, 1870).
1857. Vesperugo leisleri Blasius, Saugethiere Deutschlands, p. 56.
1878. Vesperugo leisleri Dobson, Catal. Chiropt. Brit. Mus., p. 215.
1910. Nyctalus leisleri Trouessart, Paune Mamm. d'Europe, p. 19.
Type locality. — Hanau, Hessen-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.
Colour. — The colour is usually a brown, darker than in N.
noctula, nearly the prout-brown of Ridgway, with or without a
lighter, wood-brown tinge, and usually showing drab or isabella
reflections in certain lights, the underparts not so dark as back ;
hairs everywhere blackish (seal-brown) through basal half. Ears
and membranes (dry) blackish. Two skins are a pale buffy 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
Nyctalus 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. — External measurements of adult male from
Co. Wicklow, Ireland : head and body, 60 ; tail, 39 ; tibia, 16-2 ;
foot, 8 • 2 ; forearm, 42 ; third finger, 78 ; fourth finger, 48 ; ear
from meatus, 13. Adult female from Co. Armagh, Ireland :
head and body, 63 ; tail, 42 ; tibia, 16 • 6 ; foot, 8 ; forearm, 42 ;
third finger, 76 ; fifth finger, 49 ; ear from meatus, 14 ; width
of ear, 14. Two adults from Welford, Warwickshire, England :
forearm, 40 • 4 and 41-4. Adult from Meiringen, Bern, Switzer-
land : forearm, 40. For cranial measurements see Table,
p. 255.
Specimens examined. — Eleven, from the following localities : —
Ibeland : Belfast, 1 ; Bray, Co. Wicklow, 2 ; Co. Armagh, 2.
England : Mexbro', Yorkshire, 1 ; Welford, Warwickshire, 2 ; Cleeve
Priory, 1 ; no exact locality, 1.
Roumania : Bustenari, Prahova, 1.
. Switzerland : Meiringen, Bern, 1.
Remarks. — Though its smaller size is the most obvious
character of this species as compared with N. noctula, the
bicolored fur, the different relative sizes of the upper incisors,
and the more abrupt convexity of the mandibular incisor series
would by themselves be quite sufficient to distinguish it.
9.
Belfast, Ireland.
G. Barrett-Hamilton (p.)
1. 3. 15. 1.
<5al.
Bray, Wicklow.
J. E. Harting (c & p).
90. 2. H. 1.
Sal.
Armagh.
Dr. G. E. Dobson (c & p).
89. 11. 12. 5,
Sal.
Armagh.
R. M. Barrington (c & p).
74. 5. 28. 8.
2 9.
Welford, Warwickshire,
Tomes Collection.
7. 1. 1. 386-
England.
387.
6.
Cleeve Priory, Warwick-
shire.
Tomes Collection.
7. 1. 1. 388.
1.
England.
Dr. Leach (p).
63. A.
i.
Bustenari, Prahova, 840 m.
Roumania. (W.Dodson.)
Lord Lilford (p).
4. 4. 6. 6.
1,
Meiringen, Bern, Switzer-
land.
Tomes Collection.
7. 1. 1. 389.
254
CHIEOPTEEA
NYCTALUS AZOREUM Thomas.
1901. Pterygistes azoreum Thomas, Ann. and Mag. Nat. Hist., 7th ser., vm,
p. 33, July, 1901. Type in British Museum.
Type locality. — St. Michael, Azores.
Geographical description. — Azores.
Diagnosis. — Smaller than Nyctalus 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 azoreum 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 prout-brown, the hairs with darker,
ill-defined slaty basal area, and usually with lighter tips, the
lighter colour ranging in different individuals from raw-umber
through wood-brown to a dull light buff, 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.
Shull 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 3 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, 16 ■ 6 ;
foot, 7 ; forearm, 42 ; thumb, 6 ; third finger, 72 ; fifth finger,
48 ; ear from meatus, 12 ; width of ear, 11. For cranial measure-
ments see Table, p. 255.
Specimens examined. — Seventeen, from the following localities in the
Azores : — Teroeira, i (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.
<5 al. St. Michael, Azores. F. Du Cane Godman 65. 10. 2. 1.
(o & p). (Type of species.)
2(5,2?. St. Michael. (W. B. 0. Hon. W. Eothschild 3.6.5.1-3,7.
Chant.) (p).
255
00
§
00
t
B
o
Teeth slightly worn.
„ not worn.
„ 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
CHIROPTEEA
<5, 2 9. Reguinho, Teroeira. Hon. W. Rothschild 3.6.5.4-6.
(W. B. 0. Grant.) (p).
2 1, 3 9. Magdalena,Pioo. (W.B.O. Hon. W. Rothschild 3.6.5.10-14.
Grant.) (p).
<5,S. Calheto, St. George. Hon. W. Rothschild 3.6.5.8-9.
(W. B. 0. Grant.) (p).
Genus PLECOTUS Geoffroy.
1816. ? Macrotus Leach, Catal, Spec. Indig. Mamm. and Birds Brit. Mus.
(Willoughby Society reprint, 1882), p. 1. Nomen nudum: "Euro-
pean Longear, Macrotus europasus."
1818. Plecoius Geoffroy, Description de l'Egypte, n, p. 112. For date see
Sherborn. Proc. Zool. Soc. London, 1897, p. 288.
1857. Plecotus 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 Linnaeus.
Geographical distribution. — Temperate Europe, Asia and
northern Africa.
Characters. — Dental formula : i *=|, c Jd, pm ||, m || = 36.
Skull with large, elongate and rounded brain-case, and slender
but normally formed rostrum ; auditory bullse 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 ;
nostrils opening upward, their orifices continued backward by
slit-like prolongations, the wart-like outgrowths on muzzle not
specially prominent.
Remarks. — 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 (Miniopterus)
the ears are unusually short. Seven species are known, one of
which occurs in Europe.
PLECOTUS AURITUS Linnaeus.
1758. Vespertilio auritus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden).
1816. ? Macrotus europseus Leach, Catal. Spec. Indig. Mamm. and Birds
Brit. Mus. (Willoughby Society reprint, 1882), p. 1. Nomen
nudum: " European Longear, Macrotus europseus."
1818. Plecotus auritus Geoffrey, Description de l'Egypte, n, p. 118.
1825. Vespertilio otus Boie, Isis, p. 1206 (Copenhagen, Denmark).
1826. Vespertilio cornutus Eaber, Isis, p. 515 (Jutland, Denmark).
1827. Plecotus communis Lesson, Man. de Mammal., p. 95 (Prance).
1828. P[lecotus] brevimanus Jenyns, Trans. Linn. Soc, London, xvi, p. 55
(Grunty Pen, Isle of Ely, Cambridgeshire, England).
1829. Plecotus vulgaris Desmarest, Paune Prancaise, p. 18 (France).
PLECOTUS 257
1829. [Vespertilio auritus] y austriacus Fischer, Synops. Mamm., p. 117.
Based on the " second variety" of Desmarest, Diet, des Sci. Nat.,
lviii, p. 51, 1829 (Vienna, Austria).
1832. Plecotus velatus I. Geoffroy, Guerin's Mag. de Zool., n, CI. i, pi. 2,
p. 5 (not numbered), footnote (Name applied by lapsus calami to
the common Plecotus of England).
1840. Plecotus megalotos Schinz, Europ. Fauna, i, p. 19 (Synonym of
auritus ; Brehm cited as authority).
1857. Plecotus auritus Blasius, Siiugethiere Deutschlands, p. 39.
1860. Plecotus kirschbaumii Koch, Achter Ber. Oberhess. Gesellseh.
Natur.- u. Heilkunde, Giessen, p. 40, May, 1860 (Dillenburg,
Oberhessen, Germany).
1863. [Plecotus auritus] var. typus Koch, Jahrb. des Vereins flir Natur-
kunde im Herzogthum Nassau, xviii, p. 406 (Wiesbaden, Nassau,
Germany).
1863. [Plecotus auritus] var. montanus Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviii, p. 406 (Westerwald, Nassau,
Germany).
1863. [Plecotus auritus] var. brevipes Koch, Jahrb. des Vereins fiir Natur-
kunde im Herzogthum Nassau, xviir, p. 407 (Substitute for kirsch-
baumii Koch). t
1878. Plecotus auritus Dobson, Catal. Chiropt. Brit. Mus., p. 178.
1910. Plecotus auritus Trouessart, Faune Mamm. d'Europe, p. 12.
Type locality. — Sweden.
Geographical distribution. — Prom Ireland eastward into Asia,
and from the Mediterranean north to the Scandinavian Peninsula.
Diagnosis. — Characters as in the genus ; auditory bulla?
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 narrow, 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 forward, 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 from
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
258
CHIROPTEKA
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) equal 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
one-third length of latter, the membrane thin and semi-trans-
parent, joining foot at base of outer toe ; third and fourth meta-
carpals 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
border of interfemoral 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.
Skull. — 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 bullae. 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
g AaaawJiy 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,
rh without special features, a narrow groove at
Plecotus mmtus. each side of basioccipital ; auditory bulla
Nat. stee. relatively larger than in any other European
bat, its diameter about three times the
space between bullae. Interorbital constriction narrow, scarcely
hour-glass shaped, the anterior upper rim of orbit with narrow,
slightly projecting edge ; lachrymal region scarcely 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 ofi ; rostral
depth at front of orbit about equal to distance from orbit to
PLBCOTDS 259
alveolus of inner incisor ; anteorbital foramen small, close to rim
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
rather 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
postero-internal portion ; cutting edge obscurely trifid ; i 3 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, forming 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 pi 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 about half that of canine, that of
posterior premolar nearly equal 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 squarish with antero-external corner rounded off ; 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 narrow internally,
the protocone with rather short base, the hypocone absent or
barely indicated ; metacone decidedly higher than paracone ;
styles and commissures well developed, though mesostyle of m l
s 2
260
CHIROPTEEA
does not extend outward to level of parastyle and metastyle ;
W-P a ttern normal ; m 3 with three cusps and three commissures,
its crown area about half that of m 2 , 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 ; cingulum forming a
barely indicated postero-internal cusp behind entbconid ; second
triangle of ro 3 much narrower than first, but about equal to it
in area.
Measurements. — Adult female from Chelmsford, Essex,
England: head and body, 42; tail, 45; tibia, 17; foot, 9'6;
forearm, 39 ; thumb, 7 ; third finger, 69 ; fifth finger, 52 ;
ear from meatus, 36 ; 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,
Segovia, 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, 1 7 • 6. Adult female
from Florence, 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 ; Wohurn Sands, Bedfordshire, 1 ; Pembrokeshire,
no exact locality, 1 ; Chelmsford, Essex, 1 (U.S.N.M.) ; Tring, Hertford-
shire, 1 ; Boxmore, Hertfordshire, 1 ; Hoddesdon, Hertfordshire, 1 ;
London, 4 ; Godalming, 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 : Oosterbeek, Guelderland, 4.
France : Near Barcelonnette, Basses-Alpes, 1 (Mottaz).
Germany : Wernigerode, 3 ; Ummerstadt, Thiiringen, 2 ; Magdeburg, 2 ;
Niesky, Silesia, 4 ; Damsdorf , Silesia, 1.
Austria-Hungary: Csall6k6z-Somorja, Pressburg, 7.
Switzerland : Geneva, 2 (Mottaz) ; Lausanne, Vaud, 1 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Boudry, 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: Borzoli, near Genoa, 1 (Genoa) ; Florence, 1 (Mottaz) ; Rimini, 1;
Home, 1 ; no exact locality, 1 ; Sicily, 1.
Spain : Silos, Burgos, 2 (U.S.N.M.) ; La Granja, Segovia, 1.
Bemarhs. — Plecotus auritus is so readily distinguished from all
other European bats by the great size of the ears that no special
comparisons are required.
<J, 3 ?. Montrose, Scotland. J. H. Coward (c & p). 11. 1. 3. 48-51
c5 st. Antrim, Ireland. Hon. N. C. Rothschild 1. 9. 3. 5.
PLECOTUS 261
2 6 al. Longford. Dr. G. E. Dobson (o & p). 76. 2. 12. 2.
<;. Pembroke, Wales. W.E. deWinton(o&p). 11. 1. 3. 393.
1 st. Winfarthing, Norfolk, E. Markham (c & p). 96. 7. 27. 3.
England.
6. Diss, Norfolk. (W. E. C. H. B. Grant (p). 11. 1. 3. 394.
Sherrin.)
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). 86. 9. 27. 1.
shire.
i. Tring, Hertfordshire. Dr. E. Hartert (c & p). 11. 1. 3. 56.
9 al. London. No history.
6. London. Sergt. Brown (o & p). 11. 1. 3. 57.
i. S. Kensington, Middle- 11. 1. 3. 54.
sex.
i. Godalming, Surrey. W. T. Blanford (c & p). 11. 1. 3. 58.
1. Bonehuroh, Isle of Bev.C. A. Bury (c & p). 4*. 3. 29. 6.
Wight.
c5, 9. Honiton, Devonshire, G. C. Shortridge(c&p). 11.1.3.52-53.
300 ft.
4 <5, 4 9 al. Devonshire. No history.
2 6, 2 9. Oosterbeek, Guelder- 0. Thomas (o & p). 98. 2. 1. 1-4.
land, Holland.
9. Wernigerode, Harz, W. Wolterstorff (c&p). 0.2.8.1.
Germany.
(5, 9. Wernigerode, Harz. Lord Lilford (p). 11. 1. 1. 45-46.
(Wolterstorff.)
S. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 47.
(Wolterstorff.)
(5,9. Ummerstadt, Thiirin- Lord Lilford (p). 11.1.1.43-44.
gen. (Schuchardt.)
£. Niesky, Silesia. (W. Lord Lilford (p). 99. 1. 9. 1.
Baer.)
6 al. Damsdorf, Silesia. Dr. A. Giinther (p). 66. 8. 1. 10.
7. Csall6k6.z-Somorja, Budapest Museum (e). 94.3.1.5-11.
Pressburg, Austria-
Hungary.
<5. Thurgau, St. Gallen, 0. Thomas (p). 4. 4. 5. 14.
Switzerland. (Zolli-
kofer.)
1. Switzerland. Leon 0. Galliard (p). 75. 9. 20. 2.
9 al. Bimini, Italy. Florence Museum (e). 85. 7. 6. 2.
<5. Borne. (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, M. de la Escalera (c). 8. 7. 30. 5.
Spain.
1. Europe. Leyden Museum (e). 37. 4. 28. 25.
262
CHIEOPTEEA
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BARBASTELLA
263
Genus BARBASTELLA Gray.
1821. Barbastella Gray, London Med. Repos., xv, p. 300, April 1, 1821.
1825. Barbastellus Gray, Zool. Journ., n, p. 243, July, 1825.
1839. Synotus Keyserling and Blasius, Wiegmann's Archiv fur Naturgesch.,
1839, I, p. 305.
1857. Synotus Blasius, Saugethiere Deutschlands, p. 42.
1878. Synotus Dobson, Catal. Chiropt. 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. — Vespertilio barbastellus Schreber.
Geographical distribution. — Northern Africa ; central and
southern Europe ; west-central Asia to the Himalayas.
Characters. — Dental formula : i H, c J=?! pm H> i»n= 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 bullse 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 members of
the family Vespertilionid.se. Two species are known, one peculiar
to the Himalayan region, the other occurring in Europe.
BARBASTELLA BARBASTELLUS Schreber.
1774. Vespertilio barbastellus Schreber, Saugthiere, 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. Vespertilio barbastelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.-
Band, p. 17 (Burgundy).
1836. Barbastellus daubentonii Bell, Hist. Brit. Quadr., pt. i, p. 63
(Burgundy ; based primarily on Daubenton).
1838. Barbastellus communis Gray, Mag. Zool. and Bot., n, p. 495,
February, 1838 (Renaming of barbastellus).
1857. Synotus barbastellus Blasius, Saugethiere Deutschlands, p. 43.
1878. Synotus barbastellus Dobson, Catal. Chiropt. Brit. Mus., p. 176.
1897. Barbastella barbastellus Miller, Ann. and Mag. Nat. Hist., 6th ser.,
xx, p. 385, October, 1897.
1910. Barbastella barbastellus Trouessart, Faune Mamm. d'Europe, p. 11,
Type locality. — Burgundy, France.
Geographical distribution. — Central and southern Europe,
264
CHIBOPTERA
west to England, north to southern Norway and Sweden, east
into Asia.
Diagnosis. — Characters as in the genus ; forearm about 38
to 40 mm.
External characters. — 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 upper lip behind
nostril 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
tip extending about 5 mm. beyond nostril when laid forward ;
anterior 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 except for an evident
flattening just below the rather abruptly rounded off and some-
what recurved tip ; posterior border abruptly concave above, then
nearly straight except for an abruptly projecting lobe near
middle (lobe usually about 1 • 5 mm. long Toy 1 mm. wide, its upper
border convex, its lower border concave) ; lower border of ear full
and rounded, but with no differentiated antitragus ; tragus large,
somewhat triangular in outline, its greatest width (slightly above
level of anterior base) equal to about two-thirds length of anterior
border, its upper portion rapidly 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 upper 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
BARBASTELLA 265
about as long as tibia, with small but evident terminal lobe and
posterior keel ; tail with extreme tip projecting beyond inter-
femoral membrane.
Colour. — General colour a very dark 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 region behind shoulders, those of underparts 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 base of under side of interfemoral
membrane pale ecru-drab or whitish smoke-grey nearly or quite
to base. Ears and membranes in dry specimens brown, 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 large brain-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-row. 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 bullse 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
rostrum ; lachrymal ridges rather prominent,
especially at their lower extremities. Zygoma
nearly straight, scarcely bowed outward beyond
level of outer surface of brain-case. Ante- n ,
orbital foramen relatively large, over anterior Barbastdla barba .
margin of m 1 . Rostrum short, its upper sur- status. Nat. size,
face flattened-concave, deeply emarginate in
front by the unusual backward extension of the upper 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 m 3 ,
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
auritus, though they are throughout smaller and more slender.
Incisors both above and below essentially as in Plecotus auritus
except that crown of outer upper incisor is nearly terete, and
266
CHIROPTERA
space between it and cingulum of canine is very narrow ; lower
incisors forming a less broadly U-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 cingulum
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 Plecotus
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 the following localities : —
England : Elton, Huntingdonshire, 1 ; Ellingham, Norfolk, 1 ; Arrow
Lodge, Warwickshire, 1 ; Alcester, Warwickshire, 1 ; Rugby, 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-Garonne, 1,
Germany : Biichenberg, Harz Mountains, 1 ; Niesky, Silesia, 4 ;
Bavaria, no exact locality, 3 (U.S.N.M.) ; 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 by the peculiar form
of the muzzle.
9. Llanelwedd, Radnorshire, H. E. Forrest (c & p). 4. 7. 3. 1.
Wales.
9. Elton, Huntingdonshire, Lord Lilford (p). 94. 9. 5. 1.
England.
6. Ellingham, Norfolk. E. R. Alston (p). 81. 6. 9. 1.
(Dr. W. M. Crowfoot.)
S. Arrow Lodge, Warwickshire. Tomes Collection. 7.1.1.348.
1. Alcester, Warwickshire. Tomes Collection. 7. 1. 1. 348.
(5 st. Rugby, Warwickshire. G. Dalgleish (c & p). 4. 10. 13. 2.
S. Oundle, Northamptonshire. Lord Lilford (c & p). 94. 4. 16. 1.
9 al. Epping, Essex. H. Doubleday (o & p). 44. 5. 1. 1.
<5. Milton, Hampshire. Rev. J". E. Eelsall (p). 11. 1. 3. 59.
BAKBASTELLA
267
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268
CHIKOPTERA
S. Swindon, Wiltshire. Rev. E. A. Goddard 8. 5. 12. 1.
(c & p).
<5. Caterille, Haute-Garonne, 0. Thomas (p). 6. 4. 1. 7.
1000 m. France.
(A. Robert.)
$, ?. Niesky, Silesia, 181 m. Ger- Dr. E. Hamilton (p). 9Y. 12. 4. 2-3.
many. (W. Baer.)
2 ?. Niesky, Silesia. (W. Baer.) Lord Lilford (p). 99. 1. 9. 2, 4.
S. Wernigerode, Saxony. Lord Lilford (p) . 0. 2. 8. 2.
(Wolterstorff.)
Sub-Family MINIOPTERINiE.
1878. Miniopteri Dobson, Catal. Chiropt. Brit. Mus., p. 170 (part).
1907. Miniopterinie Miller, Families and Genera of Bats, p. 227, June 29,
1907.
Geographical distribution. — Africa, southern Europe and
southern Asia, eastward to the Malay region, Japan and
Australia.
Characters. — Like the Vespertilioninse, but presternum with
median lobe enormously developed and forming the greater part
of the bone ; scapula with coracoid straight, directed con-
spicuously inward.
BemarJes. — The sub-family Miniopterinx, though widely dis-
tributed in the warmer portions of the Old World, is at present
known to contain the genus Miniopterus only.
Genus MINIOPTERUS Bonaparte.
1837. Miniopterus Bonaparte, Ioonogr. Faun. Ital., i, faso. xx, under
Vespertilio emarginatus (Sub-genus of Vespertilio).
1857. Miniopterus Blasius, Saugethiere Deutschlands, p. 45.
1866. Miniopteris Gray, Ann. and Mag. Nat. Hist., 3rd ser., xm, p. 91,
February, 1866.
1878. Miniopterus Dobson, Gatal. Chiropt. Brit. Mus., p. 347.
1892. Minyopterus Winge, Jordf undne og nulevende Flagermus (Chiroptera)
fra Lagoa Santa, Minas Geraes, Brasilien, p. 36.
1900. Minneopterus Lampe, Jahrb. Nassau Ver. Naturkunde, Jahrg. 53,
Catal. Saugeth.-Samml., p. 12.
1907. Miniopterus Miller, Families and Genera of Bats, p. 227, June 29,
1907.
Type species. — Vespertilio ursinii Bonaparte = V. schreibersii
Kuhl.
Geographical distribution. — Same as that of the sub-family.
Characters. — Cental formula : i H, c \^, vm |4 m t? = 34.
.i-3 1—1 ■*■ 3— .J 3—3
Skull with unusually high brain-case, and low, flattened rostrum.
Second phalanx of third finger nearly three times as long as first.
Ears short, separate, the upper margin (in European species)
appearing almost artificially truncate.
Remarks. — The peculiar shortening of the ears together with
the remarkable elongation of second phalanx of third finger readily
MINIOPTERUS
269
distinguish this genus among the members of the European fauna.
About a dozen forms have been described, one of which occurs
in southern Europe.
MINIOPTERUS SCHREIBERSII Kuhl.
1819. Vespertilio schreibersii Kuhl, Ann. Wetterau. Gesellsoh. Naturk., iv
(= Neue Ann., i), pt. 2, p. 185 (Hungary).
1837. Vespertilio ursinii Bonaparte, Ioonogr. Faun. Ital., I, fasc. xxi
(Monte Corno, Ascoli, Italy). Type in British Museum.
1841. Vespertilio orsinii Tern mi nek, Monogr. de Mamm., n, p. 179 (modi-
fication of ursinii).
1857. Miniopterus schreibersii Blasius, Saugethiere Deutschlands, p. 46.
1878. Miniopterus schreibersii Dobson, Catal. Chiropt. Brit. Mus., p. 348.
1910. Miniopterus schreibersi Trouessart, Faune Mamm. d'Europe, p. 34.
Type locality. — Kulmbazer Cave, mountains of southern
Bannat, Hungary.
Geographical distribution. — Southern Europe from the Iberian
Peninsula eastward, north to Switzerland and Hungary. Limits
of range not known.
Diagnosis. — Charaters as in the genus ; forearm about
43 mm.
External 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 equal to distance from eye 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 different 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
practically no " tip," the whole anterior border appearing like an
obliquely, almost artificially truncate extremity ; antitragus low,
obscurely marked off 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 exterior
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 fifth finger exceeding forearm by about one-fifth
length of latter, the tip unusually slender and elongate owing to
the great length of last bone of third finger ; third and fourth
270 CHIEOPTEHA
metacarpals sub-equal, about 3 mm. shorter than forearm ; fifth
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 body, included in the membrane to
extreme tip.
Fur and colour. — The fur is of a very soft, silky texture,
though rather short, the hairs at middle of back only about
7 mm. in length, those of head abruptly much 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 : (a) at base (3 mm.), a slaty mouse-grey ;
(&) 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
■Qf^iy 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,
fig. 47. slightly elevated posteriorly. Brain-case
Miniopterus schrewersii. -, jS j_j_i.lt i . __ 1
Nat. size. broadly ovate, the outline broken at each
side posteriorly by the slightly projecting
mastoid region ; sagittal crest low but 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 brain-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
MINIOPTERUS
271
at outer side of each pit ; auditory bulla moderately large, the •
transverse diameter about equal to distance between bullae.
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 off at sides, flattened concave along median
line, the narial emargination small, squarish, extending backward
about one-third of the distance to front of forehead ; rostral
depth at front of orbit about equal to distance from orbit to
outer incisor ; anteorbital foramen rather large, directly above
small premolar, 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 upper 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 spine 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-row, the posterior seg-
ment unusually 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-concave 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 t and i 2 with crown very low, the cutting edge
obscurely tried, the crown of each tooth wider posteriorly (in line
of booth-row) than anteriorly, but this more evident in second
than in first ; j 3 nearly terete, the middle and posterior cusps
enlarged 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
antero-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
CHIKOPTEEA
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 externally than
internally ; large upper premolar with crown area about equal
to that of first molar and double that of small premolar, its
anterior and posterior borders very slightly concave, the antero-
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 squarish, with antero-external angle tending
to become rounded off, particularly in the posterior tooth ; cusp of
first about half as high as canine, that of second slightly higher
than first, 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 inner edge of tooth, the anterior border nearly
straight ; protocone low but rather broad ; hypocone clearly
indicated but not distinct from posterior commissure of protocone ;
paracone and metacone nearer outer edge of crown than 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 anterior border, its three cusps and three
commissures normally developed. Lower molars with no special
peculiarities, the second triangle of m 3 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
tibia, 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 body, 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 93; 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.
MINIOPTBRUS
273
Specimens examined. — Three hundred and fifty-one, from the following
localities : —
Spain: Silos, Burgos, 9 (B.M. and U.S.N.M.); Seville, 1; Minorca,
Balearic Islands, 3 ; Majorca, Balearic Islands, 2.
France : Troubate, Hautes-Pyrenees, 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 oxact
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.M.); Spezia, 5 (U.S.N.M.); Monte Gorno, Ascoli, 1
(type of ursinii Bonaparte) ; Borne, 10 ; Velletri, Bome, 2 (U.S.N.M.) ; no
exact locality, 1 ; Elba Island, 5 (U.S.N.M.) ; Marsala, Sicily, 2.
Sardinia: Sassari, 3 (U.S.N.M.); Cagliari, 4 (U.S.N.M.); Mount
Gennargentu, 2 (U.S.N.M.).
Greece : Corinth, 17 (U.S.N.M.) ; Labyrinth, Crete, 1.
Remarhs. — 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 Miniopterus schreibersii as compared with other European
bats.
2. Silos, Burgos, Spain. Bev. S. Gonzalez (c). 88. 1. 5. 46-47.
<J, ?. Inca, Majorca, Balearic 0. Thomas and B. I. 0.7.1.4-5.
Islands. Pocock (c & p).
3 <5. San Cristobal, Minorca. 0. Thomas and B. I. 0. 7. 1. 32-34.
Pocock (c & p).
<$, 9. Troubate, Hautes- 0. Thomas (p). 6. i. 1. 8-9.
Pyrenees, 400 m.
Prance. (A.Robert.)
8 <5, 1 ? Dions, Gard, 70 m. 0. Thomas (p). 8. 8. 10. 1-9.
Prance. (0. Mottaz.)
1 al. Ofener Mountains, Budapest Museum (e). 94. 7. 18. 10.
Budapest.
9. Beri, Montenegro. 0. Thomas (p). 5. 8. 4. 6.
(L. FUhrer.)
9. Velgi, Czolo. 0. Thomas (p). 5. 8. 4. 5.
(L. FiXhrer.)
7 <5, 3 9. Rome. (C. Coli.) Lord Lilford (p). 11. 1. 2. 5-14.
Skeleton Monte Corno, Ascoli, Tomes Collection. 7. 1. 1. 735.
(without Italy. (Type of Vespertilio ursinii Bonaparte.)
skull).
2 £. Marsala, Sicily. 0. Thomas (p). 6. 8. 4. 24-25.
(A. Robert.)
S. Labyrinth, 200 ft. Miss D. Bate (c). 5. 12. 2. 9.
Crete.
274
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276 CHIEOPTBEA
Family MOLOSSIT)^.
1865. Molossi Peters, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 258.
1872. Molossidse Gill, Arrangement of the Families of Mammals, p. 17.
1878. Emballonuridas (part; Molossinx, part, Molossi) Dobson, Catal.
Cbiropt. Brit. Mus., p. 402.
1907. Molossidse Miller, Families and Genera of Bats, p. 241, June 29, 1907.
Geographical distribution. — Warmer parts of both hemispheres ;
in the Old World north to the Mediterranean region and
southern Asia.
Characters. — Essentially like the Vespertilionidae 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 conspicuously beyond membrane, a
character by which the only European member of the group may
be immediately recognized.
Bemarhs. — 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 Europe.
Genus NYCTINOMUS Geoffroy.
1818. Nyctinomus Geoffroy, Descr. de l'Bgypte, n, p. 114 (segyptiacus).
1821. Nyctinoma Bowdioh, Anal. Nat. Class. Mamni., p. 28 (Modification
of Nyctinomus).
1821. Nyctinomes Gray, London Med. Bepos., xv, p. 299, April 1, 1821
(Modification of Nyctinomus).
1822. Nyctinomia Fleming, Philos. of Zool., n, p. 178 (Modification of
Nyctinomus).
1842. Mops Lesson, Nouv. Tabl. B.egne Anim., p. 18 (Mops indicus Lesson
= Dysopes 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 (femorosaccus).
1907. Nyctinomus Miller, Families and Genera of Bats, p. 251, June 29, 1907.
Type species. — Nyctinomus segyptiacus Geoffroy.
Geographical distribution. — Warmer portions of both hemi-
spheres, north to the southern United States and to the Mediter-
ranean coast of Europe, east to the Philippines and Norfolk
Island.
Characters. — Dental formula : i tl or t?,* c t? , vm — , m s - = 30
2-iJ 3-3' 1-1' £ 2-a' 3-3
or 32 ; brain-case not unusually flattened, its depth at least half
its width ; bony palate with a small median anterior emargination
extending to behind level of roots of incisors.
Remarks. — As thus defined the genus Nyctinomus contains
* In the species occurring in Europe.
NYCTINOMUS 277
about forty species, two-thirds of which are peculiar to the Old
World, one of them occurring in the Mediterranean region of
Europe.
NYCTINOMUS TENIOTIS Rafinesque.
1814. Cephalotes teniotis Rafinesque, Pr6c. des D6couv. Somiol.,p.l2 (Sicily).
1825. Dinops cestoni Savi, N. Giorn. de' Letterati, Pisa, x, p. 235 (Pisa, Italy).
1840. Dysopes savii Sohinz, Europ. Fauna, i, p. 5 (Substitute for cestoni).
1871. [Dysopes cestonii] var. nigrogriseus Schneider, Neue Denkschr.
Schweiz. Gesellsch. Naturwiss., xxiv, p. 5 (articles separately
paged). Basel, Switzerland.
1877. Nyctinomus cestonii Dobson, Catal. Cbiropt. Brit. Mus., p. 423.
1891. Nyctinomus tseniotis Thomas, Proc. Zool. Soc, London, p. 182.
1897. [Dysopes] midas Schulze, Abh. Ges. Nat. iv, No. 10, p. 23 (Substitute
for cestoni). Not of Sundevall, 1842.
1910. Nyctinomus tseniotis Trouessart, Faune Mamm. d'Europe, p. 36.
Type locality. — Sicily.
Geographical distribution. — Mediterranean region of Europe
and northern Africa. Accidental ? at Basel, Switzerland.
Diagnosis. — Like the African Nyctinomus eegyptiacus but
larger, 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 beyond interfemoral
membrane, immediately distinguish the animal from all other
European bats. Muzzle projecting rather noticeably beyond
upper lip, obliquely truncate ; nostril pad well denned, 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
brush-like fringe of recurved 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 many deep
oblique wrinkles, its surface both above 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
irregularities, though the anterior border usually shows some
slight flattening and the posterior border is faintly concave above
278
CHIROPTERA
and near 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 ; posterior 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 equal 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 ; anterior and upper margin of tragus fringed with long
loose hairs. Wings longer and narrower 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 stiffened 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 projecting 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 specimens
blackish. Fringes on feet and hairy lines on ears, drab like
body.
Skull. — 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 bullae. 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 abrupt 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 first 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 lateral 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-defined
pits about as large as glenoid surface ;
auditory bullae 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 well differentiated
from interorbital region, the distance from orbit to front of pre-
maxillary about equal to depth at front of m 1 ; lachrymal ridge
short but well defined, the small, inconspicuous anteorbital fora-
men and minute lachrymal foramen opening forward under its
anterior edge, at level of middle of large premolar ; nares with
distinctly tubular lateral margins separated from roots of canines
by evident grooves, the dorsal emargination extending about half
way to lachrymal level. Palate moderately wide, noticeably
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 3 ; mesop-
terygoid space large, its width anteriorly equal to that of
Nyclinomua teniotis.
Nat. size.
280
CHIEOPTERA
temporal fossa at same 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 skull 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 of a large hypocone, quite distinct from commissure
of protocone, in m 1 and m 2 . Upper incisor simple, a little more
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-extemally ;
posterior surface of shaft somewhat
flattened ; space between incisor
and canine about 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 deeply 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.
Xyctinomiis leniotis. Teeth X 5.
NYCTINOMtTS
281
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282
CHIROPTEEA
of lower molars ; large upper premolar with crown area about
two-thirds that of m 1 , its antero-internal cusp high and distinct.
First and second upper molars alike in structure, 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.
Measurements. — Young adult male from Italy : head and
body, 87 ; tail, 56 ; free portion of tail, 20 ; tibia, 19 ; foot,
10-4; forearm, 60 - 4; third finger, 115; fifth finger, 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, 114; 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 following 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.
S. Italy (probably Pisa). Zoological Society (p). 10. 5. 24. 1.
{Savi.)
?. Greece. Parreys. 46. 6. 15. 121.
CARNIVORA
283
Order CARNIVORA.
1827. Carnivora Gray, Griffith's Cuvier. Anim. Kingd., v, p. 111.
Geographical distribution. — Continents and larger islands of
the entire world, Australia,* New Zealand, and the Antarctic
region excepted.
Characters. — Terrestrial (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 tuberculo-sectorial type, the posterior upper premolar
and anterior lower molar usually developed as special carnassial
or flesh-cutting teeth.
Bemarks. — 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 certain 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 recent families, five of which occur in Europe.
KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES
OF CABNIVOBA.
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) .. . Ursidas, 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) Felidie, p. 455.
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) Canidie, p. 303.
Represented in Australia by a species of Canis, probably introduced.
284
CARNIVOEA
Tooth-row relatively short (less than half condylo-
basal length of skull) ; number of teeth in
European members of family not more than 40.
Auditory bulla 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) Viverridm, p. 440.
Auditory bulla simple ; upper molars 1-1, the
crown wider internally than externally MusteUdas, 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) Melinse, 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) Lutrime, 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 pair capable of
shearing action ; upper carnassial with
posterior cusp narrow and trenchant ;
auditory bulla longer than broad; form
slender; feet digitigrade (Martens and
Weasels) Mustdinx, p. 364.
Dontition not highly trenchant ; small pre-
molars opposite, not capable of shearing
action, the points of all but pm? and pm i
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) Gulminte, p. 432.
Family TJRSIDiE.
1825. Ursidie Gray, Thomson's Annals of Philosophy, xxvi, p. 339,
November, 1825.
Geographical 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 type,
the crowns wide and flattened, with large terete cusps, the last
TJKSDS -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 antborbital
foramen as not to be at point of greatest mechanical efficiency ;
auditory bulla flattened, without septum ; form heavy ; size
large ; feet strictly plantigrade ; digits, 5-5.
Remarks. — The family Ursidse, containing the bears, is at
present 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 URSIDM.
Cheek-teeth relatively large ; incisors and canines not
specially enlarged and prehensive (Ordinary Bears) Ursus, p. 285.
Cheek-teeth relatively small ; incisors and canines
enlarged and unusually^ prehensive in character
(Polar Bears) .' Thalarctos, p. 297.
Genus URSUS Linnaeus.
1758. Ursus Linnaeus, Syst. Nat., I, 10th ed., p. 47 (arctos, by tautonymy).
1857. Ursus Blasius, Saugethiere Deutschlands, p. 196.
1864. Euarctos Gray, Proc. Zool. Soc. London, p. 692 (americanus).
1864. Myrmarctos Gray, Proc. Zool. Soc. London, p. 694 (eversmanni = arctos) .
1898. Ursarctos Heude, Mem. Hist. Nat. Emp. Chinois, iv, pt. i, p. 18
sis).
Type species. — Ursus arctos Linnseus.
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"J, pm ", m £§ = 42 ;
inner upper incisor well developed, permanent ; first, second and
third premolars in both jaws small, single-rooted, readily deciduous,
especially pm 2 , pm 2 and pm 3 ; molars large and robust, the length
of the two upper teeth together equal to width of palate.
Remarks. — The genus Ursus as thus restricted is a very
homogeneous group practically confined to the north temperate
region. The species are at present so imperfectly known 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 Linnseus.
1758. [Ursus] arctos Linnaeus, Syst. Nat.,*i, 10th ed., p. 47 (Sweden).
1772. [Ursus] ursus Boddaert, Kortbegrip van het zamenstel der Natuur,
i, p. 46 (Renaming of arctos).
1788. [Ursus arctos] a niger Gmelin, Syst. Nat., I, 13th ed., p. 100
(Northern Europe).
286
CABNIVORA
1788. [Ursus arctos] $ fuscus Gmelin, Syst. Nat., i, 13th ed., p. 100 (Alps).
1778. [Ursus aretes] y alhus Gmelin, Syst. Nat., i, 13th ed., p. 100
(Unknown ; based on the "ours blanc terrestre " of Buflon).
1792. U[rsus] arctos griseus Kerr, Anim. Kingd., p. 184 (Germany; also in
northern North America).
1797. Ursus arctos rufus Borkhausen, Deutsche Fauna, i, p. 46 (Swiss and
Tirolean Alps).
1798. Ursus badius Schrank, Fauna Boica, i, p. 55 (Forests on the
Bohemian boundary).
1808. Ursus fuscus Tiedemann, Zoologie, I, p. 374 (Substitute for arctos).
1814. Ursus alpinus Fischer, Zoognosia, in, p. 161 (Alps? Based on an
individual seen alive in Paris).
1820. Ursus arctos major 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] $ brunneus Billberg, Synopsis Faunae Scandinavise,
p. 15 (Northern Scandinavia).
1827. [Ursus arctos] j annulatus Billberg, Synopsis Faunae Scandinavise,
p. 15 (Northern Scandinavia).
1827. [Ursus arctos] 8 argenteus Billberg, Synopsis Faunae Seandinaviae,
p. 15 (Northern Scandinavia).
1827. [Ursus] myrmephagus Billberg, Synopsis Faunae Seandinaviae, p. 16
(Northern Scandinavia).
1828. [Ursus] formicarius Billberg, Synopsis Faunae Seandinaviae, 2nd ed.,
p. 16 (Renaming of myrmephagus).
1829. U[rsus] pyrenaicus 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] norvegicus Fischer, Synopsis Mamm., p. 142. Latinization
of "Ours de Norwege " F. Cuvier, Hist. Nat. des Mammif., n,
fasc. 7, 1819 (Norway).
1836. ? Ursus falciger Reichenbach, Regn. Anim. Icon., I, p. 32 (' ' Pyrenees" ;
afterwards supposed to be an individual of " 17. ferox." See Natur-
gesch. des In- und Auslands, Raubsaugeth., p. 299, 1852).
1842. Ursus pyrenieus F. Cuvier, Hist. Nat. des Mamm., Tabl. gen., p. 3
(Described in fasc. 44, 1824) (Asturias, Spain).
1847. ? Ursus euryrhinus Nilsson, Skand. Fauna, i, 2nd ed., p. 212 (Sweden ?
Type an individual raised in captivity).
1857. Ursus arctos Blasius, Saugethiere Deutschlands, p. 196.
1861. Ursus arctos aureus Fitzinger, Wissensch.-pop. Naturgesch. der
Saugeth., I, p. 372 (Norway).
1864. [Ursus arctos] var. 1. normalis Gray, Proc. Zool. Soc, London, p. 682
(Renaming of arctos).
1864. [Ursics arctos] sub-var. a. scandinavicus Gray, Proc. Zool, Soc,
London, p. 682 (Based on Nilsson, Ilium. Fig. Skand. Fauna,
pi. 23).
1864. ? [Ursus arctos] sub-var. c. rossicus Gray, Proc. Zool. Soc, London,
p. 682 (nomen nudum).
1864. [Ursus arctos] sub-var. f. yplonicus Gray, Proc. Zool. Soc, London,
p. 682 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, pi. xxn,
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
in the Zoological Gardens from 1852 to 1863 ").
uesds 287
1864. [Ursus arctos] var. 4. stenorostris Gray, Proc. Zool. Soc., London,
p. 685 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, 2nd var.,
pi. xxu, fig. 4).
1864. Myrmarctos eversmanni Gray, Proc. Zool. Soc, London, p. 695
(Norway).
1910. Ursus arctos, U. arctos formicarius, U. arctos alpinus, and 77. arctos
pyrenaicus Trouessart, Faune Mamm. d'Europe, pp. 67-68.
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 butfy, 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 pad
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 longest, 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
CAKNIVOEA
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 differences in colour have been
described, some of which are probably characteristic of geographical
races.
Skull. — General form of skull rather robust, the rostrum
KU. 50.
Ursus arctos.
X S.
moderately long (distance from orbit to front of premaxillary
contained about 2£ times in condylobasal length), the brain-case
uksus 289
deep (depth to level of under side of postglenoid process con-
siderably more than distance from tip of postorbital process to
middle of interparietal) but not unusually wide (mastoid breadth
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 this point to
slightly overhanging lambdal region ; ventral profile faintly and
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, the 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.
'290
CABNIVORA
runs to extremity of each postorbital 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 If times in mastoid breadth ; mastoid processes,
paroccipital processes, and condyles extending to about the same
via. 62.
XTrsus arctos. X J.
level. Floor of brain-case nearly flat, the basioccipital with
raised edges applied to inner surface of bullae, 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, postorbital 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 process usually somewhat swollen.
Zygomata moderately expanded, the greatest zygomatic breadth
opposite anterior glenoid edge ; orbital process well developed.
Rostrum equal to less than half condylobasal 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 1 or paracone of m 2 ;
palate narrow, its width between posterior molars contained about
3£ times in median length ; extension behind molars nearly
parallel-sided, its length equal to about three-quarters breadth ;
mesopterygoid space 1| times to twice as long as wide, its anterior
border squarish or rounded, 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 posterior portion
(measured to level of lower border) a little less than half length j
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 1 and i 2 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 developed and forming
a rudimentary nodule on inner and outer side of i 2 and on outer
side of j 1 near level of middle of anterior surface ; i 3 with crown
area nearly double that of i 2 , and height nearly half that of
canine, its anterior surface smoothly rounded but with pronounced
nodule on inner side, its posterior surface 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 i 3 ; posterior border slightly
292
CARMVORA
1 W
ism
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Fig. 53.
Ursus arctos. Teeth nat. size.
uitsus 293
concave, with inner, middle and outer ridges, the inner and middle
low and confluent in i v 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 developed 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 postero-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 developed
antero-external cusp nearly as high as main cusps of molars, a
rudimentary antero-internal cingulum cusp, and a tuberculated
ridge 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 molar 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
3
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296
CAENIVORA
oblique to it ; postero-external cusp slightly smaller than antero-
external, and contrast between outer and inner cusps less marked
than in m l ; 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 primitive 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 surface of the crown
occupied principally by a flattened, irregularly sculptured crushing
surface, the cusps near border ; five 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 m 2 , its surface
entirely flat except 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; no exact locality, 3 (B.M. and
U.S.N.M.).
Switzerland : Engadine, 1.
Austeia-Hungaby : Near Hatszeg, Hunyad, 3.
Remarks. — 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 specimens seen. Ursus arctos 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
THALARCTOS
297
Ursus richardsoni ; forehead in most specimens rising abruptly
above 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. horribilis 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. (b).
16. Sweden. (Lloyd.) Purchased (Stevens) . 62.3.29.7-8.
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 formicarws of Bieleb.
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) ; in form 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 Myrmaretos 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. TJialarctos Gray, Ann. of Philosophy, N.S., x, p. 62, July, 1825.
1825. Tlmlassarctos Gray, Ann. of Philosophy, N.S., x, p. 339, November,
1825.
1896. Thalassiarchus Kobelt, Bericht Senckenberg. naturforsch. Gesellsch.
Frankfurt am Main, p. 93 (Substitute for Thalarctos).
Type species. — Thalarctos polaris Gray = Ursus maritimus
Phipps.
* Compte-Bendu des Seances du Sixieme Congres Internationale de
Zoologie, Berne, 1904, p. 248. 1905.
298
CARNIVOEA
Geographical 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 upper molars not equal
to width of palate, and canines and incisors enlarged and more
prehensive in general form.
Remarks. — The genus Thalarctos, though not very strongly
differentiated from Ursus, is a well defined and perfectly natural
group.
THALARCTOS MARITIMUS PhippS.
1774. Ursus maritimus Phipps, Voyage toward North Pole, p. 185 (Spitz-
bergen).
1776. Ursus marinus Pallas, Beise duroh verschiedene Provinzen des
russisehen Beiohs, in, p. 691 (Arctic Ocean).
1792. Ursus polaris Shaw, Museum Leverianum, l, p. 7 (Benaming of
marinus).
1862. Thalarctos maritimus Gray, Gatal. Bones Mamm. Brit. Mus., p. 105.
1908. ? Thalassarctos jenaensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.
Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Jena Island,
Spitzbergen).
1908. ? Thalassarctos spitzbergensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.
Naturforsch. Preunde, Berlin, p. 184, July, 1908 (Seven Island,
Spitzbergen).
1910. Ursus (Thalassarctos) maritimus Trouessart, Paune Mamm. d'Europe,
p. 66.
Type locality. — Spitzbergen.
Geographical distribution. — 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
large ; colour uniform whitish or buffy.
External characters. — Form longer and less heavy than in
Ursus arctos, 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
points 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
arctos, 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
THALAKCTOS
299
elongate. Palate noticeably broader than in Ursus arclos, a
character made more conspicuous by the relative weakness of the
teeth. Mandible with no special peculiarities except that the
FIS. 54.
Thalarctos maritimus. X J.
lower margin is nearly straight throughout, the posterior concavity
being very slightly indicated, and lower border of angular process
only a little elevated above general outline.
Teeth. — While the general character of the dentition differs
300
CAENIVOEA
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
Fig. 55.
Thalarctos maritirmis. x \.
noteworthy. General form of upper incisors as in Ursus arctos,
but points of i l and i 2 narrower and more hooked backward ; i 3
with cusp more slender and ridges nearly obsolete ; lower incisors
THALAECTOS
301
with lobes more sharply defined. Canines both above and below
essentially similar to those of Ursus arctos, except for their greater
size. Owing to the greater width of palate the proportion of
Fig. 56.
Thalarctos maritimus. X \.
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 relatively higher
302
CAHNIVOHA
FIG. 57.
ThalaretoB maritimus. Teeth nat. size.
THALARCTOS
303
anterior cusp than in TI. arctos, its inner side more flattened,
giving the 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 m 1 not peculiar, though outer cusps are higher and narrower
and inner cusps relatively lower ; m 2 with inner cusps 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 Ursus 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 Behring Strait (mounted) :
head and body, 2670 ; tail, 90 ; hind foot, 370 ; ear, 80. For
cranial measurements see Table, p. 294.
Specimens 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^E.
1821. Canidm Gray, London Medical Eepository, xv, p. 301, April 1, 1821.
Geographical distribution. — Essentially cosmopolitan ; in E arope
west to Ireland.
Characters. — Larger cheek-teeth of a combined trenchan t 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 moderately or considerably
inflated, without septum ; form rather light, the legs long ; size
moderate ; feet digitigrade ; toes, 5-4 or 4-4.
Remarks. — Notwithstanding its wide distribution the family
Canidse 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 CANIDJE.
Interorbital region distinctly elevated ; postorbital processes
convex above ; pupil of eye round Canis, 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 Vulpes, p. 325.
Genus CANIS Linnseus.
1758. Canis Linnseus, Syst. Nat., i, 10th ed., p. 38 (type by tautonymy
G. familiaris).
1816. Lupus Oken, Lenrb. d. Naturgesch., in, pt. 2, p. 1039 (Canis lupus,
by tautonymy).
1837. Vulpicanis Blainville, Ann. Sci. Nat., Paris, 2nd ser., Zool., vni,
p. 279, November, 1837 (Canis aureus Linnseus).
1839. Lyciscus H. Smith, Jardine's Naturalists' Library, Mammals, ix,
p. 160 (Canis latrans Say).
1839. Thous H. Smith, Jardine's Naturalists' Library, Mammals, ix,
p. 193 (Canis anthus P. Cuvier).
1839. Sacalius H. Smith, Jardine's Naturalists' Library, Mammals, ix,
p. 213 (Canis aureus Linnseus).
1841. Oxygous Hodgson, Calcutta Journ. Nat. Hist., n, p. 213 (Canis
aureus Linnseus).
1857. Canis Blasius, Saugethiere Deutschlands, p. 177.
1868. Neocyon Gray, Proc. Zool. Soc, London, p. 506 (Canis latrans Say).
1869. Dieba Gray, Catal. Carn. Pachyd. and Edentate Mamm. Brit. Mus.,
p. 180 (Canis anthus P. Cuvier).
Type species. — Canis familiaris Linnseus.
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) ; interorbital region thickened
and elevated, the frontal sinuses rather 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 ; * g, e 1=5, pm tl in || = 42 ;
teeth heavy and large, the length of carnassial and upper molars
together contained about 2 J times in palatal length ; canines
robust and not specially elongated, the point of upper tooth
extending scarcely beyond middle of mandibular ramus when
jaws are closed (fig. 65).
Remarks. — 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, and the Old World jackals. Two species are known
to occur in Europe, one of which is represented by several
geographical races.
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 (length
of upper carnassial 17 to 18 mm.) ; oingulum on
outer margin of m l broad and conspicuous (South-
eastern Europe ; Jackal) C. aureus, p. 315.
Condylobasal length of skull more than 200 mm. ;
teeth larger than in the largest domestic dogs
(length of upper carnassial 25 to 27 mm.) ; cingu-
lum on outer border of m 1 narrow, tending to be
incomplete at middle (Distribution general; true
Wolves) C. lupus, p. 305.
Size rather small (exact dimensions unknown)
(Southern Spain) C. I. deitanus, p. 315.
Size large.
White of throat not extending uninterruptedly
on to cheek (Central and northern Europe) ... C. I. lupw, p. 313.
White of throat extending uninterruptedly on to
cheek (Spain except extreme south) C. I. signatus, p. 314.
CANIS LUPUS Linnaeus.
(Synonymy under subspecies.)
Geographical distribution. — Originally throughout Europe from
Ireland eastward and across Asia, now exterminated in the
British Islands, Holland and Denmark.
Diagnosis. — Condylobasal length of skull more than 200 mm.
(220 to 255 mm.) ; cheek-teeth larger than in the largest races
of 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 (fig. 61).
External characters. — General form essentially as in domestic
dogs of the " collie " type. Ear moderately long, erect, somewhat
pointed, extending about to eye when laid forward. Muzzle pad
completely bare. Fore foot with third and fourth digits sub-
equal and longest, second and fifth shorter, the large pad-like
balls fitting closely between those of third and fourth and the
slightly trilobed, heart-shaped main pad, the combined area of
balls of 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 border of main pad, its
ball scarcely indicated, no pad at its base ; wrist pad single, near
outer side, its area somewhat more than half that of ball of toes.
Hind foot essentially like fore foot, but hallux and posterior pad
absent. Claws robust, slightly curved, sub-equal throughout.
Pads and balls narrowly edged with short hair.
Colour. — General colour of upper parts, tail, and outer surface
of legs yellowish brown or buff, darker along median region of
306
CAENIVOKA
back, on posterior portion of head and outer surface of ears,
lighter and more inclined 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,
FI0. 58.
Camis lupus. X *.
especially behind middle and at base of tail ; pencil narrowly
clear black, rest of tail essentially like back. TJnderparts and
inner surface of legs pale buff or buffy white, not strongly
contrasted with sides, the chin and interramia 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 differs slightly if at all
Via. I
Canis lupus, x i.
from that of some of the larger races of domestic dogs, though
often attaining a greater size. The rostrum, however, appears
to be relatively less robust than in dog skulls of approximately
the same length. Dorsal profile rising gradually from nares to
x 2
308
CARNIVORA.
just in front of orbit, then abruptly to a little in front of bregma,
behind which it is nearly flat to strongly overhanging lambdal
region. Depth of brain-case through auditory bulla about i\
Fig. 60.
Canis lupus. X +.
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
li times that of rostrum over canines and approximately equal
CANIS 309
to distance from bregma to most posterior point of occiput.
Posterior portion of occiput strongly concave when viewed from
the side, the condyles nearly hidden beneath the projecting
lambdal region. Floor of brain-case with no specially noteworthy
features, the auditory bullre 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 moderately 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 posteriorly 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 1 and i 2 slightly more than half as
wide as high, smoothly rounded off, the cutting edge narrow but
not acute ; a small but distinct secondary lobe at each side of
front aspect, that of inner side a little below middle, that of
outer side about equally above ; posterior surface of crown con-
cave 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
CAEKIVOEA
lobes, its general form intermediate between tbat of canine and
of inner incisors. Lower incisors forming continuous row between
canines, the three teeth essentially alike in form, but increasing
Fia. 61.
Canis lupus. Teeth nat. size.
regularly in size from first 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 moderately spaced
except that pm 3 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 l subterete, that of pm 1 nearly twice as long as
broad, the height in both slightly less than length, the crown area
approximately the same as that of corresponding inner incisor, the
small cusp a little in front of middle and with slightly developed
anterior and posterior ridge. Second and third premolars two-
rooted, the crown about twice as long as wide, sub-elliptical in
outline, the inner margin sometimes (especially in pm 3 and pm 2 )
slightly concave, the long axis nearly parallel with sagittal plane
except in pm 3 , 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 i
similar to pm 3 but considerably larger, its secondary cusp
better developed and succeeded by a small but evident postero-
basal cusp springing 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
front, 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 equally trenchant commissure of main cusp at an
angle of about 75° ; cingulum complete, though low and incon-
spicuous. Lower carnassial narrower than upper but with equally
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 commissure bearing essentially the same relation to that of
protoconid as in the case of the two cusps of the upper tooth,
except that the relative lengths of the cutting edges is reversed ;
metaconid small but evident, at postero-internal base of proto-
conid ; hypoconid 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 cusps of the main triangle by a wide transverse
groove ; crown area of entoconid equal to about half that of
hypoconid, its cusp approximately 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 hypoconid 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 sharply differentiated, the
antero-posterior diameter of the outer decidedly greater than that
of inner ; paracone and metacone conical, terete, with slightly
developed anterior and posterior 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
the paracone is scarcely larger than protocone 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 1 it is usually
obsolete (compare figs. 61 and 62).
Remarks. — The material available for study has been so poor
that 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 Mr. Cabrera.
There seems to be no good reason to doubt their validity.
CANIS
313
The only known characters by which the skull of Canis lupus
can be distinguished 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 230 mm. the length of
upper and lower carnassials is respectively 21 • 6 and 25 - 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 -0(22 -8-26 -0).* In all the dog skulls which
I have examined, representing such different breeds as the pug,
fox-terrier, bloodhdund, 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 Linnaeus.
1758. [Canis] lupus Linnaaus, Syst. Nat., I, 10th ed., p. 39 (Sweden).
1792. C[anis] lupus flavus Kerr, Anim. Kingd., p. 137 (France and
Germany).
1804. Gcmis lupus niger Hermann, Observ. Zool., p. 32. Not of Kerr,
1792 (Forest of Hagenau, Alsace, Germany).
1839. ? [Canis lupus] var. canus de Selys-Longchamps, Etudes de Micro-
mamm., p. 144 (nomen nudum).
1839. 1 [Canis lupus] var. fulvus de Selys-Longchamps, Etudes do Micro-
mamm., p. 144 (nomen nudum).
1841. Lupus orientalis Wagner, Schreber's Saugthiere, Suppl., n, p. 367
(Europe).
1857. Canis lupus Blasius, Saugethiere Deutschlands, p. 180.
1863. [Canis lupus] var. major Ogerien, Hist. Nat. du Jura, in, p. 59
(Lower slopes of the Jura).
1863. [Canis lupus] var. minor Ogerien, Hist. Nat. du Jura, in, p. 69
(Higher portions of the Jura).
1897. Canis lupus minor Mojsisovics von Mojsvar, Thierleben der osterr.-
hung. Tiefebenen, p. 241 (Southern Hungary). Based on tbe
"Rohrwolf," an animal supposed to be smaller and greyer than
true lupus.
1910. Canis lupus and C. lupus lycaonf Trouessart, Faune Mamm.
d'Europe, p. 90.
Type locality. — Sweden.
Geographical distribution. — Northern and central Europe,
exact limits of range unknown ; formerly west to Ireland.
Characters. — Size maximum for the species; general colour
* Winge (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 are
255, 22 and 28. This author (p. 124) regards the domestic dogs as derived
from Canis aureus.
t Applied to the wolf of the Pyrenees; but Schreber's plate lxxxix,
the basis of the name, is a copy of Bufion'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 1 moderately large, the paracone with trans-
verse diameter of base about equal to width of large flattened
portion of crown.
Measurements. — For cranial measurements see Table, p. 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 Sassello, Liguria, 1 (Genoa).
Canis lupus signatus Cabrera.
1907. Canis lupus signatus Cabrera, Bol. Real Soc. Espan. Hist. Nat.,
Madrid, vn, p. 195.
1910. Canis lupus signatus Trouessart, Faune Mamm. d'Europe, p. 91.
Type locality. — Escorial, Madrid, Spain.
Geographical distribution. — Central Spain.
Characters. — Size and general appearance as in Canis lupus
lupus ; colour a more tawny brown than in the northern animal,
particularly on muzzle ; white of throat extending uninterruptedly
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.
Remarks. — In 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 1 conspicuously exceeds
width of the small inner portion of tooth. Mr. Cabrera informs
me that the type 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 globular form of the auditory
bullse.
<J. Seville, Spain. (A. Ruiz.) Lord Lilford (p). 95. 3. 3. 6.
<S, ?. Riocabado, Burgos. Hon. N. C. Roths- 11. 10. 5. 1-2.
(Bev. S. Gonzales.) child (p).
* In the Burgos specimens the colour is not unusually tawny : back
and sides a coarse mixture of black, whitish, ochraceous-buff , and drab grey
(underfur), the black and whitish most conspicuous along back, the
ochraceous-buff on 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.
315
Canis lupus deitanus Cabrera.
1907. Canis lupus deitanus Cabrera, Bol. Real Soc. Espan. Hist. Nat.,
Madrid, vn, p. 197.
1910. Canis lupus deitanus Trouessart, Faune Mamm. d'Europe, p. 91.
Type locality. — Moratalla, Murcia, Spain.
Geographical distribution. — Now known from the type locality
only.
Diagnosis. — Smaller than Canis lupus lupus and brighter in
colour, the general appearance much as in C. aureus.
Measurements. — Unknown .
Specimens examined. — I have seen the two living examples in Madrid
on which the form was based.
Remarks. — Tn general appearance the two Moratalla wolves
are strikingly different from Can in lupus. Unfortunately their
true characters are not yet known.
CANIS AUREUS Linnseus.
1758. [Cams'] aureus Linnseus, Syst. Nat., i, 10th ed., p. 40.
1835. Canis aureus var. moreotica I. Geoffroy, Exped. Sci. de Mor6e. Zool.,
pi. i (Morea, Greece).
1841. C[anis] dalmatinus Wagner, Sohreber's Siiugthiere, SuppL, n, p. 383
(Dalmatia).
1841. Clams'] griecus Wagner, Sohreber's Siiugthiere, Suppl., n, p. 383
(Peloponesus).
1892. Canis aureus balcanicus Brusina, Glasnik Hrvatskoga Naravoslovnoga
Drustva, Zagreb, vn, p. 317 (Drava River, Croatia).
Type locality. — Province of Lar, Persia.
Geographical distribution. — India and westward through Asia
Minor to the Balkan Peninsula, north to Heves Comitat,
Hungary.
Diagnosis. — Smaller than Canis lupus (condylobasal length of
skull less than 200 mm.) ; teeth not equal to those of the larger
domestic dogs in size, the upper molars with wide, conspicuous
cingulum on outer side (fig. 62).
Colour. — 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
ai'ound and behind base of ear ; muzzle more heavily washed
with black y 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 back, the basal half above greyish, the
terminal half and underside buffy much overlaid with black ;
pencil blackish ; underparts and inner side of legs dull buffy
grey ; chin blackish ; feet dull buffy.
316
CAKNIVOEA
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318
CAENIVORA
Skull. — The skull is much smaller than that of Ganis lupus.
In form it differs slightly in the less elevated frontal region and
somewhat more inflated auditory bullae.
Teeth. — While agreeing with those of Ganis 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 Ganis lupus, their
cutting ridges much more developed ; transverse
diameter of metacone in m 1 noticeably less than
width of inner portion of crown ; cingulum on
outer border of both upper molars wide and
conspicuous in contrast with narrow cusps,
Fig. 62. showing no tendency to become obsolete in
Canis aureus, region between paracone and metacone. Lower
La clee ™teeth ary carna ssial 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 cusps strongly developed.
Measurements. — For cranial measurements see Table, p. 316.
Specimens examined* — One from Greece (Piraeus) ; numerous others
from Asia Minor and India.
Remarks. — 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. Piraeus, Greece. (0. Mottaz.) Hon. N. 0. Roths- 8. 10. 2. 49-50.
child (p).
Genus ALOPEX Kaup.
1829. Alopex Kaup, Entw.-Gesch. und Natiirl. Syst. Europ. Thierw., i,
p. 83.
1857. Leucocyon Gray, Proc. Zool. Soc, London, p. 512.
Type species. — Ganis lagopus Linnaeus.
Geographical distribution. — Arctic region of both 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 bead-like,
overhanging edges ; dorsal profile of forehead rising abruptly
ALOPEX 319
above rostrum as in Canis ; teeth moderately heavy and large,
the length of carnassial and upper molars together contained
about 2£ times in palatal length ; canines and incisors inter-
mediate between those of Canis and Vulpes (see fig. 65) ; external
form fox-like, but ear short and rounded, not conspicuously
overtopping the surrounding fur.
Remarks. — 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. spitzbergenensis, p. 324.
ALOPEX LAGOPUS Linnaeus.
1758. [Canis] lagopus Linnaeus, Syst. Nat., i, 10th ed., p. 40 (Lapland).
1816. V[ulpes] arctica Oken, Lehrb. d. Naturgeseh., in, pt. 2, p. 1033
(Renaming of Canis lagopus).
1820. C[anis] vulpes cosrulea Nilsson, Skand. Fauna, I, p. 88 (Lapland).
182Y. [Canis lagopus] & argentms Billberg, Synopsis Faunae Scandinaviae,
p. 14 (Lapland).
1910. Vulpes lagopus Trouessart, Faune Mamm. d'Europe, p. 96.
Type locality. — Lapland.
Geographical distribution. — 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.
Diagnosis. — General characters as in the genus ; condylobasal
length of skull about 130 mm. in males, about 124 mm. in
females.
External characters. — In general external characters Alopex
lagopus resembles Vulpes vulpes, though the muzzle is less
elongated, and the low, rounded ears (not extending to eye when
laid forward) impart a somewhat un-fox-like appearance to the
head. Fur very dense, the underfur in summer about 12 mm.
deep 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
abundant underfur. Feet as in Canis, but claws longer and
more slender, and entire palm and sole covered with a dense'
woolly growth of hair, 15-17 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 difference between the two animals.
320
CAENIVOEA
summer when it sometimes wears away sufficiently to expose
balls 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. 63.
Alopex lagopus. X $.
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 interramial region
tinged with slaty grey ; each of the longer hairs of back with
321
one cream-buff sub-terminal annulation, producing a noticeably
speckled appearance throughout dark area ; flanks with a few
long, entirely buff 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 buffy
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 ; underf ur 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 Vulpes
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. vulpes) ; 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 3
and pn& ; 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 Vulpes
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 1 and i 2 much more developed and forming a
T
no. 64.
Jlopex lagopus. Teeth. Nat. size.
322
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noticeable inner and outer secondary cusp (the former soon
wearing away in i 1 ) 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. Schaanning(c). 11.6.3.12.
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 skulls. Lapland. Wheelwright (c). 64. 3. 8. 3-4.
ALOPEX SPITZBERGENENSIS Barrett-Hamilton and Bonhote.
1799. ? Canis fuliginosus Beohstein, Thomas Pennant's allgem. Uebersicht
d. vierfuss. Thiere, I, p. 270 (Iceland).
1799. ? Canis groenlandicus Bechstein, Thomas Pennant's allgem. Ueber-
sicht d. vierfuss. Thiere, i, p. 270 (Greenland).
1898. Canis lagopus spitzbergenensis Barrett-Hamilton and Bonhote, Ann.
and Mag. Nat. Hist., 7th ser., I, p. 287, April, 1898 (Spitzbergen).
Type in British Museum.
1910. Vulpes lagopus spitzbergensis and ? V. lagopus fuliginosus Trouessart,
Paune Mamm. d'Europe, p. 97.
Type locality. — Spitzbergen.
Geographical distribution. — Spitzbergen ; also Iceland and
Greenland ?
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-buff 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.) ; lips 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.): —
i skull. Spitzbergen. Stockholm Museum (e). 90. 8. 1. 2.
6, 9. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 2-3.
(96. 9. 23. 3. Type of species.)
S, ¥ skulls. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 4-5.
Genus VULPES Oken.
1816. V[ulpes] Oken, Lehrb. d. Naturgesch., in, pt. 2, p. 1033, in full on
p. 1034 and in index, p. 1268 (Vulpes communis Oken = Canis
vulpes Linnseus).
1821. Vulpes Bowdich, Anal. Nat. Olassif. Mamm., p. 40 (Canis vulpes).
1857. Vulpes Blasius, Saugethiere Deutschlands, p. 178 (Sub-genus of
Canis).
Type species. — Canis vulpes Linnseus.
Geographical distribution. — Northern portion of the northern
hemisphere from about the limit 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 processes
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 Canis, 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).
Remarks. — As thus restricted the genus Vulpes contains about
thirty-five forms, all peculiar 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) V. ichnusse, p. 336.
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 V. vulpes, 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. wipes, p. 330.
Teeth smaller and less robust, the premolars tend-
ing to be widely spaced, their secondary cusps
usually obsolete or absent.
Yellowish and reddish tints bright ; posterior
half of back seldom much frosted with whitish ;
tail never clear grey (Central Europe) V. v. crurigera, 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. silacea, p. 333.
326
CARNIVOEA
FIG. 65.
Incisors and canines from front, of Cards (a), Alopex (6), and Vulpes (c).
VULPES VULPES Linnaeus.
(Synonymy under subspecies.)
Type locality. — TTpsala, Sweden.
Geographical distribution. — Europe from the Arctic coast to
the Mediterranean, and from Ireland eastward into Asia.
Diagnosis. — 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,
v0lpes 327
thick and bushy, with abundant underfur ; longer hairs of back
normally concealing the underfur ; feet as in Ganis, but with
the soles hairy 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 middle 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 divides, 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 bullae 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 somewhat concave, the under surface so
abruptly concave that the process is much less thick than in
Alopex and Ganis. Rostrum moderately long (the distance from
orbit to gnathion about equal 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
CARNIVORA
foramen over posterior root of pm 3 or over space between pm 3
and pm i . Palate relatively narrower than in the European
Fig. 66.
Vulpes vulpes. X h
species of Alopex and Canis, its median posterior termination
about at level of middle of last molar ; mesopterygoid fossa
329
about twice as long as wide, the sides gradually converging
posteriorly ; hamulars 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
S 9
cusps. Upper incisors slender,
not closely crowded, their
crowns relatively narrower than
in either Canis lupus or Alopex
lagopus, 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 3 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£ 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 he in or nearly in the main axis of the tooth. Lower carnassial
with posterior heel essentially as broad as main portion of tooth,
the cusps essentially as in Canis, but general aspect of tooth more
trenchant. Upper molars differing from those of Canis in the
relatively smaller size of paracone and metacone, these cusps
appearing 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.
Nat. size.
330
CABNIVOEA
Remarks. — 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 vulpes vdlpes Linnseus.
1758. [Canis] vulpes Linnseus, Syst. Nat., i, 10th ed., p. 40 (Sweden).
1758. [Canis] alopex Linnseus, Syst. Nat., I, 10th ed., p. 40 (Sweden).
1798. C[anis] vulpus Thunberg, Beskrifning pa Svenske Djur, Mamm., p. 7
(Variant of vulpes).
1816. V[ulpes] vulgaris Oken, Lehrb. d. Naturgesoh., in, pt. 2, p. 1034
(Renaming of vulpes).
1820. Canis nigro-argenteus Nilsson, Skand. Fauna, i, p. 91 (Lofoten
Islands, Norway).
1827. [Canis vulpus] y nigrocaudatus Billberg, Synopsis Faunse Scandi-
navise, p. 12 (Uppland, Sweden).
1827. [Canis vulpus] £ variegatus Billberg, Synopsis Faunse Soandinavise,
p. 13 (Uppland, Sweden).
1827. [Canis vulpus] 17 Uneatus Billberg, Synopsis Faunse Soandinavise,
p. 13 (Skane, Sweden).
1830. ? [Vulpes] communis Burnett, Quart. Journ. Sci. Lit. Art, xxviii,
1829, p. 349 (Substitute for vulpes). Nomen nudum.
1898. Vulpes vulpes Thomas, The Zoologist, 4th ser., n, p. 100, March,
1898 (part).
1910. Vulpes vulpes Trouessart, Faune Mamm. 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 European
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). 64. 3. 8. 2.
11 skulls. Egersund, Stavanger, K. H. Schaanning (c). 11. 6. 3. 1-11.
Norway.
VULPES 331
Vulpes vulpbs crucigera Bechstein.
1789. [Cams] crucigera Bechstein, Gemeinn. Naturgesoh. Deutsohlands,
I, p. 250 (Thiiringen, Germany).
1792. C[anis] Vulpes alopex europaeus Kerr, Anim. Kingd., p. 142 (Bur-
gundy, France).
1797. Cams vulpes alba Borkhausen, Deutsche Fauna, I, p. 33 (Vogelsberg,
near Budigshain, Hessen, Germany).
1797. Cams vulpes nigra Borkhausen, Deutsche Fauna, I, p. 33 (Hessen
and Thiiringen, Germany).
1801. Clams'] v[ulpes] lutea Bechstein, Gemeinn. Naturgesch. Deutsoh-
lands, i, 2nd ed., p. 628 (Thiiringen, Germany).
1801. C[anis] v[ulpes] cinerea Bechstein, Gemeinn. Naturgesch. Deutsoh-
lands, I, 2nd ed., p. 628 (Thiiringen, Germany).
1832. Canis melanogaster Bonaparte, Ioonogr. Fauna Ital., i, faso. 1
(Neighbourhood of Borne, Italy).
1841. Vulpes hypomelas Wagner, Schreber's Saugthiere, Suppl., n, p. 405
(Oberbayern, Germany).
1857. Cams vulpes Blasius, Saugethiere Deutsohlands, p. 191.
1861. ? V[ulpes] vulgaris meridionatis Fitzinger, Wissensch.-pop. Natur-
gesch. der Saugeth., I, p. 194 (Dalmatia).
1910. Vulpes vulpes (part) and V. vulpes melanogaster Trouessart, Faune
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 rather 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 instances 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.
Skull and teeth. — The skull and teeth do not attain so great
332
CARNIVOKA
size as in true Vulpes vulpes, 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, 160 ; 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 : Ax-les-Thermes, Ariege, 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.M.) ; Grossgraben, Silesia, 1 (Breslau) ; Biesengiberge, Silesia, 1
(Breslau) ; southern Germany, 3.
Austria-Hungary : Trentino, 1 (Genoa) ; Gazza, Trentino, 1 (Genoa) ;
Vigolo Vattaro, Trentino, 1 (Genoa).
Switzerland : Geneva, 1 (Ghidini) ; Valais, 1 (Ghidini).
Italy : Porlezza, Como, i (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.
Remarks. — 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 meridionalis Fitzinger.
9. Ben Nie, Sutherlandshire, E. B. Alston (p). 79. 9. 25. 80.
Scotland. (H. Brown.)
6. Inversanda, Ardgour. H. Leigh (c & p). 1. 16. 12. 1.
2. Kilmanock, Wexford, G. Barrett-Hamilton 6. 5. 20. 1.
Ireland. (c & p). 9. 12. 15. i.
?. 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).
volpbs 333
2 6. Thame, Oxfordshire. Hon. N. C. Roths- 0. 10. 31. 1-2.
child (p).
2i5. Ditchling, Sussex. Guy Do 11m an (9. 11. 3. 1.
child (p).
Dollman (9. 11.
(c & p). \10. 9. 13. 1.
3. Tunbridge Wells, Sussex. C. H. B. Grant (c). 1. 2. 15. 1-3.
2 6,9. Ax-les-Thermes, Ariege, V. Builles (a & p). 8.3.27.6-7.
Prance. 8. 3. 27. 14.
9. Caterille, Haute-Garonne, 0. Thomas (p). 8. 7. 15. 2.
900 m. (A. Robert.)
9. Pic Sessire, Haute-Garonne. O. Thomas (p). 8. 7. 15. 3.
(A. Robert.)
6. St. Aventin, Haute-Ga- 0. Thomas (p). 8. 7. 15. 4.
ronne, 900 m. (A. Robert.)
6. 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.
6. Genoa, Liguria, Italy. Marquis G. Doria(p). 88. 12. 1. 3.
skull. Pisa. Zool. Soc. Mus. 58. 5. 4. 126.
1 yg. S. Italy. (Prof. Savi.) Zool. Soc. Mus. 55. 12. 24. 240.
9. Tuscany. Purchased (Dr. 45.7.22.15.
Riippell).
6, 9. Tatoi, Athens, Greece. Hon. N. 0. Roths- 8. 10. 2. 22-23.
(C. Mottaz.) child (p).
Vulpbs VULPES silacea Miller.
1907. Vulpes vulpes 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'Burope, p. 45.
Type locality. — Near Silos, Province of Burgos, 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 ; Coto Donana, Huelva, 3 (B.M. and U.S.N.M.)
Elche, Alioante, 1 ; Barracas, Castellon, 1.
Remarks. — 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.)
6, 2 9, 1. Burgos. Rev. Saturio Gon- 8. 7. 7. 12-15.
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CARNIVORA
9. Arrechavaleta, Vitoria. 0. Thomas (p). 8. 2. 9. 49.
(N. Gonzalez.)
S. Torres del Alio, Corufia. Dr. V. L. Seoane 94. 5. 29. 1.
(o & p).
5. Madrid. M. de la Esoalera (c). 8. 9. 24. 7.
<$, ?. Seville. (A. Buiz.) Lord Lilford (p). 95. 9. 4. 7-8.
2 c5. Coto Donana, Huelva. A. Chapman (c & p). 8. 3. 8. 4-5.
6. Elehe, Alicante. G. S. Miller (c). 8. 8. 4. 48.
i juv. Barraoas, Castellon. 0. Thomas (p). 8. 2. 9. 50.
(N. Gonzalez.)
VULPES ICHNUS.E Miller.
1907. Vulpes ichnusee Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 391,
November, 1907. Type in British Museum.
1910. Vulpes vulpes ichnusse Trouessart, Paune Mamm. d'Burope, p. 94.
Type locality. — Sarrabus, Sardinia.
Geographical distribution. — Sardinia and Corsica.
Diagnosis. — Smaller than any of the races of Vulpes 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 lighter and
more dull on base of ears and on neck, and fading to ochraceous-
ruf ous on shoulders and back ; sides of neck, outer surface of
upper arm and region just behind axilla still lighter, a tawny
buff, 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 buffy white subterminal areas about
5 mm. in length, the extreme tips reddish ; feet and legs
ochraceous-rufous, slightly clouded with blackish and a little
speckled with buffy white ; tail like back above, the tawny
gradually fading out through a buffy grey to the whitish buff
tip, the longer hairs except in pencil black tipped (30-40 mm.) ;
underparts to front legs buffy whitish 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 j 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, 1 27 ; ear, 74.
Adult from near St. Florent, Corsica; hind foot, 123 ; ear from
meatus, 73.
Specimens examined. — Eight, from the following localities : —
Corsica : Grotto Campu Cousule, near St. Florent, N.W. Corsica, 1
(Major).
Sardinia : No exact locality, 1 (Turin) ; Lamisei, 1 ; Sarrabus, 3 (B.M.
and Genoa) ; Zinnigas Siliqua, 2.
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Corsica : near St. Plorent
Germany : Southern Germany
Ingelheim, Rheinh
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Italy : near Genoa .
Pisa
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340
CARNIVORA
Bemarks. — This small, short-eared fox is probably a native
on the two islands where it occurs. The presence of the larger
animal, similar to that of Italy, on Sardinia may be due to recent
introduction. The Corsican specimen, though a mummy and
without fur, appears to be a typical example of Vulpes ichnusse.
6. Sarrabus, Sardinia. Marquis G. Doria (p). 88. 12. 1. 2.
(Type of species.)
6. Lanusei, Sardinia. 0. Thomas (p). 0. 2. 21. 1.
(W. Wolterstorff.)
Family MUSTELIDvE.
835. Mustelidss Swainson, Nat. Hist, and Classif. of Quadrupeds, p. vn, 361.
Geographical distribution. — Essentially cosmopolitan ; absent
from Madagascar and Australia ; in Europe 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 ; upper molars, 1-1 ; upper
carnassial with not more than two outer cusps ; auditory bulla flat
or moderately inflated, without septum ; form usually slender,
the legs always short ; size moderate or small (including the
smallest known carnivores) ; feet digitigrade or sub- plantigrade ;
toes. 5-5.
Bemarks. — The family Mustelidse is, next to the Ganidte, the
most generally distributed group of carnivores. It is divisible
into four sub-families, all of which occur in Europe, where they
are represented by six of the two dozen or more known genera.
KEY TO THE EUROPEAN GENERA OP MTJSTELIDM.
Grown of upper carnassial triangular or rhombic in
outline, its length and width sub-equal ; lower car-
nassial with anterior triangle distinct, the mefca-
conid nearly as large as the outer cusps.
Upper molar much larger than carnassial ; skull
narrow and high (normal), the rostrum much
longer than broad; tail short, bushy, not mus-
cular ; habits fossorial (Badgers, sub-family
Meles, p. 341.
Upper molar about equal to carnassial ; skull broad
and flat, the rostrum broader than long ; tail long,
densely furred, very muscular ; habits aquatic
(Otters, sub-family Lutrime) Lutra, p. 354.
Crown of upper carnassial not triangular or rhombic in
outline, its length much greater than its width ;
lower carnassial with anterior triangle obsolete or
absent (represented by the two outer cusps only),
the metaconid when present much smaller than the
other cusps.
341
Premolars t-..
4-4
Lower carnassial without metaconid ; skull robust
(in European species considerably more than
100 mm. in condylobasal length) ; form robust ;
tail short (Wolverenes, sub-family Ouloninm)... Oulo, p. 433.
Lower carnassial with evident though small meta-
conid ; skull slender (in European species con-
siderably less than 100 mm. in condylobasal
length) ; form slender ; tail long (Martens, sub-
family Mustelinie) Maries, p. 365.
Premolars ||.
Lower carnassial with evident though small meta-
conid ; hamular in contact with bulla ; back and
sides spotted and striped (Tiger Polecats, sub-
family Mustelinae) Vormela, p. 428.
Lower carnassial without metaconid ; tip of hamu-
lar widely separated from bulla ; back and sides
never spotted, rarely (in certain Asiatic species)
with median dorsal stripe (Weasels, Polecats,
&c, sub-family Mustelime) Mustela, p. 381.
Sub-Family MELINiE.
1857. Melinss Baird, Mamm. North Amer., p. 148.
Geographical distribution. — Temperate and tropical portions of
both hemispheres ; in Europe west to Ireland and north to
central Scandinavia.
Characters. — Upper carnassial with evident crushing surface,
its 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 rather high and long, the rostrum
longer than broad ; external form short and heavy, the fur long
and loose ; toes not webbed, the claws large, fossorial ; tail
variable in length (short in European members of the group),
never unusually muscular.
Bemarks. — About a dozen genera, or nearly one-half of the
family, are now placed in the sub-family Melinse. Only one
occurs in Europe.
Genus MELES Brisson.
1762. Meles Brisson, Kegn. Anim. in Classis ix, distrib., 2nd ed., p. 13
(Meles Brisson = Ursus meles Linneeus).
1780. Meles Storr, Prodr. Meth. Mamm., p. 34. First use of name by an
author following the Linnsean system ( Ursus meles) .
1795. Taxus Geoffroy and Cuvier, Magasin Encyclopedique, II, p. 184
(Ursus meles Linnaeus).
1815. Melesium Rafmesque, Analyse de la Nature, p. 59 (Modification of
1857. Meles Blasius, Saugethiere Deutschlands, p. 202.
Type species. — Meles * Brisson = Ursus meles Linnseus.
* Not "Metes meles," the form in which this and other Brissonian
monomial specific names are often cited, apparently with the intention of
palliating the absurdity of recognising in nomenclature the terms applied
to genera by an author who did not follow the Linnsean system.
342
CARNIVORA
Geographical 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: ijgj, eg, pm^* mt? = 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 anterior cusp with its slightly developed
anterior commissure and more distinct posterior commissure, the
crushing portion represented by the obliquely sloping 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.
Memarhs. — 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 POEMS OP MELES.
Maxillary tooth-row, exclusive of incisors, about
85 mm. ; auditory bullee rather strongly in-
flated, the inner border 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. meles, p. 343.
General colour moderately pale ; 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. in. marianensis, p. 352.
* In adults usually — ,, owing to the early disappearance of the small
pm l .
343
meles MELES Linnaeus.
(Synonymy under subspecies.)
Type locality. — Upsala, Sweden.
Geographical distribution. — Europe, west to Ireland, south to
the Mediterranean and north to central Scandinavia. Eastern
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-defined 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.
Mamma; : a 2-2, i 1-1 = 6.
Colour. — Back and sides a coarse grizzle of black and bufly
■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 insensibly into colour of back ; width of white
median area on face usually greater than that of the dark lateral
stripe, and about equal 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 band ; tail like back at base, soon fading to soiled white.
Skull. — Except for the greatly developed sagittal crest, the
344
CAENIVOKA
height of which in old individuals is equal to nearly half inter-
orbital breadth, the skull of Meles nieles 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. 88.
Meles meles. X 4.
developed it reduces this angle to 20° or less. It is slightly
convex anteriorly, rather abruptly convex posteriorly, the hinder-
most portion slightly overhanging. Postorbital processes 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 widely spreading posteriorly, narrow
MELES
345
anteriorly, the widest portion opposite front of glenoid fossa, the
arch very slightly bowed upward. Anteorbital foramina rather
large, sub-circular in outline, their greatest diameter about half
that of orbit. Ventral profile 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 i 3 , 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 hamular. Mesopterygoid
x h
space short and wide, its length only a little more than distance
between tips of hamulars. Auditory bull* 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 bullae about 1£ times diameter of
inflated portion ; paroccipital process short, triangular (not ridge-
like), its extreme base applied to posterior border of bulla.
346
CAENIVOKA
Mandible heavily built, the ramus nearly straight, the lower
border with the usual upward curve posteriorly, its depth at
middle about one-third that through coronoid process ; angular
fig. 70.
Meles males. X f.
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 Mustelidse. 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 1 and P 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 ; i 3 with crown area about double that of i 2 and nearly half
347
that of canine, its crown higher than in the other incisors,
owing to lower insertion of root, bujt 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
canines, the shafts straight, sloping obliquely forward, the root
of « 2 implanted behind the others ; cutting edge irregularly
2-lobed, that of i 3 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 1 early
deciduous, pm^ usually more
persistent. Second premolars
similar to each other, the crown
area about equal to that of i 2 ,
the outline irregularly oval when
viewed from above, the cusp
about as high as long, its apex
slightly in front of middle of
crown ; root of each tooth single,
that of pm 2 showing a tendency
to become divided longitudi-
nally. Other small premolars
(pm 3 , pm 3 and pm^j alike in form,
the crowns laterally compressed,
triangular when viewed from
the side, with apex slightly in
front of middle, pin 3 and pm 3
nearly alike in size, pm i some-
what longer ; cingulum very slightly developed, not forming
secondary cusps. Upper carnassial triangular in outline, the
postero-internal border longest ; outer, trenchant portion of teeth
consisting of a main anterior cusp resembling pm i 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 postero-internal border. Lower carnassial with
anterior triangle well developed, the three cusps sub-equal in
height, the metacouid subterete, the protoconid more compressed
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
no. 71.
Meles meles. Teeth. Nat. size.
348
CAKNIVORA
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 metacooid. 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 It 1 , times as long as latter; outer border occupied by two
rather large, subterete, conical cusps, the anterior (paracone)
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.
JRemarks. — 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. [Ursus] meles Linnseus, Syst. Nat., i, 10th ed., p. 48 (Sweden).
1785. [Meles] taxus Boddaert, Elenchus Animalium, i, p. 80 (Europe).
1789. [Ursus meles} £ 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
1816. Meles europmus Desmarest, Nouv. Diet. d'Hist. Nat., in, p. 465
(Eenaming of XIrsus meles).
1822. Taxus meles F. Cuvier, Hist. Nat. des Mamm., in, faso. 36, January,
1822.
1827. [Meles] communis Billberg, Synopsis Faunae Soandinaviae, p. 16
(Renaming of Ursus meles).
1827. [Meles communis] caninus Billberg, Synopsis Faunae Scandinavia,
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, Mitteilungen des Kaukasisohen
Museums, n, p. 115 (Based on the cranial measurements of
English specimens recorded by Barrett-Hamilton, Ann. and Mag.
Nat. Hist., 7th ser., 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. — Central and southern Europe from
southern Sweden to the Pyrenees and Italy, and from Ireland
eastward ; eastern limit of range not known.
Characters. — Skull and teeth moderately large, the teeth
rarely attaining maximum size for the species ; colour moderately
light, the sides not often conspicuously whitish.
Measurements. — Adult male and female from Woodpark,
Galway, Ireland: head and body, 686 and 618; tail, 153 and
150 ; hind foot, 108 and 102 ; ear from meatus, 48 and 45.
Adult female from Boueonne, 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 examined. — Thirty-five, from the following localities : —
Ibeland : Woodpark, Galway, 2.
England: Burnley, Lancashire, 1; Ross, Hereford, 1; Kentchurch,
Herefordshire, 4 ; Tetworth, Oxfordshire, 1 ; Ibstone, Buckinghamshire, 1 ;
Horsham, Sussex, 1 ; Bicton, Devonshire, 1 ; Oornwall,l; no exact locality, 4.
Norway : Holme, Mandal, 7 ; Egersund, Stavanger, 2.
Sweden : Middle Sweden, no exact locality, 1 (Stockholm) ; Takern,
Ostergotland, 1 (Stockholm) ; Smaland, 1 (Stockholm) ; Skane, 2 (Stock-
holm) ; no exact locality, 1.
Ekance : Eor£t de Boueonne, Gers, 1.
Gebmany : Ingelheim, Bheinhessen, 1 ; Strass, near Burgheim,
Bavaria, 1.
Italy : Liguria, 1 (Genoa).
<5, ?. Woodpark, Galway, Ire- R. E. Hibbert (p). 95. 6. 2. 1.
land. 95. 12. 3. 1.
(Co-types of M. m. britannicus, Satunin.)
?. Burnley, Lancashire, Hon. N. C. Roths- 0. 10. 31. 4.
England. child (p).
i. Ross, Herefordshire. Hon. N. C. Roths- 0. 10. 31. 3.
child (p).
9, 3 juv. st. Kentchurch, Hereford- W. E. de Winton
shire. (c & p).
6. Tetworth, Oxfordshire. Hon. • H. Parker
(c & p).
i. Ibstone, Buckingham- A. H. Cocks (c & p). 4. 1. 25. 1.
shire.
A. Horsham, Sussex. E.C. Hawes (c & p). 7.4.13.1.
9. Bicton, Devonshire. J.C.Stagdon(c&p). 97.2.23.1.
1 (albino) Cornwall. Lord Lilford (p). 89. 10. 28. 1.
skull. Sweden. Purchased (Wheel- 64. 3. 8. 1.
wright).
4 (J, 2 9. Holme, Mandal, Norway. R. J. Cuninghame 8. 8. 9. 7-12.
(c & p).
2 skulls. Egersund, Stavanger. K. H. Schaanning /ll. 6. 3. 13.
(o), ill. 10. 23. 2.
350
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9. Foret de Bouconne, Gers, 0. Thomas (p). 8.7.15.5.
250 m. Prance. (A.
Robert.)
9. Ingelheim, Rheinhessen, C. Hilgert (o). 8. 11. 2. 16.
Germany.
Meles meles marianensis Graells.
1897. [Meles taxus] var. marianensis Graells, Mem. Real Acad. Cien.,
Madrid, xvn, p. 170 (Central Spain).
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., iv, p. 384, November, 1899 (Seville, Spain). Type
in British Museum.
1910. Meles meles mediterraneus Trouessart, Faune Mamm. d'Europe, p. 70.
Type 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 ; Coto Dofiana, Huelva, 2.
Remarks. — 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. Bev. Saturio Gon- 8. 7. 7. 19-20.
6,9. Seville. (A. Ruts.) Lord Lilford (p). 95.3.3.7-8.1
(95. 3. 3. 7. Type of mediterraneus Barrett-Hamilton.)
<S, 9. Coto Donana, Huelva. A. Chapman (p). 8. 3. 8. 6-7.
MELES ARCALUS Miller.
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., iv, p. 131, November, 1899 (part).
1906. Meles meles mediterraneus Bate, Proc. Zool. Soc, London, 1905, ir,
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.
Type 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 OP MELES MELES AND M. ABC ALUS.
Looality.
Number.
Sex.
Tr , ' Lower molar
Upper molar. , (flrst)
M. meles meles.
i
Sweden : no exact locality
64. 3. 8. 1
9
13-0 X 11-4
15-4 x 7-0
Wegelin
Stockholm
£?
15-0 x 130
16-2 x 7-8
Skane
,,
6 1
15-2 x 13-0
18-0 X 8-2
Ulriksdal .
,,
<5juv.
13-4 x 11-6
15-8 x 7-8
Smaland .
,,
9?
14-2 x 11-8
16-2 x 7-4
Orebro . . . . !
,
5 juv.
14-2 x 11-8
15-8 X 7-0
Takern ....
„
6 ?
13-6 X 11-2
15-6 X 7-2
Norway : Holme, Mandal
152622
6
11-6 x 10-4
14-0 X 6-4
•
8. 8. 9. 9
i
12-8 x 11-8
14-6 x 7-0
» „
8. 8. 9. 10
6
12-6 x 11-2
14-6 X 7-0
„ „
8. 8. 9. 11
9
11-2 X 10-8
14-0 x 6-8
)i
8. 8. 9. 12
9
12-2 x 10-4
14-0 x 7-0
Egersund, Stavanger
11. 6. 3. 13
15-0 X 11-4
14-8 x 7-0
England : Boss, Hereford
0. 10. 31. 3
i
13-2 x 11-4
16-0 x 8-0
Cornwall
98. 10. 28. 1
9 ?
14-0 x 12-0
16-6 x 7-8
no exact locality
2116
SI
13-4 x 12-0
16-4 x 8-0
,>
211c
9?
14-0 X 12-4
16-2 x 8-2
» ..
—
14-2 x 12-2
—
—
13-6 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 : Woodparb, 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*
S
15-0 X 12-6
18-5 x 8
2
„
6
15-0 X 13-7
18-3 x 8
3
S
15-5 x 13-6
18-0 x 9
9
14-6 X 125
17-5 x 7-7
various localities, (Maxi-1
13 specimens : \ mum J
16-3 X 13-6
fMini-1
\mumj
13-0 X 12-0
—
various localities, f Maxi-1
17 specimens : \ mum |
18-7 x 8-5
"
jMini-1
\ mum /
150 x 7-0
France: Bouconne, Gers.
8. 7. 15. 5
9
14-0 X 120
16-2 X 7-2
Germany : Ingelheim, Rheinhessen .
8. 11. 2. 16
9
14-8 x 12-8
1^-2 X 7"8
Italy: Borghetto S. Spirito, Liguria
33 Genoa
9
13-6 X 110
15-6 X 72
M. meles marianensis.
Spain : Quintanar, Burgos
8. 7. 7. 19
S
15-2 X 12-6
17-4 x 8-6
Seville
95. 3. 3. 7t
i
15-0 X 12-0
16-6 x 7-8
95. 3. 3. 8
9
14-6 X 12-6
16-8 x 8-2
Coto Dofiana, Huelva .
8. 3. 8.
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
M. arcalus.
Crete ....
5. 12. 2. 17t
?juv
! 13-6 x 11-2
15-4 X 7-2
>>......
5. 12. 2. 38
ad.
13-8 X 11-6
* Measured by H. Winge.
+ Type of mediUiTanevx Barrett-Hamilton.
■2
t Type.
354
CAENIVOEA
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 1 relatively larger
than M. meles, so that breadth of crown through this cusp is as
great as or greater than that through paracone ; colour as in
Meles meles marianensis or slightly paler.
Measurements. — For cranial and dental measurements see
Tables, pp. 351, 353.
Specimens examined. — Four, all from Crete.
(5,9. 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.
9. Crete. H. 0. Jones, R.N. (c). 99. 6. 13. 1.
Sub-Family LUTEINS.
1857. Lutrinse 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.
'Remarks. — 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. Lutra Brisson, Regnum Animale in Classis rx distrib., 2nd ed., p. 13
(Lutra Brisson = Mustela lutra Linnaeus).
1780. Lutra Briinnich, Zoologiae Fundamenta, p. 34. First use of name
by an author following the Linnaean system (Mustela lutra).
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. Mus., p. 70 (L. mollis Gray
= Lutra lataxina F. Cuvier).
1857. Lutra Blasius, Saugethiere Deutschlands, p. 236.
1865. Barangia Gray, Proc. Zool. Soc. London, p. 123 (B. sumatrana Gray
= Lutra barang F. Cuvier).
LDTRA 355
1865. Lutrogale Gray, Proc. Zool. Soc. London, p. 127 (Lutra monticola
Hodgson).
1865. Nutria Gray, Proo. Zool. Soo. London, p. 128 (Lutra felina Molina).
1867. Lutronectes Gray, Proo. Zool. Soc. London, p. 180 (L. whiteleyi
Gray = L. lutra Linnseus ?).
Type species. — Lutra * Brisson = Mustela lutra Linnaeus.
Geographical distribution. — 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 g, 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 transverse 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.
Bemarks. — Although more widely distributed than any other
living genus of land mammals, Eptesicus and Myotis 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 Linnaeus.
1758. [Mustela] lutra Linnseus, Syst. Nat., i, 10th ed., p. 45 (Sweden).
1777. [Lutra'] vulgaris Erxleben, Syst. Regni Anim.,i, p. 448 (Renaming of
lutra).
1792. M[ustela] Lutra giscatoria Kerr, Anim. Kingd., p. 172 (Renaming of
lutra). \i
Not " Lutra lutra" (see footnote under Meles, p. 341).
2 A 2
356
OARNIVOEA
1816. ? Lutra fluviatilis Leach, Syst. Gatal. Spec. Indig. Mamm. and
Birds Brit. Mus., p. 6 (nomen nudum : " Kiver Otter ").
1827. [Lutra vulgaris] j8 marinus Billberg, Synopsis Faunae ScandinaviEe,
p. 28 (Coasts of Scandinavia).
1830. ? [Lutra] fluviatilis Burnett, Quart. Journ. Sci. Lit. Art. xxviii,
1829, p. 349 (Substitute for lutra), nomen nudum.
1834. [Lutra] nudijoes Melchior, Den Danske Stats og Norges Pattedyr,
p. 50 (Coast of northern Norway).
1834. Lutra roensis Ogilby, Proo. Zool. Soc. London, p. Ill (Roe Mills,
near Newton Lemavaddy, Londonderry, Ireland). Type in British
Museum.
1857. Lutra vulgaris Blasius, Saugethiere Deutschlands, p. 237.
1884. [Lutra] lutra Lataste, Actes Soc. Linn, de Bordeaux, xxxvni, 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 locality. — Upsala, Sweden.
Geographical 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 half 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 broad, 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, the 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 short 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
LUTKA
357
length) ; palm bare, a large, heart-shaped, obscurely trilobed
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 palm,
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 waterproof, 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. Mammae : 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-
RG. 72.
Lutra lutra. 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
conspicuous as well as paler, usually assuming a tinge of cream-
buff, the throat and cheeks fading to buffy white ; interramial
region and upper lip with irregular white mottlings, the hairs of
which are white to base ; whiskers and claws light horn-colour.
The exact colour is subject to considerable variation, but the
material examined is not sufficient to show whether such
differences as occur are correlated with locality or season.
* Southwell, Field, en, p. 1043, December 19, 1903.
358
CAKNIVOKA
Sometimes the brown is darker and richer than usual, or the
drab may be especially pronounced. Occasionally the long hairs
of the back are 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, broad posteriorly and with greatly
Fie.
Lutra lutra. Nat. size.
developed lambdoid crest, but low though evident sagittal crest ;
widely spreading zygomata forming a marked contrast with narrow
postorbital region ; rostrum 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 Dearly equals width of .flattened inter-orbital region.
Rostrum short, broad and deep, the distance from front of
zygoma to gnathion barely equal to greatest breadth across
canines, about equal to depth at front of orbit, and contained
about 4J 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
na. 74.
iMtra lutva. 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 equal to distance between molars and about one-third that
from molar to hamular ; mesopterygoid space narrow anteriorly,
wider posteriorly, its length about double the width between
hamulars. Auditory bulla small, flattened, though a little
360
CAENIVOBA
inflated along inner margin ; outline triangular-flask shaped, the
tubular portion of meatus not well defined from rest of bulla ;
least distance between bullae 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 off 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 areas, the
small premolars of upper and lower mandible fitting 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 1 and i 2 small,
sub-equal, the shafts compressed,
the cutting edges rounded, simple ;
i 3 higher than the others, somewhat
resembling the lower canine in form,
its apex curved outward and back-
ward beyond level of smaller teeth,
its postero-external surface with two
concavities separated by a trenchant
ridge ; lower incisors subterete, the
crown of « 3 obscurely and unsym-
metrically bilobed, that of the others
simple; cross section of i x about. one-third that of i 3 . Canines
robust though not greatly elongated ; shaft simple, without
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 1 small, single
rooted, its crown area about equal to that i 2 , its form essentially
like that of succeeding tooth, though with cusp relatively less
developed ; pm 2 and pm 3 successively larger, two-rooted, the
crown area of pm 3 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 2 elliptical, that of
pm 3 with inner margin bulging inward behind middle, though
without developing a definite inner lobe ; pm 2 and pm 3 much like
FIG. 75.
Lutra lutra. Teeth. Nat. size.
LUTRA 361
pm 2 in both size and form, the three teeth cutting against each
other when jaws are closed ; pm i larger than the other small
premolars, its cusp less 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 paraconid sub-equal, the metaconid somewhat smaller ;
crushing portion of crown wider than anterior 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 projection representing the parastyle, the
posterior lobe bearing a low but robust metacone ; inner 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.
Measurements. — 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. Por cranial and dental measurements
see Tables, pp. 362, 363.
Specimens examined. — Thirty-three, from the following localities : —
Ireland : Londonderry, 1 (type of roensis) ; Ahascragh, Oo. Galway, 1.
England : Pembrokeshire, 1 ; Eiver Stour, Dorsetshire, 1 ; Rugby,
Warwickshire, 1 ; Norfolk, 1 ; no exact locality, 1.
Nobwat : 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) ; Etupes, Doubs, 1 (Mottaz).
Germany : Southern Germany, 2.
Observations.
eth moderately worn.
, not worn.
much worn.
slightly worn.
, , not worn.
slightly worn.
>
moderately worn. g
, slightly worn. g
, not worn.
, moderately worn.
, not worn.
a
u
is
a
, moderately worn.
, not worn.
, slightly worn.
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364
CAKNIVOEA
Austbia-Hongary : Karlsbad, Bohemia, 1 ; Csehtelek, Bihar, Hun-
gary, 1.
Italy: Torriglia, Liguria, 1 (Lataste).
Spain: Castrillo de la Beina, Burgos, 3(B.M.andU.S.N.M.); Galioia, 1;
Seville, 5.
Remarks. — Although the specimens show some rather note-
worthy variations in colour, and in cranial and dental measure-
ments, the material examined is not sufficient 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.
9, 3 juv. st. Pembrokeshire, Wales.
5. Bugby, Warwickshire,
England.
9 st. B. Stour, Dorset.
1. England.
4 6, 1 9, Egersund, Stavanger,
skulls. Norway,
ad., juv. South Germany.
6. CsShtelek, Bihar, Hun-
gary.
1. Karlsbad, Bohemia.
6, 9. Castrillo de la Beina,
Burgos, Spain.
1 juv. Galicia.
skull. Seville.
9, juv. Seville. (Dr. A. Ruiz.)
skull. Seville.
Zoological Society's 57. 12. 14. 4.
Museum. (Type of L. roensis
Ogilby).
Purchased (Bow-
land Ward).
Purchased (Bow-
land Ward).
Dr.T. S.Townsend
(c & p).
J. 0. Mansel Pley-
dell (c & p).
K. H. Schaanning
(c).
Dr. A. Giinther (c).
Hon. Mrs. N. 0.
Bothschild (p).
Lord 0. Bussell
(c & p).
Bev. Saturio Gon-
zalez (c).
Dr. V. L. Seoane
(c & p).
Lord Lilford (p).
Lord Lilford (r).
Col. L. H. Irby
(c & p).
5. 7. 10. 1.
5. 5. 21. 1-4.
9. 12. 16. 1.
98. 5. 13. 1.
99. A.
11.6.3.14-18.
59. 9. 6. 62-63.
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 MUSTELINE.
1835. Mustelinn: Swainson, Nat. Hist, and Classif. Quadr., p. VII.
Geographical distribution. — Northern hemisphere, south into
northern Africa, the Malay Archipelago, and northern and
western South America ; in Europe west to Ireland.
Characters. — Dentition highly trenchant, upper carnassial
without crushing surface other than a small concave area 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
MARTBS 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.
Remarks. — As here understood the sub-family Mustelinm
contains the three genera, Martes, Mustela and Vormela, all of
which occur in Europe.
Genus MARTES Pinel.
1792. Martes Pinel, Actes Soo. d'Hist. Nat., Paris, i, p. 55 (M. domestica
Pinel = M. foina Erxleben).
1820. Martes Nilsson, Skand. Fauna, I, p. 38 (M. foina and M. sylvatica =
martes).
1829. Zibellina Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,
p. 31 (M. zibellina).
1857. Mustela Blasius, Saugethiere Deutschlands, p. 211.
1911. Martes Thomas, Proo. Zool. Soo., London, p. 139, March, 1911.
Type species. — Mustela martes Linnaeus.
Geographical distribution. — Northern hemisphere from the
limits of tree growth south to the Mediterranean, the Malay
Archipelago, and the eentral United States ; in Europe west to
Ireland.
Characters. — Skull narrow, moderately high (depth of brain-
case 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 between anteorbital foramina; auditory
bullse 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 : i t5, c t- 1 , prn ££, m t| = 38 ; cutting edges of five small
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
offset to antero -internal 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 flat, 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
366
CAKN1V0RA
a well defined postero-internal process on base of protoconid, the
posterior crushing heel slightly more than half as large as
trenchant portion 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.
Remarks. — The genus Martes contains about ten species, some
of which are represented by numerous geographical races. Three
are known from western and southern Europe.
KEY TO THE EUROPEAN FORMS OP 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 crown ; 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)... M. m. martes, p. 372.
Throat-patch buff-yellow, general colour
lighter (Mediterranean region) M.m. latvrurrum, 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. buwites, p. 380.
Condylobasal length of skull in adult male,
79 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 M.f. foma, p. 375.
General hue of upper parts wood-brown .. M.f. mediterranea, p. 380.
MARTES MARTES Linnaeus.
(Synonymy 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 premolar with outline of crown
MAETES 367
concave on outer side, irregularly convex on inner side, the region
of 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 oarnassial, the metacone more than half
as large as paracone, the outer border of the crown usually
notched; general colour yellowish brov^n, the throat patch
FIG. 76.
Ma/rtes martes. Nat. size.
always distinctly tinged with yellow ; underfur moderately long
and dense.
External characters. — Form slender and graceful, suggesting
that of both cat and squirrel ; legs moderately long ; tail about half
head and body, densely furred and bushy, the hairs at tip about
one-third as long as vertebrae. Head pointed ; eyes of medium
size ; ear rising distinctly above fur, its height from crown about
25 mm., the outline rounded but forming a slightly indicated
point above, the entire surface densely furred ; muzzle pad
368
CARNIVOKA
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 underfur
about 25 mm. long at middle of back in winter, shorter in
summer, the longer hairs reaching about 40 mm. Mammae : 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-colo'ur.
Skull. — As compared with that of the European species of
Mustela the skull of Maries mattes is characterized by a general
Fig. 77.
Maries martes. Nat. size.
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
MARTBS 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 bullse
moderately inflated, the surface smooth, the general outline
broadly flask-shaped (longitudinal diameter greatest), the meatal
tube well defined, rather wider than long ; space between bullse
about equal to diameter of bulla without tubular portion.
Postorbital region rather abruptly constricted, much as in
Mustela, its width about half that of brain-case. Interorbital
Fig. 78.
Marten martes. Nat. size.
region wider than rostrum, the discrepancy in breadth greater
than in Mustela erminea. Postorbital processes well developed
though short. Rostrum relatively longer than in Mustela
erminea or M. putorius, 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
CAKNIVOEA
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,
slightly everted.
Teeth. — In proportion to size of skull the teeth are larger
than in the members of the other genera of Mustelinse 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 1 and i 2 together ; crowns strongly
compressed, the antero-poscerior diameter fully twice transverse
diameter, the anterior face convex, the posterior concave with
faintly indicated cingulum ; i' 2 slightly
larger than i 1 , i 3 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
portion. Mandibular incisors smaller
and lower than the upper teeth, their
crowns projecting obliquely forward,
the root of i 2 implanted in jaw behind
level of the other two; crowns ob-
scurely bifid, the outer lobe smaller
than the inner ; crown area of i t
j. IG j 9 about one-third that of the com-
Martesmartes. Teeth. Nat. size, pressed i 2 or the subterete i 3 . Canines
with no special peculiarities ; diameter
of the upper tooth along alveolus equal to one-half width of
palate 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, especially 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 1 and i 2
together), subterete, the height of the single ill-developed cusp
scarcely equal to diameter of crown in pm 1 , much less in pm v
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 pm 2
about three times that of pm 1 ; outline of crown flattened-elliptical
MARTES 371
or slightly concavo-convex, the concavity, when present, on outer
side ; cusp simple, its height a little less than length of crown
along base, its posterior surface with slightly developed longi-
tudinal ridge. Crown of pm? somewhat longer than that of pm 2 ,
but its area much greater owing to widening of strongly convex
inner side ; cusp essentially as in pm 2 though somewhat higher.
Second lower premolar about equal in size to pm 2 , but axis of
shaft more anterior in position ; pm 3 essentially like pm 2 , but
cusp with faintly indicated anterior longitudinal ridge, and
posterior ridge with a slight nodule or rudimentary basal cusp
(sometimes absent) ; pm i noticeably larger than pm 3 and with a
well developed secondary cusp at middle of posterior border of
main cusp. Upper carnassial (pin 4 ) 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 1 , and separated from
main cusp 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 posterointernal 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 postero-external 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 paraoone and metacone,
the 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.
Remarks. — The well - known pine marten is so strongly
characterized as to require no special comparisons with any other
European species except Martes foina (see account of latter).
Two geographical races are currently recognized, though their
status is by no means clear.
Martes martes martes Linnseus.
1758. [Mustela] martes Linnaeus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1816. M[ustela] sylvesiris Oken. Lehrb. d. Naturgesch., hi, pt. 2, p. 1029
(Renaming of martes).
1827. M[artes] vulgaris Griffith, Guvier's Anim. Kingd., v, p. 123
(Renaming of martes).
1847. Martes sylvatica Nilsson, Skand. Fauna, i, 2nd ed., p. 41 (Renaming
of martes).
1857. Mustela martes Blasius, Saugethiere Deutschlands, p. 213.
1910. Mustela martes Trouessart, Paune Mamm. d'Europe, p. 72.
1911. [Martes] martes Thomas, Proo. Zool. Soc. London, p. 139, March,
1911.
Type locality. — Vicinity of Upsala, Sweden.
Geographical distribution. — Europe north of the Mediterranean
region, from Ireland eastward into Asia.
Diagnosis. — Throat patch cream-buff or slightly more yellow.
Colour. — 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 bufi'y drab ; tail very dark brown (nearly the seal-brown of
Ridgway), its underfur raw-umber or somewhat darker ; feet
and lower portion of legs blackish ; throat-patch varying
considerably, but as a rule rather pale, nearly cream-buff 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 ; Co. Kerry, no exact locality, 1.
Scotland : No exact locality, 1.
England : Keswick, Cumberland, 1 ; Cockermouth, 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 : Bohemia, 1.
MARTES 373
9. Kenmare, Kerry, Ire- E. Dodson (o). 94. 3. 27. 1.
land.
1. Kerry. J. H. Gurney (p). 72. 1. 10. 1.
1. Scotland. (McLeay.) Hargitt Collection. 86. 9. 9. 1.
st. Keswick, Cumberland, W. E. Wilson (c). 96. 11. 6. 1.
England.
4. Cockermouth, Cum- W. R. Wilson (c). 97. 1. 15. 1.
befland.
<$. Pontrilas, Hereford- Mrs. St. John A. 85. 6. 10. 1.
shire. Matthews (p).
4. Ludford Park, Here- H. J. Bailey (c & p). 0. 2. 23. 1.
fordshire.
2. Wales (1828). Earl Cawdor (p). 61. c. d.
2 4, 1 9, Egersund, Stavanger, K. H. Schaanning 11. 6. 3. 19-21.
skulls. Norway. (o).
2 skulls. Southern Germany. Dr. A. Gunther (c). 188i, 1299f.
i. Bohemia. Lord O. Russell 50. 12. 23. 1.
(c&p).
Martes maetes latinorum Barrett-Hamilton.
1904. Mustela martes latinorum Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xm, p. 389, May, 1904 (Nurri Mountains, Sardinia).
1910. Mustela martes latinorum Trouessart, Paune Mamm. d'Europe,
p. 72.
Type locality. — Nurri Mountains, Sardinia.
Geographical distribution. — Mediterranean region (Italy, Sar-
dinia, Balearic Islands).
Diagnosis. — Throat patch buff-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 Bidgway, the
general effect darker in certain lights, especially on posterior
half of body ; underfur a pale bluish grey, essentially as in true
martes, the tips of the hairs ranging from pale wood-brown to
clear buff; face isabella-colour streaked with whitish hairs, the
muzzle and lips between mars-brown and prout-brown ; ear
edged with buffy 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 :
head 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.
Sardinia : Nurri Mountains, 1 (type).
Spain : Mancor, Majorca, Balearic Islands, 1 ; San Cristobal, Minorca, 1.
Remarks. — The status of this form is very unsatisfactory. So
far as can be judged from the few specimens examined there is
374
CAENIVORA
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. Nurri Mountains, Sar-
dinia.
?. Mancor, Majorca, Bale-
aric Islands. (Biutort.)
9. San Cristobal, Minorca.
G. Barrett-Hamilton 11. 1. 2. 29.
B. Cavendish Taylor 5. 5. 6. 8.
E. N. Buxton (p). 95. i. 16. 1.
(Type of subspecies.)
0. Thomas (p). 1. 3. 6. 1.
0. Thomas & B. I.
Pocock (c & p).
0. 7. 1. 43.
MARTES FOINA Erxleben.
(Synonymy under subspecies.)
Geographical distribution. — Central and southern Continental
Europe, from the Atlantic coast eastward, and from the
Mediterranean to the Baltic.
Diagnosis. — Third upper premolar with outline of crown
nearly biconvex, the outer side occasionally
flattened, the greatest 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 portion of tooth behind
middle of crown ; upper molar not so large
„, f, as in M. martes, its greatest diameter less
Fiq. 80. than outer length of carnassial, the meta-
Larger cheek-teeth i of Martes cone less than half as large as paracone;
martes (re) and M.fmnaOi). .„ . ,.- s . r , . j. '
Nat. size. external form as in M. martes, but iur
of less fine quality ; colour usually more
greyish or drab than in M. martes and seldom with the rich
brown tints of the related species, the throat-patch never strongly
tinged with yellow. Mammse : 4.
Skull. — The skull resembles that of Martes 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 Mustela erminea and distinctly
less elevated than in Martes martes. Interorbital region and
rostrum wider than in Martes martes, and concavity of 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 differences.
Teeth. — Except as already pointed out the teeth agree with
those of Martes martes.
MARTES 375
Remarks. — Though readily distinguishable from Martes martes
by the character of the skull and teeth M. 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 quality 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.
Maktes foina foina Erxleben.
1777. [Mustela'] foina Erxleben, Syst. Regni Anim., I, p. 458 (Germany).
1792. Martes domestica Pinel, Aotes Soo. d'Hist. Nat., Paris, I, p. 55 (Prance).
1801. M[ustela~] f[oina~] alba Beohstein, Gemeinn. Naturgesoh. Deutsch-
lands, I, 2nd ed., p. 759 (Thiiringen, Germany).
1857. Mustela foina Blasius, Saugethiere Deutschlands, p. 217.
1869. Mustela martes var. fagorum Patio, Paune Vert. Suisse, I, p. 318
(Sweden) : name wrongly attributed to Linnseus, Syst. Nat., i,
12th ed., p. 67.
1910. Mustela foina Trouessart, Paune Mamm. d'Europe, p. 72.
Type locality. — Germany.
Geographical distribution. — 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 throughout 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 buffy drab externally, creamy white along rim, the inner
surface greyish white. Muzzle, lips and chin broccoli-brown.
Throat-patch buffy white, darkening to cream-buff along outer
edges.
Measurements. — Adult male from near St. Gallen, Switzer-
land : head and body, 453 ; tail, 260 ; hind foot, 85 ; ear from
meatus, 34. For cranial measurements see Table, p. 377.
Specimens examined. — Twenty-three, from the following localities : —
France : Manonville, Meurthe-et-Moselle, 2.
Germany : Ingelheim, Kheinhessen, 2 ; Nuremberg, Bavaria, 1
(U.S.N.M.) ; southern Germany, 7 (skulls).
Switzerland : Geneva, 1 (Mottaz) ; Vallee-de-Joux, Vaud, 1 (Mottaz) ;
Thayngen, Schaffhausen, 3 (B.M. and U.S.N.M.) ; Aarburg, Aarau, 1
(U.S.N.M.) ; Oberrich, St. Gallen, 1.
Spain: Silos, Burgos, 1 ; Dofiasantos, Burgos, 1 (U.S.N.M.).
Italy : Near Rome, 1.
376
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CARNIVORA
Greece : Mt. Vuno, Cephalonia, 1 (colour normal ; size apparently
rather small).
9, juv. Manonville, Meurthe-et- Lord Lilford (p). 95. 9. 5. 1-2.
Moselle, France.
5. Ingelheim, Rheinhessen, C. Hilgert (o). 8. 11. 2. 17.
Germany.
6. Ingelheim, Rheinhessen. G. Barrett-Hamilton 11. 1. 2. 102.
7 skulls. South Germany. Dr. A. Gunther (o). 1299, a-g.
2 6 juv. Schaffhausen, Switzerland. 0. Thomas (p). 2. 8. 4. 22-23.
(E. H. Zollikofer.)
S. 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. E. Cavendish Taylor 5. 5. 6. 7.
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1. Mt. Vuno, Cephalonia, J. I. S.Whitaker (p). 8.10.1.9.
Greece. (0. Mottas.)
Martes foina mediterranea Barrett-Hamilton.
1898. Mustela 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 distribution. — Southern Spain.
Characters. — Longer hairs of back tipped with a light
yellowish brown, the general hue of upper parts lighter, yellower
and less drab than in true foina.
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.
Remarks. — 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 approaches them, a
specimen from Rome.
1 juv. Sierra de Jerez, Cadiz, Spain. A Chapman (o & p). 98. 3. 18. 1.
(Type of subspecies.)
MARTES BUNITES Bate.
1899. Mustela foina leucolachnsea Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., iv, p. 383 (Not of Blanford).
1906. Mustela foina bunites Bate, Proc. Zool. Soc. London, 1905, u, p. 318,
April 5, 1906.
1910. Mustela foina bunites Trouessart, Faune Mamm. d'Europe, p. 73.
Type locality. — Kontopalo, Kania, Crete.
Geographical distribution. — Crete.
Diagnosis. — In general like Martes foina, but size not so
MUSTBLA 381
large (condylobasal length of skull in adult male not reaching
80 mm.), rostral portion of skull not so broad, and pale throat-
patch so much encroached on by brown of surrounding parts,
that it is occasionally absent.
Colour. — The colour is lighter and more yellowish (less drab)
than in Martes foina foina, 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, however, 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
seven 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 interramial region.
Skull and teeth. — The skull and teeth resemble those of
Martes foina except for their uniformly smaller size and the
relatively somewhat less breadth of the rostrum.
Measurements. — External measurements of type (young-adult
male) : head and body, 403 ; tail, 255 ; hind foot, 79 ; ear from
meatus, 39. For cranial and dental measurements see Tables,
pp. 377, 379.
Specimens examined. — Seven, all from Crete.
<5, 2. Kanea, Crete. Miss D. Bate (c). 5. 12. 2. 18-20.
1. Katharo. Miss D. Bate (c). 5. 12. 2. 21.
6. Kontopalo. Miss D. Bate (c). 5. 12. 2. 22.
(Type of species.)
skull. Crete. Miss D. Bate (c). 5. 12. 2. 37.
2. Crete. H. 0. Jones, B.N. (c). 99. 6. 13. 2-3.
Genus MUSTELA Linneeus.
(Synonymy under sub-genera.)
Type species. — Mustela erminea Linnaeus.
Geographical distribution. — Northern hemisphere from 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. — Skull in general resembling 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 flask-
shaped ; paroccipital process small and flattened, closely applied to
posterior margin of bulla ; dental formula : i ! ||, c 1=?, pm ~ 61
* A 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.
382
CARNIVOEA
m ? * = 34 ; upper carnassial as in Martes, its posterior cusp 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 heel crossed by a longitudinal
trenchant ridge ; other teeth essentially as in Martes ; 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.
Remarks. — 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 PORMS OP MUSTELA.
Brain-case broad, the mastoid width decidedly
greater than distance between basion and
palation ; auditory bullae triangular in outline
(Sub-genus Putorius, Polecats) M. putorius, p. 419.
Underfur usually bufly grey (Central Europe) M. p. putorius t p. 423.
Underfur usually yellowish (Southern Spain) M.p. aureolus, p. 425.
Brain-case narrow, the mastoid width less than
or about equal to distance between basion and
palation ; auditory bulte not triangular in
outline.
Rostrum flattened above ; inner margins of
auditory bullae strongly divergent pos-
teriorly ; tail bushy ; habits semi-aquatic
(Sub-genus Lutreola, Minks) M. lutreola, p. 415.
Rostrum convex above ; inner margins of auditory
bullse nearly parallel ; tail not bushy ; habits
terrestrial (Sub-genus Mustela, 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. hibemica, p. 398.
* At least so far as Lutreola and true Mustela are concerned. Putorius
appears to be sharply circumscribed, so that it might readily be regarded
as a distinct genus (see remarks on p. 419).
MUSTELA
383
Brown of sides not encroaching on yellowish
or whitish of underparts, the line of
demarcation straight ; upper lip wholly
or partly white M. erminea, p. 385.
Eegion 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) M. t. erminea, p. 387.
Begion extending 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) M. e. ricfiue, p. 397.
Zygomatic arches moderately spreading
(ratio of zygomatic breadth to condy-
lobasal length about 54) ; posterior
extension of palate not unusually
short and broad.
White winter ooat 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
Sweden to the Alps and Pyrenees) M. e. sestiva t p. 389.
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
and thickened (Great Britain) M. *. stabilis, p 390.
Tail without conspicuous black terminal area,
though a tuft of black or blackish hairs
may be present in pencil ; skull with ros-
trum usually wider than interorbital
region ; condylobasal length of skull, except
in 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 (africanus
group).
384 ■ CAENIVORA
Brown of sides noticeably encroaching on
yellowish or whitish of underparts,
the line of demarcation irregular
(Malta, Azores) M. africana, 412.
Brown of sides not encroachingon yellowish
or whitish of underparts, the line of
demarcation straight (Crete) M. galinthias, 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. n. nivalis, 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 46' 6 mm. in condylobasal
length ; hind foot of adult male 34 to
41 mm. ; colour rather dark (Central
Mediterranean region) M. n. botxamda, 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. iberica, p. 407.
Sub-genus MUSTELA Linnseus.
1758. Mustela Linnseus, Syst. Nat., i, 10th ed., p. 45 (Type by tautonymy
M. erminea Linnseus).
1829. Arctogale Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,
p. 30 (erminea).
1829. Ictis 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., n, p. 234 (vulgaris =
nivalis).
1857. Fcetorius Blasius, Saugethiere Deutschlands, p. 219 (part).
1865. Neogale Gray, Proc. Zooi. Soc, London, p. 114 (brasiliensis, aureo-
ventris and xanthogenys).
1871. Mustelina, Bogdanow, Tpjx OOm. Ectcctb. Ka3ancit. ymiBepc. I, Mem. i,
p. 167.
1899. Eumustela Acloque, Paune de France, Mammiferes, p. 62 (vulgaris
and erminea).
1911. Mustela Thomas, Proc. Zool. Soc. London, p. 138, April, 1911.
Type species. — Mustela erminea Linnseus.
Geographical distribution. — Northern hemisphere, south in the
Old World to northern Africa, and in the New "World to the
Andes ; in Europe west to Ireland.
Characters. — Form very slender ; tail not bushy ; fur not
modified for aquatic life ; skull without noticeably projecting
mastoid processes ; auditory bullse not triangular, the inner and
outer borders nearly parallel ; posterior border of pin 3 shearing
MUSTELA 385
against anterior border of pm 3 ; other small premolars not
capable of trenchant action.
Ben/iarks. — 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 LinniBUS.
(Synonymy under subspecies.)
Geographical distribution. — Europe from the Arctic coasst
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 bullse 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 denned, 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 vertebrae ; fur turning white in winter
except in warmer parts of range. Mammae : 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
OAKNIVOEA
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 fifth 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
FlO. 81.
Mustela ei'minea. Nat. size.
exception of the black area of the tail, which remains as in the
dark coat.
Skull. — In general appearance the skull resembles that of
Mustela putorius (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 anteorbital foramen as in M. putorius, but
orifice relatively larger, the width of plate separating 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. putorius, 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. putorius. 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 in 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
bullae 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 hamular, and separated from foramen
ovale by a broad nearly flat area, the length of which is
about one-half distance between foramina ; inner extremity of
glenoid surface marked off by a wide, shallow notch. Postorbital
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
Jlf. 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 m 1 with metacone more reduced, and m 2
with crown area distinctly smaller than that of heel of carnassial.
Mustela erminea eeminea Linnseus.
1758. [Mustela] erminea Linnseus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1792. M [ustela] erminea hyberna Kerr, Anim. Kingd., p. 181 (name applied
to the northern true ermine).
1816. M[ustela] herminea Oken, Lehrb. d. Naturgesch., in, pt. 2, p. 1026
(Benaming of erminea).
1827. [Mustela erminea] $ maculata Billberg, Synopsis Faunse Scandinaviae,
p. 8 (Scandinavia).
1857. Fcstorius erminea Blasius, Saugethiere Deutschlands, p. 228 (part).
2 C 2
388
CAENIVOEA
1877. Putorius erminea Coues, Fur-bearing Animals, p. 109 (part).
1910. Putorius (Ictis) ermineus Trouessart, Faune Mamm. d'Europe, p. 78
(part).
Type locality. — Upsala, Sweden.
Geographical 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 TJpsala, 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, Fillef jeld, Bergen, 1,
Bgersund, Stavanger, 13.
Remarks. — 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 ricin'se from
the islands of Islay and Jura, west Scotland. The white winter
coat is normally assumed throughout the range of this form.
<5, 6 juv. Aker, Christiania, Nor- Christiania Museum 93. 3. 1. 2, 4.
way. (e).
9. Brekkebygden, Trond- Christiania Museum 93. 3. 1. 3.
hjem. (e).
<5, 9. Nordre Fron, Gudbrands- Christiania Museum 93. 3. 1. 5-6.
dal. (e).
9. Marestuen, Fillef jeld, E. B. Alston (p). 79.9.25.21.
2,500 ft.
13 skulls. Egersund, Stavanger. K. H. Schaanning(c). 11.6.3.22-34
* In eight skulls of males from Egersund, Stavanger, Norway (all
sutures closed, teeth slightly or not worn), received after the Tables were in
typo, 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 estiva Kerr.
1792. M[ustela] erminea mstiva Kerr, Anim. Kingd., p. 181 (Germany;
based on Schreber's pi. 137a).
1820. Mustela erminea major Nilsson, Skand. Fauna, i, p. 34 (Carlskrone,
Blekinge, Sweden).
1857. Fcetorius erminea Blasius, Saugethiere Deutschlands, p. 228 (part).
1910. Putorius (Ictis) ermineus Trouessart, Faune Mamm. d'Europe, p. 78
(part).
Type locality. — Germany.
Geographical distribution. — Continental Europe 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 ; 
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