(N.H.)

f^'^W^

FOR THE PEOPLE

FOR EDVCATION

FOR SCIENCE

LIBRARY

OF

THE AMERICAN MUSEUM

OF

NATURAL HISTORY

#^1

3

tjyi^ <£=,-^-

CATALOGUE

OP THE

MAMMALS

OF

WESTERN EUROPE

(EUROPE EXCLUSIVE OF RUSSIA)

IN THE

COLLECTION

OF THE

BRITISH MUSEUM

^^

GEEBIT S. MILLEK

LONDON

PRINTED BY ORDER OF THE TRUSTEES OP THE

BRITISH MUSEUM

SOLD BY

Longmans, Green & Co., 39, Paternoster Row, E.G.

B. QuARiTCH, 11, Grafton Street, New Bond Street, W.

DuLAU & Co., Ltd,, 37, Soho Square, W.

and at the

British Museum (Natural History), Cromwell Road, S.W.

1912

All rights reserved,

3j^ l^y^^l S /Y^. 17

LONDON :

PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,

DUKE STREET, STAMFORD STREET, S.E., AND GREAT WINDJIILL STREET, W.

CD

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. MiJler, of the United
States National Museum at Washington, has for some years
been doing independently on the subject.

It is true that European Mammals had not been neglected
here, and that the collection had begun to grow, both by
the help of the Lilford Eund and by the efforts of Major
G. E. H. Barrett-Hamilton, who published many papers on
the subject, and of Mr. Oldfield Thomas, 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, Eev. S. Gonzalez and
N. Gonzalez, the cost of whose services were contributed by
friends of the Museum, notably Mr. Oldfield Thomas, the
Hon. N. C. Eothschild and Mr. J. I. S. Whi taker. 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 delinition 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. HARMEE,

Keeper of Zoology.

British Museum (Natural History),
London, S.W.

October, 1912.

INTEODUCTION

The collection of European Land-mammals in the British
Museum consists of about five thousand specimens. One
hundred and twenty-four of these are types. It has for the
most part been brought together during the past thirty years
through the efforts of the late Lord Lilford, of Mr. Oldfield
Thomas, and of Major G. E. H. Barrett-Hamilton. The older
material, though not extensive, includes much that is of historic
interest, such as the numerous specimens received from the late
Baron E. de Selys-Longchamps, the types of various species
described by Gray and Bonaparte, and Darwin's Porto Santo
rabbits which have been the subject of so much gi^oundless
speculation. It is, however, from the recently-obtained material
that the collection derives its true value. These specimens are
almost without exception carefully-prepared skins accompanied
by skulls and measurements, together with full records of sex,
date, and exact locality. The more important sources from
which they were obtained are as follows : collections brought
together by Professor W. Wolterstorff from central and eastern
(jermany, and presented by the late Lord Lilford ; collections
made in Roumania by the late E. Dodson under the direction
of Major G. E. H. Barrett-Hamilton, and presented by the late
Lord Lilford ; material from southern 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 IST. 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

VI INTRODUCTION

Mr. J. I. S. Whitaker and the Hon. N. C. Rothschild ; a
collection from Spain and southern France made by G. S.
Miller and purchased by the Museum ; several collections
from Transylvania made by C. G. Danford ; collections from
Hungary made and presented by the Hon. N. C. 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, Lunnberg). 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 viseful 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 3Ii/otis
oxygnathis 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 Vll

specimens of Lynx), Lausanne Agricultui-al School (skull of
Xlrma "formicarius " from the Alps), Munich (type of Spalnx
(jrsernn Nehring), Berlin Agricultural High School (type of
Arvicoln ratticeps stimmingi JSTehring), Breslau (skulls of foxes),
Leiden (co-types of Arvicola arenarius de Selys-Longchamps),
Copenhagen (Miis fseroensis and small carnivores), Christiania
{Sorex, Evotomys, etc.), Stockholm (Swedish carnivores and
rodents), Cambridge [Mustela erminea ricinse, Lemtnui^ lemmns
" rrassidens") 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. f

For the purposes of this Catalogue, "Western Europe" is
regarded as including the continent of Europe outside the
frontiers of Russia; also the immediately adjacent islands, and
Spitzbergen, Iceland, and the Azores. The members of the
living mammal fauna of this region, exclusive of the cetaceans,
pinnipeds, and species such as Buhalus hubalis in Italy and
Simia sylvanus | on the Rock of Gibraltar, which certainly owe

* In these lists 11,372 specimens are recorded. They are distributed
as follows: Insectivora, 1,777; Chiroptera, 2,210; Carnivora, 877 ; Rodentia
Duplicidentata, 379, Rodentia Simplicidentata, 5,854 ; Ungulata, 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.

VJll

INTRODUCTION

their presence to artificial introduction, are treated mono-
graphically on the basis of the material already enumerated.
This material has been found sufficient, in most of the groups, to
give what appears to be a fairl}?- satisfactory idea of the essential
features of the fauna. In the ungulates and the larger car-
nivores, however, it is so totally inadequate that no attempt
could be made to revise the genera by which they are repre-
sented. This is especially to be regretted on account of the fact
that some of these larger mammals are nearly extinct, while
others are being modified by the introduction of foreign stock
to replenish exhausted game preserves. Immediate action is
necessary if the final opportunity to gain a clear understanding
of this part of the European fauna is not to 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 diflerent from those underlying the
present work, that an amount of labour incommensurate with
the importance of the results would be required to prepai'e
extended bibliographical Tallies 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 diiferent 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. Tei'zi ; 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 IX

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,

July 1, 1912.

Order.

Number of

genera
recognized.

Numljer of

forms
recognized.

Number of

forms not

represented

in B.M.

Number of

recognized

forms not

seen.

Insectivoba

7

45

5

0

Chiroptera

10

33

0

0

Carnivora

15

47

1

0

RODENTIA DuPLICIDENTATA

2

19

1

0

„ Simplicidentata

26

139

6

2

Ungulata

9

31

9

4

Total

G9

314

22

G

SYSTEMATIC INDEX

ORDER INSECTIVORA.

3. russula Hermann .

PAGE

99

Family TALPID^E . . .

PAGE

2

a. russula Hermann

b. pulchra Cabrera .

101
103

Sub-Family 1. Talpin.-e . .

2

c. cintrae Miller. ...

108

1. Talpa Linnaeus

3

4. sicula Miller ....

108

1. europsea Linnaeus .

3

5. caueae Miller ....

109

2. caeca Savi ....

15

6. caudata Miller ....

110

3. occidentalis Cabrera .

15

7. cyrnensis Miller

111

4. romana Thomas

18

8. balearica Miller ...

112

Sub-Family 2. Desmanin.e .
1. Galemys Kaup

1. pyrenaicus Geoffroy
a. pyrenaicus Geoffroy

20
20
21
26

Family ERINACEID^ . . .
1. Erinaceus Linnaeus .
1. europaeus Linu;ieus.

114
114
115

b. rufulus Graells .

26

a. europEeus Linnaeus .

b. hispanicus Barrett-

120

Family SORICID^ . . .

28

Hamilton

122

1. Sorex Linnaeus

29

c. italicus Barrett-Hamil-
ton

d. consolei Barrett-Hamil-

123
126

1. araneus Liun^us .
a. araneus Linnaeus

31
35

h. castaneus Jenyns

37

ton

c. santonus Mottaz

40

2. roumanicuB Barrett-

d. euronotus Miller

41

Hamilton

127

e. bergonsis Miller .
/. tetragonurus Hermann
g. pyrenaicus Miller
h. fretalis Miller

41

42

3. nesiotes Bate ....

4. algirus Duveruoy and

129

44
45

Lereboullet ....
a. algirus Duvernoy and

130

i. granarius Miller .
2. minutus Linuasiis .

52
53

Lereboullet ....
b. vagans Thomas .

131
133

a. minutus Linu;eus .

55

b. lucauius IMiller .

60

3. alpinus Schinz .

60

ORDER CHIROPTERA.

a. alpinus Schinz .

62

b. hercynicus Miller

63

Sub-Order MICROCHIROPTERA.

2. Neomys Kaup ....

65

1. fodiens Schreber

66

Family RHINOLOPHID^ . .

136

a. fodiens Schreber.

69

1. Rhinolophus Lac^pede .

137

b. bicolor Shaw .

73

1. ferrum-equinum Schreber

139

2. miller i Mottaz .

78

a. ferrum-equinum Schre-

3. anomalus Cabrera .

81

ber

142

3. Pachyura de Selys-Long

b. insula n us Barrett-

champs

81

Hamilton

147

1. etrusca Savi ....

83

2. hipposideroR Bechstein

147

4. Crocidura Wagler .

86

a. hipposideros Bechstein .

149

1. leucodon Hermann

88

b, minimus Heuglin .

151

2. mimula Miller .

94

c. minutus Montagu .

154

a. mimula IMiller .

95

3. euryale Blasius ....

155

b. iculisma Mottaz .

98

4. mehelyi Matschie .

159

c. cautabra Cabrera

99

5. blasii Peters

162

SYSTEMATIC INDEX

aud

Order Chiroptera — Sub - Order

MiCROCHIROPTERA — COIltd.

Family VESPERTILIONID.E . 165

Sub-Family Vespertilionin.:

1. Myotis Kaup .

1. mystaciuus Kuhl .

2. nattereri Kuhl .

3. emarginatus GeolTroy

4. bechsteinii Kuhl .

5. daubentonii Kuhl .

6. capaccinii Bonaparte

7. dasycneme Boie

8. myotis Borkhauseu

9. oxygnathus Monticelli

2. Pipistrellus Kaup

1. pipistrellus Schreber

2. nathusii Keyserling

Blasius.

3. kuhlii Kuhl . . .

4. savii Bonaparte .

3. Eptesicus Rafinesque

1. serotinus Schreber

2. sodalis BarretT;-Hamiltou .

3. nilssonii Keyserling and

Blasius .

4. Vespertilio Liunteus
1. murinus Linnaeus

5. Nyctalus Bowdich

1. maximus Fatio.

2. noctula Schreber

3. leisleri Kuhl

4. azoreum Thomas

6. Plecotus Geoffroy .
1. auritus Linnaeus

7. Barbastella Gray .
1. barbastellus Schreber

Sub-Family Miniopterin.^l
1. Miniopterus Bonaparte
1. schreibersii Kuhl .

Family MOLOSSID^ .

1. Nyctinomus Geoffroy

1. teuiotis Rafinesque

ORDER CARNIVORA.

165
166
169
174
177
179
184
187
189
192
199
202
204

213
215
219
224
226
231

284
238
238
242
244
245
252
254
256
256
263
263

268
268
269

276
276

277

, Alopex Kaup .

1. lagopus Linnaeus

2. spitzbergenensis Barrett

Hamilton and Bonhote
Vulpes Oken

Family URSID/E ....

. 284

1. Ursus Linnaeus

. 285

1. arctos Linnaeus.

. 285

2. Thalarctos Gray .

. 297

1. maritimus Phipps .

. 298

Family CANID^ ....

. 308

1. Canis Linnaeus

. 304

1. lupus Linnaeus .

. 305

a. lupus Linnaeus .

. 313

b. signatus Cabrera

. 314

c. deitanus Cabrera

. 315

2. aureus Linnaeus

. 315

vulpes Linnaeus

a. vulpes Linnaeus .

b. crucigera Bechsteiu.

c. silacea Miller.
2. ichnusae Miller .

Family MUSTELID.^ . .
Sub-Family Melin^ .
1. Meles Brisson.

1. meles Linnaeus .

a. meles Linnaeus .

b. marianensis Graells.

2. arcalus Miller .
Sub-Family Luteins; .

1. Lutra Brisson ....
1. lutra Linnaeus .
Sub-Family Musteline .

1. Martes Pinel ....

1. martes Linnu3us

a, martes Linnaeus .

b. latinorum Barrett-Ha
milton

2. foina Erxleben .
a. foina Erxleben .
h. mediterranea Barrett

Hamilton ....

3. bunites Bate

2. Mustela Linnaeus.
Sub-gemis Mustela Linnaeus

1. ermiuea Linnseus .

a. erminea Linnaeus .

b. gestiva Kerr .

c. stabilis Barrett-Hamil
ton

d. ricinae Miller.

2. hibernica Thomas and

Barrett-Hamilton .

3. nivalis Linnaeus

a. nivalis Linnaeus .

b. boccamela Bechsteiu

c. iberica Barrett-Hamilton

4. africana Desmarest

5. galinthias Bate
Sub-Genus Lutreola Wagner

6. lutreola Linnaeus .
Sub-Genus Putorius Cuvier

7. putorius Linnaeus .

a. putorius Linnaeus .

b. aureolus Barrett-Hamil

ton

3. Vormela W. Blasius .

1. peregusna Gueldenstaedt
Sub-Family Gulonin.e .

1. Gulo Storr

1. gulo Linnaeus .

SYSTEMATIC INDEX

Order Carnivora — continued.

Family VIVERRIDtE . .

440

1

ISIungos Geoffrey and Cuvie

• 440

2

1. widdringtonii Gray
Genetta Oken ....

441
446

1. genetta Linnaeus .

447

a. genetta Linnteus

451

b. balearica Thomas

452

c. rhodanica Matschie .

452

Faiv

[iLY FELID^ ....

455

1

Felis Linnseus.

456

1. silvestris Schreber .

457

a. silvestris Schreber .

462

b. grampia Miller .

c. tartessia Miller .

464
465

2. sarda Lataste .

468

3. agrius Bate ....

470

2

Lynx Kerr

1. lynx Linnaeus .

470

471

2. pardellus Miller

476

ORDER RODENTIA.
Sub-Order DUPLICIDENTATA.

Family LEPORID^ .... 484

1. Oryctolagus Lilljeborg . . 484
1. cuniculus Linnfeus . . 485

a. cuniculus Linnaeus . . 490

b. huxleyi Haeckel . . . 491

2. Lepus Linnaeus .... 495

1. europaeus Pallas . . . 498

a. europteus Pallas. . . 502

b. occidentalis de Winton . 504

c. pyrenaicus Hilzheimer . 506

d. meridiei Hilzheimer . 506

e. corsicanus de Winton . 507
/. hybridus Desmarest. . 508
g. transsylvanicus Mat-
schie 509

2. creticus 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 Linnffius . . . 522

a. timidus Linnaeus . 526

b. varronis Miller . . . 528

c. scoticus Hilzheimer. . 529

6. hibernicus Bell. . . . 531

Family HYSTRIGID^ . .
1. Hystrix Linnaeus .
1. cristata Linnaeus .

Family IMUSCARDINIDiE .

1. Eliomys Wagner .

1. quercinus Linnaeus

2. gymnesicus Thomas .

3. pallidus Barrett-Hamilton

4. sardus Barrett-Hamilton

5. lusitanicus Reuvens .

2. Dyromys Thomas.

1. nitedula Pallas .

a. nitedula Pallas .

b. intermedius Nehring

c. wingei Nehring .

2. robustus Miller.

3. Glis Brisson ....
1. glis Linnaeus ...

a. glis Linnaeus ...

b. italicus Barrett-Hamil
ton

c. melonii Thomas .

d. pyrenaicus Cabrera .

4. Muscardinus Kaup

1. avellanarius Linnaeus, .

2. pulcher Barrett-Hamilton

Family MURID^ . .

Sub-Family Cricetin.e .

1. Cricetulus Milne-Edwards
1. atticus Nehring

2. Cricetus Leske
1. cricetus Linnaeus .

a. cricetus Linnaeus

b. canescens Nehring

c. nehringi Matschie

3. ]\Iesocricetus Nehring
1. newtoni Nehring .

Sub-Order SIMPLICIDENTATA.

Family ZAPODID^ .... 535

Sub-Family Sicistin.^
1. Sicista Gray

1. loriger Nathusius

2. trizona Pet^nyi .

Sub-Family Microtin^e .

1. Myopus Miller . .
1. schisticolor Lilljeborg

2. Lemmus Link.
1. lemmus Linnaeus .

3. Evotomys Coues .

1. glareolus Schreber .

a. glareolus Schreber

b. britannicus IMiller

c. suecicus Miller .
rf. istericus Miller .
c. uorvGgicus Miller
/. vasconiae Miller .
g. helveticus Miller
h, nageri Schinz
i. hallucalis Thomas

2. skomerensis Barrett -
536 j milton ...

536 j 3. caesarius Miller . .

537 4. rutilus Pallas . .
539 5. rufocanus Sundevall

Ha-

I-ACE

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 Rodentia — Sub - Order
SiMPLiciDENTATA — Family
MuEiDiE — Sub- Family Mi-
CBOTiN-E — continued.

4. Microtus Schrank . . . 658

Sub-Genus Microtus Schrank 659

1. agrestis Linnseus . . . 662

a. agrestis Linnaeus . . 668

b. exsul Miller .... 669

c. levernedii Grespon . . 671

d. bailloni de S61ys-Loug-
champs 672

e. hircus Bellamy . . . 673
/. neglectus Jenyns . . . 675
g. rozianus Bocage . . . 680

2. arvalis Pallas .... 681

a. arvalis Pallas . . . 683

b. meridianus Pallas . . 686

c. duplicatus Rorig and
Borner 686

d. levis Miller .... 687

3. incertus de S61ys - Long-

champs 690

4. asturianus Miller . . . 693

5. orcadensis Millais . . . 694

6. sandayensis Millais . . 696

a. sandayensis Millais . . 697

b. westrae Aliller ... 698

7. sarnius aiiUer .... 700

8. cabrerae 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

■ "' " . . 713

13. nivalis Martins

a. nivalis Martins .

b. aquitanius Miller

14. lebruuii Grespon .

a. lebrunii Grespon.

b. leucurus Gerbe .

15. ulpius INIiller .
5. Arvicola Lac6pede

1. amphibius Linnaeus

a. amphibius Linnaeus

b. reta Miller

2. sapidus Miller .

a. sapidus Miller

b. tenebricus Miller

3. terrestris Linnaeus.

4. italicus Savi 740

5. illyricus Barrett-Hamilton 741

6. musignani de Selys-Long-

champs

7. scherman Shaw

a. scherman Shaw

b. exitus Miller . . ,

c. monticola de Selys-
Longchamps ....

716
717
718
719
722
723
723
725
730
732
732
733
735
738

744
744
745
746

749

PAGE

6. Pitymys McMurtrie ... 752

1. subterraneus de S61ys-

Longchamps .... 755

a. subterraneus de Selys-
Longchamps .... 758

b. capucinus Miller . . 760

2. dacius Miller .... 760

3. druentius Miller . . . 762

4. fatioi Mottaz .... 763

5. multiplex Fatio . . . 764

6. savii de Selys-Longchamps 768

7. nebrodensis ]\Iina-Palumbo 770

8. pyrenaicus de S61ys-Long-

champs 770

a. pyrenaicus de S^lys-
Longchamps .... 771

b. brunneus INIiller ... 772

9. planiceps MiUer . . . 772

10. gerbii Gerbe .... 773

11. lusitanicus Gerbe. . . 776

12. mariae Major .... 777

13. pelandonius Miller . . 778

14. depressus Miller . . . 779

15. ibericus Gerbe . . . 780

a. ibericus Gerbe . . . 782

b. centralis Miller ... 782

c. pascuus Miller . . . 783

d. regulus ]Miller . . . 784

16. duodecimcostatus de

S61ys-Longchamps . 784

17. provincialis Miller . . 785

18. thomasi Barrett-Hamil-
ton 786

19. atticus Miller .... 787

Sub-Family Murin.^ .... 791

1. Apodemus Kaup .... 791

1. epimelas Nehring . . . 794

2. sylvaticus Linnaeus . . 797

a. sylvaticus Linnaeus . . 803

b. callipides Cabrera . . 809

c. dichrurus Rafinesque . 810

d. creticus Miller . . . 813

3. hebridensis de Winton . 824

4. hirtensis Barrett-Hamil-

ton 825

5. fridariensis Kinuear . . 825

6. flavicollis ]\Ielchior . . 828

a. flavicollis Melchior . . 829

b. wintoni Barrett-Hamil-

ton 831

7. agrarius Pallas .... 836

2. ]\Iicromys Dehne .... 840
1. minutus Pallas .... 841

a. soricinus Hermann . . 844

b. pratensis Ockskay . . 846

3. Epimys Trouessart . . . 848

1. rattus Linnaeus .... 849

a. rattus Linnaeus . . . 853

b. alexandrinus Geoffroy . 854

2. norvegicus Erxleben . . 858

SYSTEMATIC INDEX

Order Eodentia — Sub - Order
SiMPLiciDENTATA — Family
MuRiD^— Sub-Family Mu-
BiNiE — contimted.

4. Mus Linnaeus ....

863

1. musculus Linnaeus

865

a. musculus Linnseus .

869

b. azoricus Schinz .

871

2. muralis Barrett-Hamilton

874

3. fseroeusis Clarke .

875

4. spicilegus Pet^nyi .

877

a. spicilegus Pet^nyi .

878

b. hispanicus Miller

879

c. lusitanicus Miller

882

5. Acomys GeoSroy .

883

1. minous Bate

888

Family SPALACID^ . .

887

1. Spalax Gueldenstaedt

887

1. dolbrogese Miller .

889

2. hungaricus Nehring .

894

3. graecus Nehring

895

Family SCIURID^ . . .

897

1. Sciurus Linnteus .

898

1. vulgaris Linnseus .

898

a. vulgaris Linnseus

905

b. varius Gmelin .

906

c. leucourus Kerr .

907

d. russus Miller.

909

c. fuscoater Altum .

910

/. italicus Bonaparte .

912

g. lilseus Miller .

913

li. alpinus Desmarest .

914

i. numantius Miller

914

j. infuscatus Cabrera .

916

k. segurse Miller

917

I. bffiticus Cabrera .

923

2. Citellus Oken ....

924

1. citellus Linnseus

924

2. suslica Gueldenstaedt .

929

3. Marmota Blumenbach .

931

1. marmota Linnseus .

932

2. bobak Miiller . . .

937

Family PETAURISTID^E .

940

1. Sciuropterus F. Cuvier .

941

1. russicus Tiedemann .

941

Family CASTORID^ . .

947

1. Castor Linnseus .

947

1. fiber Linnseus .

947

ORDER UNGULATA

Family SUID^ ....
1. Sus Linnseus ....

1. scrofa Linnseus.
[attila Thomas] .

2. meridionalis JMajor

Family CERVID^ . . .

1. Cervus Linnseus .
1. elaphus Linnseus .

a. germanicus Desmarest

b. elaphus Linnseus

c. atlanticus Lonnberg

d. scoticus Lonnberg .
c. hispanicus Hilzheimer
/. corsicanus Erxleben.

2. Dama Hamilton Smith .
1. dama Lirmseus .

3. Capreolus Gray
1. capreolus Linnseus .

a. capreolus Linnseus .

b. transsylvanicus Matschie

c. canus Miller .

d. thotti Lonnberg.

4. Alces Gray

1. alces Linnseus .

5. Rangifer Hamilton Smith

1. tarandus Linnseus .

2. fennicus Lonnberg.

3. platyrhynchus Vrolik .

Family BOVID^ . .

1. Ovis Linnseus
1. musimon Pallas

2. Capra Linnseus .

1. ibex Linnseus .

2. pyrenaica Schinz

a. pyrenaica Schinz

b. lusitanica Franc^a

c. victorise Cabrera.

d. hispanica Schimper.

3. segagrus Erxleben

3. Rupricapra Blainville

1. rupicapra Linnseus

2. ornata Neumann .

3. pyrenaica Bonaparte .

4. parva Cabrera .

PAGE

956
956
957
960
960

962
963
964
965
967
967
968
969
969
970
970
972
973
974
975
975
975
976
978
979
980
981
985

986
986
987
988
989
989
990
991
991
991
992
992
993
994
995
995

APPENDIX.

Crocidura ichnusse Festa . . . 998
Evotomys glareolus norvegicus —

cranial measurements . . . 999
Spalax — Forms recognized by

M^hely 1000

CATALOGUE

OF THE

LAND-MAMMALS OF WESTERN
EUROPE.

Order INSECTIVORA.

1S27. Insectivora Gray, Griffith's Cavier, Anim. Kingd., v, p. 100.

Geographicdl distribution. — Africa (including Madagascar),
Europe, Asia (including the Malay Archipelago), North America,
I Greater Antilles, and extreme north-western portion of South
4. 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 difiicult 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) Soricidx, p. 28.

B

^ INSECTIVORA

Anterior lower incisor not elongated in axis of jaw ;
mandibular articulation single (normal) ;• zygoma
present ; floor of brain-case bony tbrougbout ; a
small auditory bulla ; general form not mouse-
like, tbe neck concealed between the greatly
enlarged shovdders ; no external ear (Moles and

Desmans) Talpidse, p. 2.

Teetb in front of molars sbarply differentiated by
form into incisors, canines and premolars, the
upper incisors small, sub-eqvial ; front feet
higbly modified for burrowing, the palms
everted ; tail scarcely as long as head (Moles) Talpinx, p. 2.
Teeth in front of molars not differentiated by
form into incisors, canines and premolars, the
inner upper incisor greath' 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 TALPID^.

1825. Talpidas Gray, Thomson's Annals of Philosophy, xxvi, p. 339.

Geographical distrihution. — 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 bvilla ;
mandible with single articulation, the glenoid surface normal ;
no external pterygoid plate ; crowns of upper molars low, much
narrower internally than externalh', the paracone and metacone
near middle of crown, the commissures and styles well developed
and forming with corresj^onding portions of lower teeth an
efiective 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.

Memarhs. — 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 Talpndse 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 TALPINX.

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. Tal2)a Linnseus, Syst, Nat., i, 10th ed., p. 52.

Type species. — Talpa europsea Linna3us.

Geographical distribution. — Europe and western and central
Asia. Eastern limits of range not known.

Characters. — Dental formula : i j^i,, c jEi, pw iir„ m ^^ — 44.
Upper premolars small, distinctly spaced, showing no tendency
to become imbricated. Lower canine slightly but evidently
difierentiated from incisors in size and form ; auditor}^ 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.

RemarJcs. — The genus Talpa contains half a dozen or more
species. Four of these occur in Europe, one of them generally
distributed, the three others confined to the Mediterranean
region.

KEY TO THE EUROPEAN SPECIES OF TALPA.

Greatest diameter of m^ about 4 mm. ; three lower
molars together about 8 mm. (vicinity of

Rome) T. romana, p. 18.

Greatest diameter of in^ about 3 mm. ; three lower
molars together about 7 mm. or less.
Condylobasal length of skull 33 to 37 mm. (dis-
tribution general) T. ciircpxa, p. 3.

Condylobasal length of skull 29 to 32 mm.
(southern).
Posterior border of anteorbital foramen over

front of last molar (Italian) T. cxca, p. 15.

Posterior border of anteorbital foramen over

middle of second molar (Iberian) T. occiclentalis,'Q. 15.

TALPA EUROPiEA Linnseus.

1758. [Talpa7\ 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. [Talpta] caudata Boddaert, Kortbegrip van het zamenstel der Natuur,

I, p. 50 (Renaming of europsea).

li 2

4 INSECTIVORA

1777. [Talpa europxa] a albo-maculata Erxleben, Syst. Regni Anim., i,

p. 117 (Ostfriesland).
1785. [Talpa] vulgaris Boddaert, Elenchus Anim., i, p. 12G (Europe).
1789. [Talpa etirop:m] ^ variccjata Gmelin, Syst. Nat., i, 13tli ed., p. 110

(Sweden).
1789. [Talpa europsea:] y alba Gmelin, Syst. Nat., i, 13th ed., p. 110

(Sweden).
1789. Talpia europxa e cinerea Gmelin, Syst. Nat., i, 13th ed., p. 110 (Eifel,

Germany).
1792. Talpia mrop[se.a\ nigra Kerr, Anim. Kingd., p. 200 (Renaming of

europsea).
1797. Talpa europsea rtifa B[orkhause]n, Der Zoologe (Compendiose Biblio-

thek gemeinniitzigsten Kenntnisse fiir alle Stande, pt. xxi).

Heft v-viii, p. 13 (Southern France).
1836. Talpa europxa flavescens Reichenbach, Pracht.-gemeinn. der

Saugeth. des In- und Auslandes, fig. 473 (Saxony).
1852. Talpa europxa albida Reichenbach, Vollstandigste Naturgesch. des

In- und Auslandes, iv, p. 336 (Germany).
1852. Talpa europxa lutea Reichenbach, Vollstandigste Naturgesch. des

In- und Auslandes, iv, p. 336 (Germany).
1857. Talpa europxa Blasius, Saugethiere Deutschlands, p. 109.
1869. Talpa europxa, flavescens Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lix, pt. i, p. 400.
1869. Talpa europxa, maculata Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lix, pt. i, p. 401

(Renaming of albo-maculata).
1869. Talpa europxa 5 grisea Fitzinger, 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, Abb. u. Vortr. Gesammtb. Naturw. iv,

no. 10, p. 19. (Substitute for europxa.)
1910. Talpa europxa Trouessart, Faune Mamm. d'Europe, p. 61.

Type locality. — Upsala, Sweden.

GeograpJdcal 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 fiist 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 m^ about
3 mm. ; mesostyle of m^ and m^ 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 integviment, 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, "w^th
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. Mamnife :
p 2 - 2 ; i 2 - 2 = 8.

Colour. — Fur everywhere dark slaty grey, sometimes almost
blackish, at others more nearly a dark smoke-grey, the hairs
everywhere with 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.

Sk-idl. — 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.

INSECTIVORA

Outline when viewed from the side long wedge-shaped, rounded
off posteriorlj. 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 antero-pos-
terior diameter about one-third
transverse diameter. Its anterior
and lateral sutures remain visible,
but the lambdoid suture is nearly
obliterated earh^ in life, though
its position is usually marked by
a slight ridge representing the
lambdoid crest. Base of brain-
case smooth, without cousf)icuous
ridges, furrows or open sj^aces,
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 r/<-, the plate forming outer wall of canal usually
much narrower than foramen Lachrymal foramen above anteor-

FIG. 1.

Talpa europeea. Nat. size.

bital foramen and slightly in front of its middle ; its orifice over
metastyle of m^. 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 in^, 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^ and m''. Mandible I'ather 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 e(iual to least bi'eadth 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. Up2:)er
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 canine very small, resembling a
fourth incisor, but with shaft conical instead of chisel-shaped,

* The roots of this tooth as well as those of the small premolars are
distinctly visible in old individuals.

KIG. 2.
Talpa europxa. Teeth x 5.

8 INSECTIVORA

and set in the jaw at a slightly different angle posterior
surface of shaft with well developed longitudinal ridge. First,
second and third upper premolars two-rooted, small, their
points on level with those of incisors and inner cusps of
molars, their crowns separated from each other as well
as from canine and large premolar by narrow equal spaces ;
crowns rather higher than long, compressed, with slightly
developed posterior cutting edge, narrowly triangular in outline
when viewed from the side, the first more slender than the
others, the second and third with slight though evident postero-
external angle on cutting edge. Fourth premolar essentially
like the other three in form, but much larger and three-rooted,
the length of crown greater in proportion to height ; no secondary
cusps. Lower premolars two-rooted, similar to the upper teeth
in form, the second and third the same size as smaller upper
premolars, the first nearly as large as pm'^, but narrower,
owing to absence of third root, the fourth intermediate. Upper
molars with crowns much wider externally than internally,
and completely divided into two unequal sections by a deep
longitudinal groove 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'-.
Paracone and metacone sub-equal in m'^ and m^, the metacone
the larger in the former, the paracone in the latter. In in^ 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.j. Styles and outer commissures well
developed in m^ and forming a distinct VV "Pattern ; mesostyle
entire or with apex slightly notched. In m^ the parastyle is
reduced to a minute though usually evident cusplet on the
cingulum, and the mesostyle to a thickening or angle in
commissure connecting the two main cusps ; metastyle well
developed.! In m'^ the parastyle, mesostyle and their commis-
sures are well developed, metastyle and fourth commissure
absent ; mesostyle with apex usually bifid. Middle lower molar
largest, its protoconid the highest cusp in the series. First lower
molar broader posteriorly than anteriorly, second and third
slightly broader anteriorly than posteriorly, the two triangles
essentially alike in form. Metaconid of m^ 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.

+ In this tooth there is no anterior V, the outer surface of paracone
essentially resembling that of pin' except for its smaller size.

TALPA 9

than a slight thickening of the cinguhmi. 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 ; Grautown-on-Spey, Elgin, 1:3 (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 localitj', 2 ; Fulbourn, Cambridge, 1 ; Holloway,
Somersetshire, 2 ; Somersetshire, no exact locality, 1 ; Banstead, Surrey, 2 ;
Coombe, Surrey, 1 ; Cobhaiu, 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.).

France : Guines, Pas-de-Calais, 1 ; Pont-de-Briques, Pas-de-Calais, 1 ;
Trinity, Jersey, 2 ; St. Lawrence, Jersey, 1 ; Barbizon, Seine-et-Marne, 2 ;
Meluu, Seine-et-Marne, 1 (Mottaz) ; Lignieres, Charente, 1 (Mottaz) ;
Huelgoat, Brittany, 1 ; Cadillac-sur-Garonne, Gironde, 1 (U.S.N.M.) ;
Foret de Bouconne, Gers, 13 ; Solferino, Landes, 10 ; Caterille, Haute-
Garonne, 10; Legouvin, Haute-Garonne, 3; Luchon, Haute-Garonne, 3;
Bareges, Hautes-Pyr^n^es, 5; I'Hospitalet, Ari^ge, 1; Port6, Pyr6n6es-
Orieutales, 3 ; St. Gilles, Gard, 1 ; Valescure, Var, 1 ; Agay, Var, 1 ;
Etupes, Doubs, 3 (Mottaz) ; Lucinges, Haute-Savoie, 9 ; Moutauban,
Haute-Savoie, 15 ; Cranves-Sales, Haute-Savoie, 12.

Spain: Pajares, Leon, 2; Castrillo de la Reina, Burgos, 9; Castafiares,
Burgos, 8 ; L6rida, 1 ; Barracas, Castellon, 24 ; Catarroja, Valencia, 4 ;
Dehesa de Valencia, Valencia, 17.

Germany : Konigsberg, 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, Rheinhessen, 4; Strass, near Burg-
heim, Bavaria, 7 ; Niesky, Silesia, 1 ; Kalbe, Saale, 2 ; Magdeburg,
Saxony, 1 ; Strassburg, 3.

Austria-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 2 ; Hatszeg,
Hunyad, Transylvania, 2.

10

INSECTIVOKA

EOUMANIA : 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.) ; Miirren, 1 ; Thurgau, 1 ; Oberhasli Valley
1 ; St. Gallon, 3 ; Rheinthal, St. GaUen, 2; Degersheim, St. Gallen, 3
Gossau, St. Gallen, 8 ; Untervatz. Grisons, 1 ; Breganzona, Ticino, 1
Cortivallo, Ticino, 6 (B.M. and U.S.N.M.) ; Comano, Ticino, 2 (U.S.N.M.)
Lugano, Ticino, 4 (B.M. and U.S.N.M.); Muzzano, Ticino, 2 (B.M. and
U.S.N.M.) ; Sorengo, 1 (U.S.N.M.) ; Gremignone, Ticino, 1.

Italy : Turin, 4 (U.S.N.M.) ; Certosa di Pesio, Cuneo, 1 (Genoa) ;
Parma, 2 (U.S.N.M.) ; Gozzauo, Novara, 3 (Genoa) ; Frugarolo, Ales-
sandria, 4 (Genoa) ; Vaccarezza, 1 (Genoa) ; Perti, Finalborgo, 10 (Genoa) ;
Florence, 10 (U.S.N.M.).

BemarJcs. — With the possible exception of certain bats, the
common mole shows less tendency to vary geographically than
any other European mammal of equally wide range.

2 S. Black Isle, Cromarty, Scot-
land.
?. Gordonstown, Elgin.

6. Stockbriggs, Lanarkshire.
?. Parsop, Herefordshire, Eng-
land.
2 6. Rugby, Warwickshire.
2. Warwickshire.
6. Fulbourn, Cambridgeshire.
2 al. Holloway, Somerset.
2. Somerset. [Hiigcl.)

2 al. Cobham, Surrey.
1 al. 1. Egham, Surrey.

?. Knockholt, Kent.
2(5. al. Bromley, Kent.

3 6. Devonshire.

2 S. Trinity, Jersey, Channel
Islands. [R. H. Bunting.)

9. St. Lawrence, Jersey.

S. Guines, Pas-de-Calais, 80 ft.
France.

5. Pont -de-Briques, Pas -de-

Calais.

6. Huelgoat, Brittany, 600 ft.
2c5, 2 9. Foret de Bouconne, Gers,

250 m. [A. Robert.)
2(5,2 9. SolfMno, Landes. (-4. Ro-
bert.)
2 (5, 2 9. Caterille, Haute - Garonne,
900-1000 m. {A. Robert.)
2 S, 9. Luchon, Haute - Garonne.
[A. Robert.)
2 6,19. Bareges, Hautes-Pyrt^nees.
6. L'Hospitalet, Ariege. [A.

Robert.)
6. Port6, Pyrenees-Orientales.
2 S Porte, Pyr^'u6es - Orientales,
1600-1700 m. (.4. Robert.)
6. Valescure, Var.

W. R. Ogilvie-

Grant (p).
W. R. Ogilvie-

Grant (p).
E. R. Alston (p).
E. A. Denny (p).

E. E. Austen (p).
Tomes Collection.

H. King (p).

E. R. Alston (p).
Dr. Leach (p).

F. Heiss (p).

W. Blackwell (p).
H. E.Rawson (p).
Oxley Grabham

0. Thomas (p).

Mrs. Power (p).
0. 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.

0. Thomas (c & p). 98. 1. 9. 3.

0. Thomas (c & p)
0. Thomas (p).

0. Thomas (p).

0. 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. 36-87.

G. S. Miller (c). 8. 8. 4. 134.

TALPA 1 1

6. Agay, Var. ~ G. S. Miller (c). 8. 8. 4. 135.

1. Moutauban, Haute - Savoie, A. Robert (c & p). 97. 1. 9. 1.

1000 m.
3 6, 19. Cranves-Sales, Haute-Savoie, A. Robert (c & p). 5. 4. 4. 1. 5. 4,

900 m. 9. 3-5.

5 c5, 3 9. Cranves-Sales, Haute-Savoie, 0. Thomas (p). 5. 11. 18. 1-8.

909-1200 m. Prance. (A.

Bohert.)
6, 9. Pajiires, Leon, Spain. {N. O. Thomas (p). 8. 2. 9. 26-27.

Goii:;ale2.)

2. Castrillode la Reina, Burgos, G. S. Miller (c). 8. 8. 4. 20-21.

Spain.
2 6, 5 9. Gastrillo de la Reiua, Burgos. N. Gonzalez (c.) 8. 7. 7. 1-7.

6. L6rida, Spain. (N. Gonzalez.) 0. Thomas (p). 8. 2. 9. 28.

6 c5, 6 9. Barracas, Castellon. (N. Con- 0. Thomas (p). 8. 2. 9. 29-40.

zalez.)
3 6, 9. Catarroja, Valencia. {N. Gon- 0. Thomas (p). 8. 2. 9. 12-15.

zalez.)
4<i, 5 9. Dehesa de Valencia, Val- 0. Thomas (p). 8.2.9.17-25.

encia. (A^. Gonzalez.)
8(5,9. Ingelheim,Rheinhessen,Ger- C. Hilgert (c). 8,11.2.11-14.

many.
6,9. Ummerstadt, Thliringen. Lord Lilford (p). 11.1.1.115-116.
{Schucliardt.)
6. Ummerstadt, Thliringen. Lord Lilford (p). 11.1.1.137.
{Schucliardt.)
c5, 3 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.1-4.

{Kijrbitz.)
2 (5, 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.132-134.
[Kurbitz.)
6. Niesky, Silesia. (W. Baer.) Dr. E. Hamilton 97. 12. 4.18.

9. Kalbe, Saale. {W.Bauer.) Lord Lilford (p). 11.1.1.5.
6. Kalbe, Saale. {W. Bauer.) Lord Lilford (p). 11. 1. 1. 136.
6. Magdeburg, Saxony. {Wolter- Lord Lilford (p). 11. 1. 1. 135.

storff.)
6,9. Strassburg, Alsace. {C. Mot- 0. Thomas (p). 8.8.10.10-11.

taz.)
2. Csallokciz-Somorja, Pressburg, Budapest Museum 94. 3. 1. 27-28.

Hungary. (p).

2 6. Hatszeg, Hunyad, Transyl- C. G. Danford (c). 3. 2. 2. 20-21.

vania, Hungary.
6. Bustenari, Prahova, Rouma- Lord Lilford (p). 4. 4. 6. 13.

nia. {W. Dodson.)
6. Oomana, Vlasca, Roumania. Lord Lilford (p). 4. 4. 6. 14.
1. Miirren, Switzerland. W. Gartner (p). 92. 10. 5. 3.

9. Thurgau, 400 m. Switzerland. O. Thomas (p). 4. 4. 5. 26.

{E. H. Zollikofer.)
1. Oberhasli Valley, Switzerland. Tomes Collection. 7.1.1.130.

{Koesej-man.)
2 6,9 St. Gallen, 650 m. Switzer- 0. 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).
6. Cortivallo, Ticino. {E. H. 0. Thomas (p). 2. 8. 4. 15.

Zollikofer).
9. Lugano, Ticino, 300 m. O. Thomas (c & p). 2. 7. 1. 1.

6. Muzzano, Ticino. {E. H. O. Thomas (p). 2. 8. 4. 16.

Zollikofer).
9. Cremignonc, Ticino. {E. H. 0. Thomas (p). 2. 8. 4. 20.

Zollikofer).

12

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15

TALPA C^CA Savi.

1822. Talpa cxca Savi, Nuovo Giorn. de' Letterati, Pisa, i, p. 265.
1857. Talpa cmca Blasius, Stiugethiere Deutschlands, p. 115.

1906. Talpa. cmca Camerauo, Boll. ]Mus. Zool. ed Anat. Comp. della K.

Univ. di Torino, xxi, No. 530, p. 1, June 22, 1906.
1910. Talpa cxca Trouessart, Faune 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 europgea (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 /;*'- and m^ with
apex deeply notched (this character not visible in specimens with
much worn teeth).

Measurements. — Average and extremes of five specimens from
Cortivallo, Ticino, Switzerland : head and body, 133 • 8 (125-142) ;
tail, 25-2 (21-30) ; hind foot (dry), 15-6 (15-16). Average and
extremes of three specimens from Reggello, Tuscany, 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: —
Switzerland : 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.

Italy : Regello, Tuscany, 4 (U.S.N.M.) ; N.S. della Vittoria, Ligurian
Appenines, 1 (Genoa) ; no exact locality, 2.

6. Agmizzo, Ticino, Switzerland. O. Thomas (p). 2.8.4.18.

{E. H. Zollikofer.)
i. Stabio, Ticino. {E. H. Zol- 0. Thomas (p). 2. 8. 4. 19.

likofer.)
c5, 9 al. Italy." Dr. Riippell (c). 45.7.22.32-38.

TALPA OCCIDENTALIS Cabrera.

1907. Talpa cxca occidentalis Cabrera, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 212, September, 1907. Type in Cabrera collection.
1907. Talpa ca&ca occidentalis Cabrera, Bol. Real. Soc. Espaii. Hist. Nat.,

Madrid, vii, p. 222, October, 1907. (For date see Cabrera, Ann..

and Mag. Nat. Hist., 8th ser., i, p. 189, February, 1908.)
1910. Talpa cmca occidentalis Trouessart, Faune Mamm. d'Europe, p. 63.

Type locality. — La Granja, Province of Segovia, Spain.
Geographical distribution. — Ibei'ian Peninsula.

16

INSECTIVORA

Dia(/no.sis. — Like Talpa cseca but skull rather robust, the
breadth of I'ostrum over roots of canines usually more than
4 mm. ; anteorbital foramen contracted, its posterior border over
mesostvle of »r, 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 Taljja csecn.

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
is noticeabl}^ smaller than in Talpa
cseca, 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 cseca. Zygoma
about as long as in Talpa cseca, 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 cseca, but otherwise the teeth show no peculiarities.

Measurements. — Type (from Cabrera) : head and body, 102 ;
tail, 24; hind foot, 15-5. Average and extremes of five
specimens from La Granja, Segovia, Spain (in spirit, body
contracted): head and body, 101-6 (98-107); tail, 25-6
(25-27); hind foot, 16-4 (16-17). Adult male from Galicia :
head and body, 112; tail, 26; hind foot, 15 "4. Two males
from Cintra, Portugal: head and body, 118 and 120; tail, 26
and 26 ; hind foot, 16 and 17. For cranial measurements see
Table, p. 17.

Specimens examined. — Ten, from the following localities : —
Spain : Galicia, 1 ; La Granja, Segovia, 5.
POBTUGAL : Cintra, 4.

Fig. 3.
Talpa ceeca (upper flgiu'e), aud
T. oceidentalis (lower figure).
Xat. siz3.

lal. Galicia, Spain.
2(J,2?al. La Granja, Segovia,
Spain.
Skull. La Granja, Segovia.
2 <J, 1 al. Cintra, Portugal.
IskuU.

Prof.Seoane (c&p). 9d. 1. 1. 23.
M.delaEscalera(c). 8. 7. 30. 19-22.

M.de la Escalera(c) .
O.Thomas (c&p).

6. 11. 4. 2.
98.2. 2.8 9,58,
59.

17

Observations.

Teeth not woru.

,, slightly worn.

„ moderately worn.

,, not worn.

,, slightly woru.

,, moderately worn.

,, not worn.

,, slightly worn.

„ much worn.

,, moderately woru.

,, slightly worn.
„ not woru.

, slightly worn.
, not woru.

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18

IXSECTIVOEA

TALPA ROMANA Thomas.

1902. Talpa roinana Thomas, Aun. and Mag. Nat. Hist., 7th ser., x, p. 517,

December, 1902. Type in British Museum.
1904. Talpa romana Camerano, ilem. Eeale Accad. Sci. di Torino, 2Dd ser.,

Liv, p. 81.
1910. Talpa romana Trouessart, Faune Mamm. d'Europe, p. 64.

Type locality. — Ostia, Rome, Italy.

Geographical distrihution. — Vicinity of Rome, Italy.

Diagnosis. — Externally similar to Talpa europsea but a trifle
larger ; skull and dentition more robust than in the related
animal (greatest diameter of m} about 4 mm. ; length of upper
tooth-row, exclusive of incisors, 14 to 15 mm.) ; mesostyle of all
three upper molars bifid at tip.

Skull and teeth. — Except that it is rather more robust the
skull does not differ appi'eciably 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 vipper
molar and in the large lower cheek-teeth (the combined length of

Fig. 4.
Crown of molars in Talpa europsea («), and T. romatia {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^ 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. europita, its edge fi-equently
forming an evident cusp, especially in m.^.

Measurements. — External measurements of type (from
Thomas): head and body, 126; tail, 29; hind foot, 19 (all
measurements from skin). Four males from the vicinity of
Rome (in alcohol) : head and body, 138-3 (130-145); tail, 28-6
(27-30) ; hind foot, 19-6 (19-20). Three females from the same

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20

INSECTIVOEA

locality: head and body, 127-5 (125-130) ; tail, 29-5 (29-30) ;
hind foot, 18"4 (18-18"6). For cranial measurements see Table,
p. 19.

Specimens exatnined. — Fourteen, from the vicinity of Rome (B.M. and
U.S.N.M.) ; also about fifty from the same region in Turin Museum.

Rtmarks. — 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 ol species.)
1. Frascati, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 9.

6, 9 al. Rome. Genoa Museum (e). 3. 1. 31. 1-2.

2 6. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 1-2.

Sub-Family DESMANIN^.

Geographical distribution. — South-western France and northern
half of Iberian Peninsula ; eastern Russia and western Siberia.

Characters. — Teeth in front of molars not differentiated by
form into incisors, canines, and premolars ; anterior upper
incisor greatly enlarged, canine-like, directed downward, its
outer edge highly trenchant, the two anterior lower incisors
slightly elongated, projecting obliquely forward ; external form
less evidently mole-like than in the 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.

Bemarks. — 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 TaJpinse.

Genus GALEMYS Kaup.

1829. Galemys Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,

p. 118 {Mygale pyrenaica Geoffroy).
1846. Galomys Agassiz, Nom. ZooL, Index Univ., p. 149 (Emendation of

Galemys).
1849. Mygalina I. Geoffroy, D'Orbigny's Diet. Univ. d'Hist. Nat., iv, p. 709

{Mygale pyrenaica Geoffroy).

Type species. — Mygale pyrenaica Geoffroy.

Geographical distribution. — Northern half of Iberian Peninsula,
and Pyrenean region of south-westei-n 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.

GALEMi'S 21

Remarls. — The genus Galemys is readily distinguishable from
the Russian and Siberian Desmana* in which the tail is flattened
laterally throughout, the unicuspid teeth are low and thick, their
width about equal to height of crown, the main cusp of large
premolar is rounded anteriorly, and the ridges on brain-case are
unusually developed. It contains a single species, peculiar to
south-western Europe.

GALEMYS PYRENAICUS Geofi"roy.

(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 equalh^ short (so that auditory oritice
appears to be at shoulder) and the muzzle similarly produced ;
legs less shortened than in the moles and front feet not
specially enlarged ; tail rat-like, longer than head and body.
Fur less 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

* Tlie synonymy of this geuus is as follows : —
1777. Desmana Gueldenstaedt, " Beschiift. Berliner Gesellsch. Naturforsch.
Freunde, iii, p. 108." [Castor moschatus Linnaeus.)

1799. Desman Lacepede, Tabl. Mamm., p. 7. Same type.

1800. MygrflZt'Cuvier, Lec.d'Anat. Comp., i.Tabl. 1. Same type. (Described

in Tabl. Elem. d'Hist. Nat. des Anim., p. 109.)
1815. Desmanus Rafinesque, Analyse de la Nature, p. 59 (Emendation of
Desvian).

1829. Myogalea Fischer, Synops. Mamm., p. 250 (Substitute for Mygalc).

1830. Caprios Wagler, Nat. Syst. Amphibien, p. 14 (Substitute for Mygale).
1836. Myogale Brandt, Wiegmann's Archiv fiir 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 fineh^ 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. Front 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

I'lG. 5.

Galemys pyrenaicus. Xat. size.

GALEMYS 23

a rudimentary dorsal and ventral keel. Mammie ijsl — 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 ellect 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.

Shiill. — In general the skull resembles that of Talpa europsea,
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 ^jentagonal, 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 antero-posterior
diameter nearly equal to its width. Base of brain-case with
deep but broad median furrow, the surface of the bones more
angular and less inflated than in Talpa ; no pits in basioccipital
in front of foramen magnum. Bullse low and flattened, less
perfectly formed than in Talpa, the tympanic bone annular and
retaining its distinctness, though joined with surrounding parts ;
meatus large, occupying about one-half surface of bulla.
Mesopterygoid space short, its length scarcely equal to width of
palate between posterior molars, its width about one-third length,
its anterior border double rounded, encroached on by slight
median spine. Outer pterygoid plate reduced to a small but
evident ridge. Interorbital region hour-glass shaped, widening
moi-e 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'^. Posterior border of
anteorbital foramen over parastyle of m'-. 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 Talpn.

Teeth. — Dentition relatively heavier than in Talpa europeea,
the teeth, with exception of anterior upper incisors, less trenchant

in general aspect. An-
terior upper incisor much
the largest of all the
teethj the two together
closing entire front of
palate ; shaft triangular
in cross section, and all
three faces sub-triangular
in outline, the posterior
and antero-external faces
widest, the former slightly
concave, the latter
slightly convex, the two
forming a perpendicular
external cutting edge
about 4 mm. in length
along their line of con-
tact ; a much shorter but
well developed cutting
edge along line of contact
of posterior and antero-
internal faces, extending
from acutely triangular-
pointed apex of tooth to
point of contact with
tooth of opposite side,
a distance of about
0 • 6 mm. ; line of contact
between antero-external
and antero-internal faces
marked by a slight though evident ridge ; height of shaft
about equal to width of palate ; first and second lower incisoi's
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 w^hich

Fig. 6.
Galemys 2)yrenaicus. Teeth X 5.

acts in opposition to combined posterior surface of large upper
incisors. Upjier 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 tits when jaws are closed. Two anterior
unicuspids (/'- and P) minute, terete, single-rooted, their axes
directed backward and falling in same line with those of two
anterior lower unicuspids when jaws are closed. Third unicuspid
(canine) two-rooted, its crown perpendicular, compressed, larger
than those of first and second combined, and distinctly greater
in height. Fourth unicuspid (pm^) single-rooted, subterete,
scarcely larger than second. Fifth and sixth unicuspids (jjto-
and jjiH^) essentially like canine, their crowns distinctly higher
than wide, comjiressed obliquely to the tooth-row, with slightly
developed anterior and posterior cutting edge. Height of third
and fifth sub-eqvial, 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 2mt'^ 3,nd j^fP 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 Soricidte.
First and second ({3 and ^) higher than third, their form suggest-
ing that of anterior lower incisor, all three single-rooted.
Fourth larger than any of the first three, obscurely two-rooted ;
fifth slightly smaller, single-rooted ; sixth (pm^) lai'gest 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 nP. Upper
molars with crowns wider and less oblicjue than in Talpa
etiropsea, and main cusps not so high. Transverse groove
between bases of main cusps converted into a median pit by
better development of commissures of protocone and larger size
of protoconule and metaconule. Paracone and metacone about
equal in height, the latter slightly the more robust. Styles well
developed, except the reduced parastyle of m^, the mesostyle in
each tooth completely divided into two cusps, the W-pattern thus
changed into two V-shaped figures. Third upper molar with
crown area about two-thirds that of second, its metastyle and
fourth commissure absent. Lower molars essentially as in
Talpa europsea, but contrast in height of outer and inner
cusps very slight.

26 INSECTIVORA

Galemys pyrenaicus pykenaicus Geoffroy.

1811. Mygale pyrenaica Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii,

p. 193.
1910. Myogale pyrenaica Trouessart, Faune IMamm. d'Europe, p. 60.

Type localiti/. — Near Tarbes, Hautes-Pyrenees, France.

Geographical distribution. — Pyrenees and adjacent portion of
southern Prance ; probably also north-eastern Spain to the
Ebro ; Asturias ?

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-bufi", 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,
p. 27.

Specimens examined. — Fifteen, from the following" localities :■ —
France : As-les-Thermes, Ariege, 5 ; Pyrenees, no exact locality, 8
(B.M. and U.S.N.M.).

Spain : Pajares, Leon, 2.

Sal. Ax-les-Thermes, Ariege, Toulouse Museum 1.7.27.1-2.

720 m. France. (p).

6, ?. Ax-les-Thermes, Ariege. V. Builles (p). 8. 3. 27. 4-5.

2. Pyrenees. (Verreaux.) Tomes Collection. 7. 1. 1. 15-16.

1. Unknown. F. Maxwell Lyte(p). 62.1.13.2.
1 al. Pyrenees. (No history.)

1. Pyrenees. Purchased (Parzu- 41. 918.

daki).

1. Pyrenees. Dr. J. E. Gray (p). 43. 10. 14. 1.

S, ?. Pajares, Leon, Spain. 0. Thomas (p). 8. 2. 9. 47-48.
{N. Gon-alcz.)

Galemys pyeexaicus rupulus Graells.

1897. Myogalea riofula Graells, Mem. Real. Acad. Sci., Madrid, xvii, p. 460.
1910. Myogale pyrenaica riifula Trouessart, Faune Mamm. d'Europe, p. 61.

Tyle locality. — Rio Balsain, above the Venta de los Mosquitos,
Sierra de Guadarrama, Segovia, Spain.

Geograpldcal distrihution. — Central Spain, south of the Ebro
Valley.

Diagnosis. — Hind foot, 36 to 38 mm. ; condylobasal length

27

Observations.

Teeth much worn.
„ moderately worn.
,, not worn.

„ moderately worn.

„ much worn.
,, slightly worn.

,, moderately worn.
„ slightly worn.

•(ajT'^ua)

A\0J-l[100l

a^inqipacj;^

OOOCOOOOOOOO 00 O O-IOl'^fO'^

^COCO^CO'*'*Cr:) CO CO ^^'tl-*^
iHrHi-liHiHrHT-lTH iH iH t-ItHi-Ii— ItH

•(aanua)
Avoj-mooj

XjBIUXBIt

o 00 CM to CO o o o o o o -^ o cq CO

t-COCOCOOt^t^t^ CO t^ oot^oot^t-

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oq(MoqcqcMCMcqcq oi cm oqc^tMoicM

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OCOOOOCOCOCOOC O (M toooo-*

•qipBgjq

CO-^'*-*-*COCOtO 00 1 OtM-^OOOl

IB^iqjoja^ui

COtOCOCOCOtOCOO CO ' cocOOOtO

•(uBipara)

aSBO-UIBjq

JO mdaa

O O CD i£i O ■ O O cq O O
1 1 Kh tH iH -H Al ! ^ Ah ^ W 1-1

JO q,p^8Jg I^I^SSSSS ' ^^^^'

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CO ooocq-^Ttio ^ocooocM

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COCOCOCOCOCO COCOCOCOCO

1

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CO -^

Number.

43.10.14.1

41.918

62. 1. 13. 2

1. 7. 27. 1

1.7.27.2

8. 3. 27. 4

8.3.27.5

152522

i 9099 "l

\ Merriam /

8. 2. 9. 47

8.8.4.44
8. 8. 4. 43
172120
8.7.7.8
8.7.7.9

Locality.

G. pyrenaicus pyrenaicus.

France: no exact locality

Ax-les-Thermes, Ariege

Pyrenees .
Spain : Pajares, Leon

G. pyrenaicus rufulus.

Spain : Silos, Burgos

28 INSECTIVORA

of skvill about 34*5 to 35*5 mm. ; colour apparently not so dark
as in the Pyrenean race.

Colour. — Upper parts essentially as in G. j^yrenaicus 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 foi*m.

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.).

Hemarlis. — The central Spanish form of Galemys appears to
be well ditt'erentiated from true pyrenaicus by its greater size,
a character which is particularly noticeable in the larger,
more massive skull. Two specimens from the Asturias (IS^os. 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 SORICIDtE.

1821. Soricidai Gray, London Med. Eepos., 5V, p. 300, April 1, 1821.

Geographical distribution. — Throughout tropical and temperate
Africa, Europe, Asia (including the Malay Archipelago), North
America, and the extreme northern portion of South America.

Characters. — Skull long and narrow, strongly tapering an-
teriorly, most of the sutures disappearing early in life ; zygomatic
arch incomplete, represented by a slight though usually evident
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

SOREX 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.

RemarJcs. — The members of the family Soricidse are at once
recognizable among European Insectivora by their mouse-like
form, small eyes, and sharply pointed mvizzle. They are all of
small size, the largest (Neomys fodiens) not so large as a house-
mouse, while the smallest [Pachym-a etrusca) is one of the least
of known mammals. Notwithstanding their manifestly primitive
general structure, the Soriei'lse present a very high degree of
specialization in the form of the anterior teeth, the absence of
the zygoma, and the remarkable double articulation of the jaw.
About fifteen genera are known. Four of these are represented
in Europe.

KEY TO THE EUROPEAN GENERA OF SORICID^^.

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 Sorcx, 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 wliite
throughout ; tail with noticeable sprinkling of
elongated hairs.

Upper unicuspid teeth 4-4 Pacliyura, p. 81.

Upper unicuspid teeth 3-3 Crocidura, p. 86.

Genus SOREX Linnteus.

1758. Sorex Linnaeus, Syst. Nat., i, 10th ed., p. 53.

1829. Oxyrhin Kaup, Eutw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,

p. 119 (Included the undeterminable Sorcx constrictas Hermann

and S. tetragonurus Hermann ; the latter may be chosen as type).
1835. Amfhisorcx Duvernoy, M6m. Soc. du Mus. d'Hist. Nat., Strasbourg,

II, p. 23 (hermanni = Neomys fodiens skull -f- Sorex araneus

tetragonicrus animal) Part.
1838. Ccyrsira Gray, Proc. Zool. Soc. Loudon, 1837, p. 123. June 14, 1837

{vulgaris = araneus).
1842. Otisorcx De Kay, Zool. of New York, i, Mamm., p. 22 (platyrhinits

= iicrsonatus).
1857. So7-ex Blasius, Saugethiere Deutschlands, p. 124.
1890. Homalurus Schulze, Schriften Naturwiss. Vereins Harzes in Werni-

gerode, v, p. 28 {nlpimis).

Type species. — Sorex araneus Ijinn.ieus.

Geographical distribution. — Northern portion of both hemi-
spheres ; in Europe west to Ireland and south to central Spain
and southern Italy.

30

INSECTIVORA

Characters. — Upper unicuspid teeth 5-5 (deutal formula :
' 1^1' ^i^» i""f=ij ''*S ~ ^^) ' 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 ; jjoints 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.

Hemarhs. — 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 FORMS OF 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 ?»' and m" ; tail about
as long as head and body ; colour uniform dark

slaty grey. (Alpine Shrews) S. aljnnus, 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. hercynicus, 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 /»' and m" ; tail shorter than
head and body ; colour never uniform dark slaty
grey, the under parts usually much lighter than
back.
First, second and third upper uuicuspids sub-equal ;
condylobasal length of skull 14 "8 to 16 '6 mm.;
head and body usually about 50 to 60 mm.

(Pigmy Shrews) S. nmiittus, p. 53.

Molars and anterior upper incisor normal (Dis-
tribution general) S. m. minutux, p. 55.

Molars and anterior upper incisor enlarged

(Southern Italy) 5. m. lucanhos, p. 60.

First and second upper unicuspid much larger than
third; condylobasal length of skull 17-4 to
20 mm. ; head and body usually about 65 to
80 mm. (Common Shrews) S. aranetis, -p. Bl.

SOREX 31

Palate broad anteriorly, its width at level of first
unicuspid nearly equal to that of tooth.
Condylobasal length of skull 18 "4 to 19 "2 mm.;

anterior teeth enlarged (Island of Jersey)... S. a. fretalis, p. 45.
Coudylobasal length of skull about 17 '5 mm. ;
anterior teeth not enlarged (Mountains of

central Spain) S. a. granaruis, p. 52.

Palate narrow anteriorly, its width at level of
first unicuspid barely more than half that
of tooth.
Condylobasal length of skull usually 19 to 20
mm.; hind foot usually 13 to 14-4 mm.
(mountain and northern forms).
Hind foot 1-3 '6 to 14*4 mm.; colour in
summer pelage very dark, the tricolor
pattern usually conspicuous (South-
western Norway) S. a. bergensis, p. 41.

Hind foot 13 • 0 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. 4:2.

Back rarely if ever blackish in summer

pelage (Pyrenees) S. a. pyrenaicus, p. 44.

Condylobasal length of skull usually 17 "4 to
19 mm. ; hind foot usually 11 to 13 mm.
(lowland forms).
Underparts blackish, not contrasted with

back (Charente, Prance) S. a. 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 France) 5. 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 Eucope and Scan-
dinavia, except south - western

Norway) s. a. araneus, p. 35.

Average colour less dark, the back
ranging from hair-brown tinged
with bister to seal-brown (Great
Britain) S. a. castaneus, p. 37.

SOREX ARANEUS Linnasus.

(Synonymy under subspecies.)

Geographical distribution. — Northern portions of Europe and.
Asia from England and Scotland eastward ; exact limits of
range unknown ; in western Europe south to central Spain, and
central Italy, north to northern Scandinavia.

Dlafjnosis. — Size medium, head and body usually about
65-80 mm, the tail 20-35 mm. shorter, condylobasal length of
skull, 17 -8-20 mm.; posterior lobe of anterior upper incisor

32 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 tine 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 posteriori)-,
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 :
«• 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 veiy 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 sufiYised with slaty (particularly
in the dark S. a. santonus). Between colour of back and sides
there is usualh' 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 invaiiable 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.

SJciill. — 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- „ """' ' , ,

orbital region (more marked than m Sorex Nat. size.

minutus and *S'. alpinm). 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 ni^. 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

34

INSECTIVOKA

Q^===^^^^>^^^*^^^

Sorex aratuu-s. 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

i^'-

Fig. 9.
Sorex araneus. I'eetli X 10.

SOREX 35

low. Hypocone small but well developed. Third uj^per 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.

3Iensure)itenis. — 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 subsj^ecies. >

Bemarhs. — Sorcx 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 aljjinus, 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 j^repared 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 Linna3US.

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 jpersonatus
I. Geoffroy, 1827.

1832. Sorex concinnus Wagler, Isis, p. 54 (Bavaria).

1832. Sorex rhinolopJms Wagler, Isis, p. 54 (Bavaria).

1832. Sorex melanodon Wagler, Isis, p. 54 (Bavaria).

1838. Sorex vulgaris Nathusius, Wiegmann's Archiv fiir Naturgesch.

IV, I, p. 45.

1839. S[orex] macrotrichus de S61ys-Longchamps, Etudes de ]Micromamm.,

p. 20. (Specimen of S. araneus briefly described as agreeing with

the <S. inucrotrichus Mehlis MSS. No locality given.)
1839. S[orex'] labiosus Jenyns, Ann. Nat. Hist., ii, p. 326, January, 1839.

(Frankfurt a/M., Germany.)
1857. Sorex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part).
1895. Sorex araneus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,

1895.
1910. Sorex araneus Trouessart, Faune Mamm. d'Europe, p. 51 (part).

Tijpe locality. — Upsala, Sweden.

Geographical distribution. — Western Continental Europe,

D 2

36 INSECTIVORA

except Atlantic waterslied 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 in^ and »i- 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 difi'erences
are best seen in the hypocones of the three large upper cheek
teeth and protocone of m^, as these small cusps, unlike the larger
ones, may be completely without brown colour. A comparison
of seventy-five topotypes of Sorex araneus with eighty Swiss
specimens of S. araneus tetragonurus gives the following results : —

araneus. tetrofjonurus.

Large premolar with pigment on hypocone . '0% ... 38"7 %

First molar with pigment on hypocone . . 22*6 % ... 93 '7 %

Second molar with pigment on hypocone . 21"2% ... 8S"7 %

Third molar with pigment on protocone . 45"3 % ... 100 "0 %

None of the small cusps pigmented . . 54'6 % ... "0 %

All of the small cusps pigmented . . . '0% ... 37*2%

Measurements. — Average and extremes of twenty specimens
from the type locality : head and body, 77 • 5 (72-85) ; tail, 40* 1
(38-43); hind foot,' 12-6 (12-13). "^ Average and extremes of
nine specimens from Lillehammer, central Gudbrandsdal, ISTorway
(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 (36-4.5); 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,
Telemarkeu, 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: Hillerod, Zealand, b; Nvstad, 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.) ; Bahreuberg, Harz
Mts., 9 (U.S.N.M.) ; Tharand, Saxony, 1 ; Magdeburg, Saxony, 5 ; Moritz-
burg, Saxony, 1 (U.S.N.M.); lugelbeim, llbeinhessen, 3; Nuremberg,
Bavaria, 10 (U.S.N.M.); Marxbeim, near IMoubeim, Bavaria, 15; Strass,
near Burgheim, Bavaria, 1 ; Euleugrund, Riesengebirge, Silesia, 2
(U.S.N.M.) ; Wolfshau, near Sneekoppe,iiiesengebirge, Silesia, 8 (U.S.N.M.);
Niesky, Silesia, 5 ; near Konigsberg, 6 (U.S.N.M.) ; no exact locality, 1.

AusTRiA-HuNGAEY : Haida, Arva, Bobemia, 9.

S,9.

jMolmen, Gudbrandsdal.

R. J. Cuninghame

98. 5. 2. 1-2.

Norway.

{^)-

2 6,2 9.

Holaaker, Gudbrands-

R. J. Cuninghame

98. 2. 28. 1-4.

dal, 1900 ft.

(P).

6.

Lesjevark, Gudbrands-
dal.

Miller Collection.

7. 7. 7. 4452.

i, 5 9.

Holme, Mandal, 200 ft.

R. J. Cuninghame

8. 8. 9. 1-6.

6a\.

Norway.

(P)-

8. 8. 9. 40.

9.

Upland, Sweden.
{G. Kollhoff.)

Lord Lilford (p).

8. 9. 8. 19.

4c5, 2 9.

Hillerod, Zealand, 10 m.
Denmark.

0. Thomas (c & p).

98. 6. 7. 2-7.

4 6, 2 9.

Oosterbeek, Guelderland,
10-15 m. Holland.

0. Thomas (c & p).

98. 2. 1. 6-8.

9.

Hastiere, Namur, Bel-

G. A. Boulenger

94. 7. 9. 1.

gium.

(c & P).

3 6, 1 9.

Guines, Pas-de-Calais,
10 m. France.

0. Thomas (c & p).

94. 6. 6. 4-7.

2 J, 2 9.

Barbizon, Seine - et-
Marne.

G. S. Miller (c).

8. 8. 4. 155-158,

6, 3 9, 2.

Auerum Forest, Bruns-

G. Barrett-Hamil-

11. 1. 2. 75-80.

wick, Germany.

ton (c & p).

9.

Tharandt, Saxony.

Lord Lilford (p).

99. 1. 9. 14.

5 al.

Magdeburg, Saxony.

Dr. W. Wolterstorff

92. 12. 1. 3-7.

6, 2 9.

Ingelbeim, Rbeinhessen.

C. Hilgert (c).

8. 11. 2. 6-S.

3 J, 4 9.

Marxbeim, Bavaria.

Lord Lilford (p).

«. 9. 8. 7-13.

2 6.

Bayreuth, Bavaria.

Miller Collection.

7. 7. 7. 2368-
2869.

6.

Strass, Burgheim, Ba-
varia.

Lord Lilford (p).

8. 9. 8. 20.

3(5,2 9.

Niesky, Silesia, 200 m.
[W. Baer.)

Lord Lilford (p).

99. 1. 9. 9-13.

2al.

No exact locality.

Zool. Soc. Collec-

55. 12. 26. 300-

tion.

301.

1.

Germany.

Stockholm Museum

46. 6. 2. 36.

3 6, 4 9.

Haida, Bohemia.

Lord Lilford (p).

8. 9. 8. 14-18.

3.

Haida, Bohemia.

Lord Lilford (p).

8. 9. 8. 21-22.

SOREX ARAXEUS CASTANEUS Jeiiyn.S.

1838. S\prex] tetragonurus var. ;8 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. Swex araneus Trouessart, Fauna 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 frequently
seen in the typical race.

Shdl and teeth. — The skull and teeth resemble those of true
Sore.i- 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) ; Cromlis,
Stirling, 10; Islay, 3; Dunkeld, Perthshire, 2; Loch Earn Head, Perth-
shire, 1 ; Stockbriggs, Lanarksliire, 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-
Tjme, Northumberland, 3 ; Newby Bridge, Lake Windermere, Cumber-
land, 1 ; Grimsby, Lincolnshire, 17 ; Whitnash, Warwickshire, 1 ; Rugby,
Warwickshire, 2; Filey,- Yorkshire, 3; Wellersey Hill, Broadway, Wor-
cestershire, 1 ; West Cheshire, 1 ; Shropshire, 1 ; Staffordshire, 1 ; Swith-
land, Leicestershire, 10 ; Bishopstoke, Herefordshire, 1 ; Leominster, Here-
fordshire, 1 ; Graftonbury, Herefordshire, 19 ; Lilford, Northamptonshire,
2 ; Drinkstone Park, Bury St. Edmunds, Suffolk, 2 ; Lowestoft, Suffolk, 6 ;
Wormsley, Oxfordshire, 1 ; Stokenchurch, Oxfordshire, 4 ; Cambridge-
shire, 1 ; Kensington Gardens, London, 1 ; Hillingdon, Middlesex, 3 ;
Bletchingley, Surrey, 4 ; Godalming, Surrey, 2 ; Merstham, Surrey, 1 ;
Richmond Park, Surrey, 1; Crowborough, Sussex, 3; St. Leonard's,
Sussex, 1 ; Tunbridge Wells, Sussex, 1 ; Eastwell, Kent, 3 ; Lyndhurst
Road, Hampshire, 2 ; New Forest, Hampshire, 17 ; Basingstoke, Hamp-
shire, 1 ; Alum Bay, Isle of Wight, 3 ; Clifton Bridge, Gloucestei-shire, 2 ;
Leigh Woods, Clifton, Gloucestershire, 3 ; Blandford, Dorsetshire, 1 ;
Combmartin, Devonshire, 4 ; Chagford, Devonshire, 5 ; Northlew, Devon-
shire, 6 ; no exact locality, 3.

Remarks. — While the British common shrew is an incom-
pletely differentiated form, the average characters of the large
series of specimens examined seem important enough to warrant
the use of Jenyn's name. As compared with the Continental

SOREX 39

race the dark extreme is less dark and less freqvient, while the
light extreme is more light and more frequent. The constantly
small size of this animal as compared with the large Continental

races (hergensis, tetragonnrus 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.

?. Black Isle, Cromarty, W.R. Ogilvie-Grant 94.10.6.1.
Scotland. (c & p).

9 6. Black Isle, Cromarty. W. K Ogilvie-Grant 11. 1. 3. 103-

(c & p). 107. 109-112.

2 6. South Sutor, Cromarty. W.R. Ogilvie-Grant 11. 1. 3. 108,

(c & p). 113.

2 6,2 9. Nairn, Morayshire. W.E. Ogilvie-Grant 11. 1. 3. 99-102.

(c & p).

6. Morayshire. E. R. Alston (c & p). 79. 9. 2.5. 79.

9. Dunphail, Elginshire. W. R. Ogilvie-Grant 11.1.3.97.

(c & P).

1. Lhanbride, Elginshire. Miller Collection. 7. 7. 7. 3598.

2 6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.95-96.

shire. (c & p).

6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11. 1. 3. 98.
shire. (c & p).

3 6,19. Cromlix, Stirlingshire. W. R. Ogilvie-Grant 11.1.3.114-123.

(c & p).

3 al. Islay. H. Russell (c & p). 9. 9. 11. 1-3.

6, 9. Stockbriggs, Lanark- E. R. Alston (c & p). 79. 9. 25. 11-12.

shire.

6,39. Kirtle Bridge, Dumfries- Miss D. Bate (c&p). 11.1.3.88.

shire. 11. 1. 3. 92-94.

3 9. Wyseby, Dumfriesshire. Miss D. Bate (c&p). 11.1.3.89-91.

5 6,39. Aberia, Merionethshire, G. H. Caton Haigh 11. 1. 3. 156-

Wales. (c&p). 163.

3^,3 9. Bridgend, Glamorgan- R. I. Pocock (c& p). 11. 1. 3. 164-

shire. 169.

2 al. Berwick-on-Tweed, -J. H. Fryer (c & p). 47.11.11.6-7.

Northumberland,
England.

1 al. Newby Bridge, Cumber- J. Paul (c & p). 94. 9. 3. 2-3.
land.

6,5 9,2. Grimsby, Lincolnshire. G. H. Caton Haigh 11. 1. 3. 124-

(c & p). 131.

6,6 9. Swithland, Leicester- P. A. Butler (c&p). 11. 1. 3. 132-

shire. 138.

1 al. Shropshire. T. C. Eyton (c & p). 63. 10. 12. 4.
9 al. Bishopstoke, Hereford- S. 0. Ridley (c&p). 84. 10. 6. 1.

shire.

6 6,8 9. Graftoubury, Hereford- W. do Winton 11. 1. 3. 72-87.

shire. (c & p).

3 St. Graftonbury, Hereford- W. de Winton 96. 4. 28. 17-19.

shire. (c&p).

2 al. Bury St. Edmunds, J. H. Powell (c & p). 80. 5. 22. 1-2.

Suffolk.

3<5, 3 9. Lowestoft, Suffolk. 0. Thomas (c & p). 11. 1. 3. 139-

144.

4 9. Stokenchurch, Oxford- W. R. Ogilvie-Grant 11. 1. 3. 145-

shire. (c & p). 148.

3 9. Hillingdon, Middlesex. Miller Collection. 7. 7. 7. 3595-

(0. Thomas.) 3597.

40 INSECTIVOEA

1. Kensington Gardens, N. Churton (c & p). 161. a.

London.

4 9. Bletchingley, Surrej'. W.R. Ogilvie-Graut 11. 1. 3. 149-

(p). 152.

1. Crowborough, Sussex. Miller Collection. 7. 7. 7. 3593.

{W. B. Ogilvie-Giant.)

? al. Richmond Park, Surrey. Prof. Owen (r). 75.9.17.1.

7 6, 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.)

6,29. Alum Bay,Isle of Wight. O. Thomas (c & p). 11.1.8.153-155.

9 (albino) Winscombe, Somerset. F. A. Knight (c. & p). 4. 8. 9. 1.

6, 4 9. Northlew, Devonshire. E. C. Wroughton 11. 1. 3. 170-

(c & p). 174.

2 6,3 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.

SOREX 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, Faune Mamm. d'Europe, p. 54.

Type locality. — Lignieres-Sonneville, Charente, France.

GeograpJdcal distribution. — Known only from the vicinity of _
the type locality.

Diagnosis. — Size as in Sorex araneus nraneus and the other
small races ; colour throughout a nearly uniform dark sooty
brown.

Colour. — Upper parts dark sepia anteriorly, deepening to
blackish postei'iorly, 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.

SJcull and teeth. — The skull and teeth do not difier 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).

RemarJcs. — 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 araneiis euronotus Miller, Proc. Biol. Soc, Washington, siv,
p. 44, April 25, 1901 (Montrejeau, Haute-Garonne, France). Type
in U.S. National Museum.

1910. Sorex arancus 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 arancus 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 arancus bergensis Miller, Ann. and Mag. Nat. Hist., 8th ser.,

Ill, 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 arancus araneus, except in their larger size,
as shown by the detailed measurements. The j^igmentation of

42 INSECTIVORA

the teeth is scarcely, if at all, more extensive than in the typical
i-ace.

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 : Vefseu, 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.).

Remarks. — This large race of Sorex araneus, closely resembling
the large Alpine and Pyrenean forms, but decidedly darker in
colour, appears to be strictly contined 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 \vinter 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,
velvet}^ summer coat.

1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 2.

way. Waldo (c & p).

6, 4 ?. Skjserdal, Nordfjord. G. H. Stephenson 8. 1. 5. 1-5.

(c & p).

2 6,29. Opheim, Bergen. Miller Collection. 7. 7. 7. 4453-

4456.

6, 2 9. Bergen. Miller Collection. 7. 7. 7. 4457-9.

9. Bergen, 2,700 ft. G. Barretfc-Hamil- 8. 9. 21. 1.

ton (p).

SoEEX ARANEUS TETRAGONURUS Hermann.

1780. Sorex tetragomcrtis Hermann in Zimmermann, Geogr. Gesch., ii,
p. 383 (Strassburg, Germany).

1792. Sai-ex quadricaudatiis 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, M6m. 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. Nafcurwiss. Classe, lvii, pt. i, p. 488
(Based on specimen from unknown locality, probably in Italy,
figured by Bonaparte, Iconogr. Faun. Ital., fasc. xxix, pi., fig. 5.)

1869. [/Sorex vulgaris'] var. nuda Fatio, Faune Vert. Suisse, i, p. 127

(Bernese Oberland, Switzerland).

SOREX 43

1869. [Sorex vulgaris'] var. 7iigra Fatio, Faune Vert. Suisse, i, p. 127
(Lucerne, Switzerland).

1900. S[orex:] vulgaris var. vei subsp. mollis Fatio, Rev. 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 ]\Iuseum.

1905. S[orex] vulgaris crassicaudatus Fatio, Arch. Sci. Phys. et Nat.,
Geneve, 4th ser., xix, p. 201, February 15, 1905 (Zermatt, Switzer-
land). Cotypes in Geneva Museum.

1905. Crossoims ou Sorex ignotus Fatio, Arch. Sci. Phys. et Nat., Geneve,

4th ser., xix, p. 202, February 15, 1905 (Switzerland). Part:

mandible, not skull. Type in Geneva Museum.
1905. Sorex araneus carpathicus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., xv, p. 506, May, 1905 (Hatszeg, Hunyad, Hungary).

Type in British Museum.
1910. Sorex araneus tetragonurus and S. araneus carpathicus Trouessart,

Faune Mamm. d'Europe, pp. 52, 54.

Type locality. — Strassburg, Germany.

Geograflikal distribution. — Alps and neighbouring portions of
Germany, France 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^ and m^ usually brown at tip (see tabular
comparison with true araneus on p. 36).

Measurements. — Average and extremes of eight specimens
from the type locality : head and body, 75 '8 (71-79) ; tail, 47-5
(45-50); hind foot, 13-5 (13 •2-14 -2). Average and extremes
of twenty specimens from Andermatt, Uri, Switzerland : head
and 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 oi Sorex
araneus alticola Miller (Meiringen, Switzerland, No. 85,930
U.S.N.M.) : head and body, 76 ; tail, 55 ; hind foot, 14. Average
and extremes of ten specimens from Meiringen, Switzerland : I
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.

* Cotype of Sorex vulgaris crassicaudatus Fatio.
t Essentially topotypes of Sorex vulgaris nigra and mollis Fatio.
X Topotypes of Sorex araneus alticola JMiller and essentially topotypes
of Sorex vulgaris nuda Fatio.

44

IXSECTIVORA

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,
Yaud, 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.il.) ; 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. Galleu, 16 (U.S.N.M.) ; Degersheim, St. Gallen, 3
(U.S.N.M.); Uzwil, St. GaUen, 1 (U.S.N.M.); Sitterwald, St. GaUen, 5
(U.S.N.iM.) ; Ziiberwangen, St. GaUen, 5 (U.S.N.M.) ; Wildkirchli, Appen-
zell, 1 (U.S.N.LI.) ; Albulapass, Grisons, 1 (U.S.N.M.) ; Untervatz, Grisons,
5 (U.S.N.M.) ; Grisons, no exact locality, 4 (U.S.N3I.) ; 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 ; Csallokoz-
Somorja, Pressburg, Hungary, 3 ; Schwaz, Tirol, 2 (U.S.N.M.).

Italy : Near Turin, 4 (Turin) ; Unerzio, Cuneo, 1 ; Vallombrosa, near
Florence, 2.

8(5,2?.

Etupes, Doubs, 350 m.
France. {G. Mottaz).

0. Thomas (p).

8. 8. 10. 14-24.

3(5.

Barcelonnette, Basses-
Alpes. (C. Mottaz.)

0. Thomas (p).

8. 8. 10. 25-27.

4 <5, 1 9.

Strassburg, Alsace.
(C. Mottaz.)

0. Thomas (p).

8. 8. 10. 28-32,

lal.

Zermatt, Valais, Swit-

Dr. J. Anderson

91. 10. 15. 26.

zerland.

(c & P).

3 6.

Vitznau, Lake Lucerne.

0. Thomas (c & p).

5. 8. 2. 32-34.

lal.

Zurich.

C. Mosch (p).

89. 11. 8. 3

?.

Faido, Ticino.

0. Thomas (c & p).

5. 8. 2. 19.

2(5,2 9.

Locarno, Ticino.

0. Thomas (c & p).

5. 8. 2. 2-5.

9.

Switzerland.

E. R. Alston (p).

79. 9. 25. 10.

8 (5, 1 9.

Hatszeg, Hunyad, Tran-

C. G. Danford (c).

3. 2. 2. 3-10.

sylvania, 1500-2000 ft.

3. 11. 8. 15.

Hungary.

3.

Csallokdz-Somorja,

Budapest Museum

94. 3. 1. 22-25.

Pressburg, 400 ft.

(E).

^ Hungary.

2al.

Vallombrosa, Florence,

Dr. G. Cecconi

1. 8. 2. 2-3.

Italy.

(c & P).

SOEEX AEANEUS PYRENAICUS Miller.

1909. Sorex araneus pyrenaicus Miller, Ann. and Mag. Nat. Hist., Sth ser..

Ill, p. 416, May, 1909. Type in British Museum.

1910. Sorex aranetis pyrenaictis 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

SOREX 45

pelage (winter coat not known), in which the back rarely if
ever assumes the blackish-brown tints often seen in the Alpine
form.

Colour. — The colour scarcely needs detailed description. As
compared with that of the other large races it is characterised
by extreme dulness and lack of noticeable contrast between the
dorsal area and sides. In none of the thirty-two skins examined
is the back so dark as in the dark individuals frequently found
in Switzerland in summer.

Skull and teeth. — As in 8. araneus tetragonunis.

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 : —

Port6, Pyren^es-Orientales (Spanish watershed), 9; I'Hospitalet, Ariege,
12 ; Ax-les-Thernies, Ariege, 2 ; Bareges, Hautes-Pyr^nees, 9.

BemarTcs. — ^In its dull colour the Pyrenean shrew differs from
the other large members of the group in much the same way as
the small Sore.v araneus euronotus of the neighbouring lowlands
differs from true araneus. It thus represents the opposite
extreme from the dark Norwegian form.

8, ?. Porte, Pyr6nees-0rien- G. S. Miller (c). 8. S. 4. 142-143.

tales, 1600m. France.
2 6,9. Porte, Pyrenees-Orien- 0. Thomas (p). 8.9.1.49-51.

tales, 1600-1700 m.

[A. Robert).
(5,4?. 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. 4. 141.

2600 ft.
6, 6 ?. Bareges, Hautes-Pyre- G. S. Miller (c). 8. 8. 4. 144-150.

nees, 1300-1500 m.

SoREX ARANEUS FRETALIS Miller.

1909. Sorex araneus fretalis Miller, Ann. and Mag. Nat. Hist., 8th ser., in,

p. 416, May, 1909. Type in British Museum.

1910. Sorex araneus fretalis Trouessart, Fauna Mamm. d'Europe, 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

46

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50

IXSECTIVORA

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S . araneus tetragonurus (cont
Switzerland : Zermatt .

no exact localitj
Austria-Hungary : Schwaz, Ti

Hatszeg, H

S. araneus pyrenaicus

Ax-les-Thermes, Ariege .

I'Hospitalet, Ariege

Porte, Pyr6n6es-Orientales
Bareges, Hautes-Pyren^es

51

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52 IN'SECTIVORA

teeth (incisors both above and below, and first and second uj^per
unicuspids) enlai'ged.

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 cvitting 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.

Remarhs. — 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.

9,6. Jersey, Channel Islands. 0. Thomas (p). 8.9.2.1-2.

(R. H. Bunting.) (S. 9. 2. 1. Tijpe of subspecies.)

Sorex araneus graxarius Miller.

1910. Sorex araneus granarius ]Miller, Ann. and Mag. Nat. Hist., Sth 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 iS'. 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 fi'om 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.

BemarJcs. — 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.dela Escalera(c). 6.11.4.3-4.
Spain. (6. 11. 4. 4. Tijpe of subspecies.)

SOREX MINUTUS Linnfeus.
(Synonymy iinder subspecies.)

Geugmphical 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 1 4 • 8-^1 6 • 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
hiinutus 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 varj'ing depth, some-

54

INSKCTIVORA

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 dai'k
below.

SJcuU. — Apart from its conspicuously smaller size (condylo-
basal length, 14 -8 to 16 mm. instead of 17 '8
to 20 mm.) the skull differs notably from
that of Sorex arancus and S. alpinus in the
narrower, more elongate brain-case, the outline
of which when viewed from above is distinctl}^
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 ofl' from interorbital region than in
Sorex minutus. ^^^® other European species. Depth of brain-case
Nat. size. usually less relatively to breadth than in

S. araneus. Lachiymal 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 ^'*^' ^^'

„ T 1 i 1 • J.1 J.T • -i" Sorex minutus. Anterior

square. In lateral view the outline is quite teeth in profile, x 5.
as in the larger animal. The relative size

of the unicuspids differs markedly from that in the larger
animal. The first, second and third are sub-equal, with the
second usually a trifle smaller than the third, and the first with
distinctly the greatest crown area. Fourth slightly smaller than
third, and fifth equally smaller than fourth'"', between which and
large premolar it is tightly crowded. First lower unicuspid
diflering 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.

SOREX 00

the side thus noticeably different from that of first upper
unicuspid. Second unicuspid, together with other mandibuhxr
teeth, essentially as in S. a ran ens. Maxillary cheek-teeth as in
S. ar emeus, except that hypocones are less developed, that on
large premolar obsolete.

Bemarks. — Sorex minutus is at once distinguishable from
S. araneus by its smaller size and relatively longer tail, as well
as by the more technical characters of the skull and teeth.
Immature individuals of araneus might sometimes be mistaken
for minutus, but their larger feet will serve to indicate their
identity ; while if the skull and teeth can be examined, a positive
identification is easily obtained.

SoREX MINUTUS MINUTUS Linnseus.

1766. [Sorex] minutus Linnaeus, Syst. Nat., i, 12th ed., p. 73 (Siberia).
1769. Sorex pygmastis Laxmann, Sibirische Briefe, p. 72 (Barnaul, Tomsk,

Siberia).
1789. [Sorex] exilis Gmelin, Syst. Nat., i, 13th ed., p. 115 (Yenesei River,

Siberia).
1806. Sorex canaliculatus Ljungh, Koiigl. Vetensk. Akad. Nya Handl.,

xxvii, p. 263 (Lommaryd Vicarage, northern Vedbo district,

Jornkoping, Sweden).
1811. Sorex jjygmains Pallas, Zoogr. Rosso-Asiat., i, p. 134 (Ob and Yenesei

Rivers, Siberia).
1811. Sorex minimus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 186

(Accidental renaming of minutus).
1832. Sorex lyumilio Wagler, Isis, p. 54 (Bavaria).
1838. S[ore.v] rusticus Jenyns, Ann. Nat. Hist., i, p. 423, August, 1838

(England).
1838. S[o7-ex] rusticus var. B S[orex] hibernicus Jenyns, Ann. Nat. Hist., i,

p. 423, August, 18.38 (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 pygmxus Blasius, Saugethiere Deutschlands, p. 133.
1895. Sorex minutus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,

1895.
1910. Sorex minutus Trouessart, Faune Mamm. d'Europe, p. 55.

Tijije locality. — Vicinity of the Yenesei River, Siberia.

Geographical distribution. — The entire Eurojaean 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
(49-5.5); 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*0 (10*8-11 '2). Average and extremes of seven
specimens from Hatszeg, Hunyad, Transylvania : head and bodv,
51-3 (47-55); tail, 40-6 (32-46); hind foot, 11-2 (10-12).
For cranial measurements see Table, p. 58.

Specimens examined. — Oue hundred and twenty-five, from the following
localities : —

Scotland : Lossiemouth, Elgin, 2 ; Gordonstown, Elgin, 1 ; Lhanbryde,
Elgin, 2; Grantowu-on-Spey, Elgin, 13; Cromlix, Dunblane, 1; Aber-
deen, 1; Dunvegan, Skye, 1; Stornoway, Lewis, Hebrides, 1; Newton,
North Uist, Hebrides, 1 ; Barra Island, Hebrides, 1 ; Stockbriggs, Lanark-
shire, 1; Kirtle Bridge, Dumfriesshire, 1; Wyseby, Dumfriesshire, 2.

Wales : Aberia, Merionethshire, 2 ; near Bridgend, Glamorganshire, 1 ;
no exact locality, 1.

England : Spurn Head, Yorkshire, 1 ; Grimsby, Lincolnshire, 3 ;
Waltham, Lincolnshire, 2 ; Thornhaugh, Nortbants, 2 ; Swithland, Leices-
tershire, 1 ; Graftoubury, 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, I
(U.S.N.M.).

France: Barbizon, Seine-et-I\Iarne, 1; Montr^jeau, Haute-Garonne, 1
(U.S.N.M.); Ax-les-Thermes, Ariege, 2; I'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 Konigsberg, 2 (U.S.N.M.).

Austria-Hungary: Csallokoz-Somorja, Pressburg, Hungary, 1 ; Hatszeg,
Hunyad, Transylvania, 7.

Switzerland : St. Cergues, Vaud, 2 (U.S.N.M.) ; Stalden, Valais, 4
(Geneva) ; Untervatz, Grisons, 5 (U.S.N.M. and Mottaz) ; St. Gothard,
Uri, 1.

Italy: YaUombrosa, near Florence, 1.

1. Lossiemouth, Moray- G. Denson (c & p). 8. 9. 17. 1.
shire, Scotland.
?, 1. Lhanbrvde, Elgin, 200 ft. W\ Taylor (c & p). 8. 9. 3. 1-2.

?. Dunblane, Stirlingshire. W.R. Ogilvie-Grant 8.9.6.1.

(c & p).
1. Aberdeen. Dr. Macgillivray 52. 7. 10. 13.

(c & p).
6. Dunvegan, Skye. J. S. Elliott (c & p). 8. 9. 4. 1.

6. Stornoway, Lewis. R. M. D. Hawker 8. 9. 18. 1.

(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

6. Stockbriggs, Lanark-
shire.

1. Bridgend, Glamorgan-

shire, Wales.

1 al. Wales.

9. Grimsby, Lincolnshire,

England.

2. Waltham, Lincolnshire.

2 st. Graftonbury, Hereford-
shire.

?. Wilbraham, Cambridge-

shire. (S.F.Harmer.)

1. Clifton, Gloucestershire.
(R. I. Pocock.)

6. Lundy Island, Devon-
shire.

2 6. Chagford, Devonshire,

856 ft. (W.P.Stark.)
6, 4 9. Ramsay, Isle of Man.

6. Sulby Glen, Isle of Man.

9. Clonbroch, Galway, Ire-

land.

1 al. Longford.

6 St. Knock, Down.

2 al. Dunkettle, Cork.

2 c5, 1 9. Duncannon, Wexford.
(Rev. Dr. Martin.)

1 al. Ireland.

6. Holaaker, Gudbrandsdal,
1900 ft. Norway.

9. Barbizon, Seine-et-
Marne, France.

2 9. Ax-les-Thermes, Ariege.
6. r Hospitalet, Ariege,

4700 m.
6. V Hospitalet, Ariege,

1450 m. (A. Robert.)
9. Port6, Pyrenees-Orien-

tales, 1600 m.

2. Quorum Forest, Bruns-

wick, Germany.
1 al. Frankfort - on - INIaiu,
Nassau.

9. Niesky, Silesia, 181 m.

(W. Bacr.)

6. ]\Iarxheim, Bavaria,

1. Csallokciz-Somorja, Press-

burg, 400 ft. Hungary.

1. Platszeg, Hunvad, Tran-
sylvania, " 2000-
5500 ft.

3. Hatszeg, Transylvania,

2000-5500 ft.

3 6. Hatszeg, Hunyad, 1500-

2000 ft.

E.R. Alston (c&p).

R. I. Pocock (c & p).

G. Stokes (c & p).
G. H. Caton Haigh

(c & p).
G. H. Caton Haigh

(c & p).
W. de W i n t 0 n

(c & p).
Miller Collection.

]\Iiller 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).

;\Iiss O'Keefe (c&p).

G. Barrett-Hamilton

(P)-.
Zoological Society's

Collection.
R. J. Cuuinghame

(c & p).
G. S. Miller (c).

V. Bullies (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 IMuseum

(E).

C. G. Danford (c).

C. G. Danford (c).
C. G. Danford (c).

79. 9. 25. 18.

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. 18. 288.

98. 2. 28. 5.

8. 8. 4. 160.

8. 3. 27. 2-8.
8. 8. 4. 159.

8. 9. 1. 45.

8. 8. 4. 161.

11. 1. 2. 86-87.

7. 1. 1. 46.

99. 1. 9. 15.

11. 1. 1. 146.
94. 3. 1. 25.

3. 2. 2. 11.

3. 2. 2. 13-15.
3. 11. 8. 16-18.

58

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60 INSECTIVOKA

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, Ann. and Mag. Nat. Hist., Sth ser.,

Ill, p. 417, May, 1909. Type in British Museum.

1910. Sorcx minutus lucanius Trouessart, Paune Mamm. d'Europe, p. 55.

Type locality. — -Monte Sirino, Lagonegro, Italy.

Geographical distribution. — At present known from the type
locality only.

Diacjuosis. — Similar to Sorex minutus minutus, but with molars
and anterior upper incisor noticeabl}' 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, 0. 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,
Riesengebirge 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

SOREX 61

scantily haired elsewhere, a variation not wholly dependent on
season.* At middle of tail there are about 18 annulations to the
centimeter. Feet relatively larger than in Sorex araneus, but
not i^eculiar 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 sux-faces not conspicuous, and sides
entirely without line of demarcation. Feet scantily clothed with
silvery hairs. Tail sharjjly 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 sorexaiphms.

first and second molars. i^at. size.

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 INSECTIVORA

are all longer in proportion to their height than in Sorex
araneus, and the posterior border is more concave. In size there
is a f'radual and regular diminution from first unicuspid to third ;
fourth somewhat more abruptly smaller ; '"' fifth slightly smaller
than fourth when viewed from the side, but with larger crown
area due to the presence of a well developed postero-internal
crushing surface. This tooth is rela-
tively larger and more functional than
in Sorex araneus, showing no tendency
to become subordinate to paracone of
large premolar. Lower unicuspids
narrower and more trenchant than in
Sorex araneus. Except for this general
tendency the second shows no special
^_^^' ^^' . peculiarities of form. The first, how-

"'^^teeth.'* X 5" '^''° ever, is strikingly diiferent 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. Schweiz. Gesellsch.

Naturwiss., Neuchatel, i, p. 13 (St. Gothard Pass, Switzerland).
1840. ? Sorex anthiorii 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 intermedius Cornalia, Catal. Descrit. Mamm. Ital., p. 27 (Hills

of Brianza, Como, Italv). Part : body ; see Sordelli, Atti Soc. Ital.

Sci. Nat. e del Mus. Civ. Stor. Nat., Milano, xxxviii, p. 364, 1899.
1899. ? Sorex alpinus var. longobarda Sordelli, Atti Soc. Ital. Sci. Nat. e del

Mus. Civ. Stor. Nat., Milano, xxxviii, p. 363 (MS. synonym of

interynedius) .
1910. Sorex alpinus Trouessarfc, 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, xii, 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.

SOREX 63

Diagnosis. — Skull and teeth of maximum size for the sjDecies
(condylobasal length of skull, 19*4 to 20' 6; upper tooth-row,
8 • 6 to 9 • 0 ; lower tooth-row, 8 • 0 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 A^itznau, Switzerland : head and body, 72 "3 (69-75) ;
tail, 74-0 (72-75); hind foot, 15-5 (15-16). Adult male from
Vitznau, Switzerland : head and body, 69 ; tail, 68 ; hind foot,
15 '3. Adult female from Chamonix, Haute-Savoie, France:
head and body, 73; tail, 74; hind foot, 15 "4. For cranial
measurements see Table, p. 64.

Specimens examined. — Thirty-one, from the following localities: —

France: Chamonix, Haute-Savoie, 1 (U.S.N.M.).

Switzerland : St. Cergues, Vaud (Jura), 4 (U.S.N.M. and Mottaz) ;
Briinig, 5 (U.S.N.M.); Vitznau, 4 (B.M. and U.S.N.M.); Audermatt, 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.) ; Siintis, St. Gallen, 1
(Mottaz) ; no exact locality, 3.

Austria-Hungary : Hatszeg, Hunyad, Transylvania, 4.

6. Vitznau, 440 m. Swit- O. Thomas (c & p). 5. 8. 3. 15.

zerland.
1. St. Gothard, Uri. Baron E. de Selys- 45. 7. 5. 1.

Longchanips (p).
1, 1 al. St. Gothard, Uri. Purchased (Brandt). 46. 2. 1.3. 13.

46. 2. 13. 14.
1. St. Gothard, Uri. Purchased (Stock- 46. 6. 2. 82.

holm Museum).
1. Switzerland. E. R. Alston (p). 79. 9. 25. 9.

1. Switzerland. Purchased (Ger- 76. 9. 18. 13.

rard).
9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 2. 2. 2.

sylvania, 2000 ft.
Hungary.
6, 2 ?. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 12-14.

sylvania, 1500-2000 ft.

SoREX ALPINUS HERCYNICUS Miller.

1909. Sorcx alpimis liercynicns Miller, Ann. and Mag. Nat. Hist., 8th ser..

Ill, p. 417, May, 1909. Type in U.S. National Museum.

1910. Sorcx alpinus hercynicus Trouessart, Faune Mamm. d'Europe, p. 51.

Type locality. — Mauseklippe, Bode Valley, Harz Mountains,
Germany.

Geographical distribution. — Harz Mountains and Riesen-
gebirge.

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

INSECTIVORA

Observations.

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,, not worn.

,, slightly worn.
,, uot woru.

,, slightly worn.
,, not woru.

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,, not worn.

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S. alpinus alpinus.

France : (Jhamonix
Switzerland : St. Cergues, Vai

Vitznau .
]->riinig

Audcrmatt

S. alpinus hercynicus.

Germany : Wolfshau, Silesia

Eulcugrund .

Bahrenherg, Harz
Miiuscklippc ,

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, Riesengebirge, Silesia, 2.

Rcraarlcs. — 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. Keomys Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Tbierwelt, i,

p. 117 {Sorex daubentonii Erxleben).
1829. Leucorhynchus Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Tbier-
welt, I, p. 117 {Sorex lineatus Geoffroy and S. Icucodon Hermann;

the first chosen as type by Thomas, The Zoologist, 4th ser., ii,

p. 102, March, 1898).
1829. Hydrogale Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Tbierwelt, 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. l7).
1835. Amphisorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg,

II, p. 28 {hermanni = fodiens skull + Sorex araneus tetragonurus

animal). Part.
1838. Pinalia Gray, Proc. Zool. Soc, Loudon (1837), p. 12G, June 14, 1838

(MS. synonym of Crossopjis).
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. Myosicfis Pomel, Catal. Meth. Vert. Foss. 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., ii, p. 100, March 15, 1893.

Type species. — Sorex daubentonii 'Evxlehen = S. fodiens Schreber.

Geographical distribution. — Pala^arctic 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 j^re-
hensory than in ^S'orea;; dental formula : i^^, ("j-Ejji"" |Ei> ^"^3^3— '^^'>
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 INSKCTIVOEA

the liind feet enlarged and turned somewhat outward ; in one
species a longitudinal ridge of stiti'ened hairs along under surface
of tail ; ear small, completely hidden in the fur, the meatus
closed by two valves, one of which lies on inner surface of
antitragus, the other on inner surface of conch ; habits aquatic.

Bemarks. — 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 FORjMS OP NEOMYS.

Tail with median keel of stiffened hairs extending

entire leDgth of under surface N. fodiens, p. 66.

Underparts whitish or yellowish, occasionally
brownish (Continental Europe except Iberian

Peninsula) X. f. fodien.i, p. 69.

Underparts usually brownish (England and Scot-
land) -V./. 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.)

Geograpldcal distribution. — From Norway and England to the
Pyrenees, northern Ital}^ 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 1 7 mm. ; lachry-
mal foramen opening over posterior half of m^ ; anterior upper
incisor robust.

E.dernal characters. — Fur very dense, its depth at middle of
back about 6 mm. in summer, 9 mm. in winter, its textui'e 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 aranens, 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. Mamrn* : 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 suftused with buffy, 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 oif 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 ^, ' "

. , , ,, ■ .1 !-■ ■ £ ^ eomys fodiens.

Widely open than m the iiuropean species or jjat. size.

Sorex. Lachrymal foramen over posterior
half of m^. ]VIesopterygoid space decidedly more than half as
wide as long ; hamular processes shorter than in Sorex araneus

68 INSECTIVORA

and more abruptly turned outward. Foramen ovale appearing
rather conspicuously in region between hamular and postglenoid
process. Rudimentary zygomatic process of maxillary well
developed, essentially as in Sorex araneus.

Teeth. — Though in general not strikingly different from those
of Sorex araneus the teeth show a tendency towards greater
elevation of the cusps as well as a more pronounced hooking
forward of the anterior upper incisor, peculiarities which impart
to the entire dentition, though 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 onh^ 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
3com?/s,^oyM7s.^ iiteiioi ee i between tips of unicuspids greater

than in the European species of
Sorex. Each cusp is nearly terete, but with a small postero-
intei'nal 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.

HemarJcs. — 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. AVhile 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. Sorcx aquaticus P. L. S. Milller, Natursyst. Suppl. u. Regist.-Band,

p. 36 (France ; based on Buffon, viii, pi. 11). Not Sorcx aquaticus
Linnpeus, 1758.

1777. Sorex fodiens Schreber, Saugthiere, iii, 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 I'Acad. Roy. des Sci., Paris, 1756, p. 42).
1780. Sorex carinatus Hermann in Zimmermann, Geogr. Gesch., ii, p. 382
(Strassburg, Germany).

1792. Sorex liricaudatus Kerr, Anim. Kingd., p. 208 (Strassburg, Germany ;

based on Pennant's account of Sorex carinatus Hermann).

1793. [Sorer] fluviatilis Bechstein, Gemeinn. Naturgesch. Deutschlands,

III, p. 746 (Suggested but not adopted as preferable to fodiens).
1793. Sorex eremita Meyer, Zool. Annalen, i, p. 823 (Thiiringen, Germany).
1793. Sorex fluviatilis Meyer, Zool. Annalen, i, p. 323 (Published as a

synomym of Sorex eremita, perhaps from Bechstein ]\IS.).
1800. Slorex] f[odie7is'] albus Bechstein, Thomas Pennant's Allgem. Uebers.

vierfiiss. Thiere, ii, p. 723.
1811. Sorex hydrophilus Pallas, Zoogr. Rosso-Asiat., p. 130 (Berlin,

Germany).
1811. Sorex lineatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 181

(Paris, France). Type in Paris Museum.
1811. Sorex remifer Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xvii, p. 182

(Abbeville, Somme, France). Type in Paris Museum.
1818. Sorcx collaris Desmarest, Nouv. Diet. d'Hist. Nat., xxii, p. 65

(Holland : islands at mouth of Escaut and Meuse). Described

but not named bv Geoffroy, M6m. 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 INSECTIVORA

1822. Sonw macrourns Lchmann, Observ. Zoologicie in faunam Hamburg-
ensem, i, p. 5 (Sachsenwald, near Friedrichsruh, Schleswig-Holstein,
Germany).

1826. So)-ex amphibitcs Brehni, Ornis, ii, p. 44 (Reuthendorf, Thiiringen,
Germany).

1826. Surex natans Brehm, Ornis, ii, p. 44 (Renthendorf, Thiiringen,
Germany).

1826. Sorcx stagnatilis Brehm, Ornis, ii, p. 47 (Renthendorf, Thiiringen,

Germany) .
1830. Sorex rivalis Brehm, Isis, p. 1128 (Renthendorf, Thiiringen, Germany).
1832. Sorex vntsculus Wagler, Isis, j). 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. Sorcx fodieiis Duvernoy, Mem. Soc. du JNIus. d'Hist. Nat., Strasbourg,

II, p. 17. Part : animal, not skull.
1835. Sorex hermanni Duvernoy, I\Iem. Soc. du Tilus. d'Hist. Nat., Stras-
bourg, II, p. 23 (Strassburg, Germany). Part : skull, not animal.

1838. Ampliisorex linneana Gray, Ann. Nat. Hist., ii, p. 287, December,

1838 (North Bothnia, Sweden).

1839. \ Sorex fodiens] var. leucotis de Selys-Lougchamps, Etudes de ]\Iicro-

mamm., p. 142 (described on p. 25) (St. Gervais, at foot of Mt.

Blanc, Haute-Savoie, France).
1839. ? [Sorcx fodiens] var. albivcntris de Selys-Lougchamps, Etudes de

Micromamm., p. 142 (nomen nudum).
1845. ? [Sorcx fodiens'] var. nigricans Nilsson, Atti della sesta Eiunione

degli Sciendati Italiani, Torino, 1844, p. 357 (Sweden). Nomen

nudum.

1857. Crossopus fodiens Blasius, Saugethiere Deutschlands, p. 120 (part).

1868. Sorcx fimhriatiis and Crossopus fimhriatns Fitzinger, Sitzungsber.
kais. Akad. Wissensch. Wien, llath.-Naturwiss. Clasre, lvii, pt. 1,
p. 610 (Synonyms of " Crossopus daubentonii nigripes,'" wrongly
attributed to Giebel, Siiugethiere, p. 899). Not Sorex Hmhriatus
Wagler, 1832.

1868. Crossopns ciliatus, griscogiilaris Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvii, pt. 1, p. 623

(Chartres, Eure-et-Loire, France).
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, xxxviii,

p. 364, 1899).

1895. Neomys fodiens Thomas, The Zoologist, 4th ser., ii, p. 100, ]March,
1908.

1899. Sorcx alpiniis var. longobardus Sordelli, Atti Soc. Ital. Sci. Nat. e

del Mus. Civ. Stor. Nat., Milano, xxxviii, p. 363 (MS. synonym

of intcrmedixts).
1901. Neomys fodiens minor Miller, Proc. Biol. Soc, Washington, xiv,

p. 45, April 25, 1901 (Montr^jeau, Haute-Garonne, France). Type

in U.S. National Museum.
1905. Crossop)us ou Sorex ignotus Fatio, 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). Tj-pe

in British Museum.

NEOMYS 71

190G. Ncomys fodiens nanus Lydekker, Zoological Record, xlii (1905),
jMamm., p. 34, August, 1906 (Accideutal renaming of naias).

1910. Ncomys fodiens, N. fodiens naias and N. fodiens minor Trouessart,
Faune ]\Iamm. d'Europe, pp. 66, 57.

Ti^pe localitij. — 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 suftused 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 (18-20j. 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.

Siycmnens 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.).

Fbance : Guines, Pas-de-Calais, 2 ; Abbeville, So nme, 2 (B.M. and Paris ;
the latter type of remifer Geoffrey) ; near Paris, 1 (Paris ; type of lincatus
Geoffrey) ; Dinan, C6tes-du-Nord, 1 ; Cadillac-sur-Garonne, Gironde, 2
(U.S.N.M.) ; Montrejeau, Haute-Garonne, 3 (U.S.N.M.) ; Port6, Pyr6n6es-
Orientales, 5 ; I'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) ; Chesieres, Vaud, 4 (Mottaz) ; Les
Plans, Vaud, 3 (U.S.N.M.) ; Meiringen, Bern, 9 (U.S.N.M.) ; Grindelwald,

* " Neomys fodiens ■minor " Miller.

t ^^ Neomys fodiens naias ^^ Barrett-Hamilton.

72

INSECTIVORA

Bern, 1 (U.S.N.M.) ; Miirren, 2; Goschenen, Uri, 2 (U.S.N.M.) ; St. Got-
hard, Uri, 1 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 8 (B.M. and U.S.N.M.) ;
St. Fiden, St. Gallen, 2 (U.S.N.M.); Murgsee region, St. Gallen, 5
(U.S.N.M.) ; Au, St. Gallen, 1 (U.S.N.M.) ; Uzwil, St. Gallen, 1 (U.S.N.M.) ;
Sitterwald, St. Gallen, 1 (U.S.N.M.) ; Paido, Ticino, 1 ; Gordola, Locarno,
Ticino, 3 ; Porlezza, Ticino, 1 (Mottaz) ; Muzzano, Ticino, 1 (Mottaz) ; no
exact locality, 1.

Italy : Busalla, Liguria, 3 ; Vallombrosa, Florence, 1.

Remarks. — There is much individual variation in colour as
well as in external measurements and proportions,* though the
size of the skull and teeth, as may be seen from the Tables,
remains very constant. The degree of development of the fringes
on feet and keel on tail is also variable, the depth of the keel in
some individuals being about equal to diameter of tail, while in
others it is too slight to be measured. This is partly, if not
entirely, due to season, as the deepest keels are found in winter
specimens. With regard to the variations in external measure-
ments shown by the averages and extremes in a preceding
paragraph, it is probable that they are much exaggerated by
different methods in taking the measurements. This is well
illustrated by the two series of Pyrenean specimens measured
by A. Robert at an interval of six years. While much
material has been examined it is not wholly satisfactory, con-
sisting chieHy of small lots taken by 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

1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 1.

way. Waldo (c & p).

6. Kvikne, Hedemarken, G. Barrett-Hamilton 11. 1. 2. 88.

2200 ft. [N. F. Ticc- (p).

hurst.)

9. Brekkebygden, Trond- G. Barrett-Hamilton 11. 1. 2. 89.

bjem [N. F. Ticehurst.) (p).

1. Botbnia, Sweden. Purchased. (Wahl- 38. 9. 24. 14.

berg.)

9. Medstugan, Jemtland, Lord Lilford (p). 11. 1. 1. 147.

550 m. (G. KoWioff.)

2 9. Guines, Pas-de-Caiais, 0. Thomas (c & p). 94: 6. 6. 1-2.

France.

6, 2 9. Porte, Pyr^n^es-Orieu- 0. Thomas (p). 8. 9. 1. 39-41.

tales, 1600-1700 m.

France. [A. Robert.)

* This tendency to variation, coupled with the animal's peculiarly
attractive and interesting appearance, has led to the creation of the most
formidable synonymy (33 distinct names) yet applied to a European
mammal.

73

9.
6,2.3 9.

2 9.

6 c?, 4 9.

4 9.

9.
3 6.

].

6,2 9.

1 al.

L'Hospitalet, Ariege,
1450 m. (.4. Robert.)

L'Hospitalet, Ariege.

Luclaon,Haute-Garonue,
600-900 m. (A.Eobert.)

Bareges, Hautes-Pyre-
nees, 1300-1500 m.

Barcelounette, Basses-
Alpes. (C. Mottaz.)

Cranves-Sales, Haute-
Savoie.-

Abbeville, Somme.

Frankfort, Wiesbaden,
Germany.

Niesky, Silesia, 181 m.
(W.Baer.)

Strassburg, Alsace.
(C. Mottaz.)

H a i d a, Bohemia,
Austria.

Hatszeg, Hunyad, Tran-
sylvania, 1500 ft.
Hungary.

Hatszeg, Transylvania,
1500-2000 ft.

Miirren, Bern, Switzer-
land.

Ziiberwangen, St.Gallen.
(E. Zollikofer.)

Faido, Ticino.

Locarno, Ticino.

Switzerland.

Busalla, Liguria, Italy.

Vallombrosa, Florence.

0. Thomas (p).

G. S. Miller (c).
O. Thomas (p).

G. S. 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).

G. G. Danford (c).

W. Gurtner (p).

0. Thomas (p).

0. Thomas (c & p).
0. Thomas (c & p).
Purchased(Parreys) .
O. Thomas (c & p).
Dr. G. Cecconi (p).

8. 9. 1. 42.

8. 8. 4. 139.
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 bicolor Shaw.

1791. Sorex hicolor Shaw, Naturalists' Miscellany, ii, pi. 55, named in

index (Oxford, England).
1805. Sorcx ciliatus Sowerby, British Miscellany, xlix, p. 103 (Norfolk,

England).
1837. Amphisorex pennanti Gray, Proc. Zool. Soc, London, p. 125 (England).
1840. Crossopus sowcrhyi i Bonaparte, Iconogr. Faun. Ital., i, fasc. 29, in

text under G. fodiens.
1857. Crossopus fodiens Blasius, Siiugethiere Deutschlands, p. 120 (part).
1905. Ncomys fodiens ciliatus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., xv, p. 508, May, 1905.
1910. NeoniTjs fodiens ciliatus Trouessart, Faune ]\Iamm. d'Europe, p. 56.

Type locality. — Oxford, England.

Geographical 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 (Gi-antown-on-Spey and Cortachy, Forfar) : head

74

INSECTIVORA

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 foot, 17. For cranial measurements
see Table, p. 77.

Specimens examined. — Forty-six, from the following localities : —

Scotland : Gordonstown, Elgin, 1 ; Dunphail, Elgin, 1 ; Granto\vu-on-
Spey, Elgin, 1 (Wilson) ; Cortachy, Elgin, 1 (Wilson) ; Cromlix, Stirling-
shire, 1 ; Knibruck, Stirlingshire, 1 ; Aberdeen, 1 ; Penkill, Ayrshire, 1.

Wales: Fishguard, Pembrokeshire, 1; no exact locality, 1.

England : Mill-on-Tyne, Northumberland, 1 ; Leeds, Yorkshire, 1 ;
Wellington, Somerset, 1 ; Great Grimsby, Lincolnshire, 2 ; Shaftesbury,
Dorset, 1 ; Hales^Yorth, Suffolk, 1 ; Cambridgeshire, 2 ; Thoruhaugh,
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 ; IBasingstoke, Hampshire, 2 ;
Hampshire, no exact locality, 2 ; Yalding, Kent, 1.

i. Dunphail, Elgin.

1 al. 9 Aberdeen, Scotland.

1. Penkill, Ayrshire.

9. Fishguard, Pembroke-

shire, Wales.

1. Wales.

9. Mill-on-Tyne, North-
umberland, England.

1 al. W^ellingborough, North-

amptonshire.
1. Halesworth, Suffolk.

?. Cambridgeshire.
1. Cambridgeshire.

(J. Baker.)
1. Epping, Essex.

2 al. Henley, Oxfordshire.

1 al. Bur y St. Edmunds,

Suffolk.

3 al. Bury St. Edmunds,

Suffolk.
6 al. Esher, Surrey.
9 al. Godalming, Surrey.

(5,3 9. New Forest, Hampshire.

2 9. Basingstoke, Hampshire.

2 al. Hampshire.

Miller Collection.
Hon. N. C. Roths-
child (p).
E. R. Alston (p).
Y. H. Mills (c & p).

S. Stokes (c & p).
Rev. H. H. Slater

(c & p).
R. R.Orlebar (c&p).

Rev. W. R. Tate

(c & p).
J. Baker (c&p).
E. R. Alston (p).

J. Baker (p).

W. Royal Dawson

(c & p).
J. H.Powell (c&p).

Duncan Parker (p).

E. Dalgleish (c&p).
W. T. Blanford

(c & P).
Miller Collection.

Miller Collection.

E. Bartlett (p).

7. 7. 7. 2848.

10. 11. 28. 1.

79. 9. 25. 8.

11. 1. 3. 376.

48. 9. 24. 4.
11. 1. 3. 375.

84. 4. 21. 1.

87. 6. 7. 1.

39. 9. 29. 26.
79. 9. 25. 7.

40. 4. 11. 9.
11. 1. 3. 377.

83. 8. 8. 1.

84. 5. 16. 1-3.

11. 1. 3.378.
94. 8. 5. 2.

7. 7. 7. 2845-

2849.
7. 7. 7. 4470-

4471.
74. 11. 24. 1-2.

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78

INSECTIVORA

NEOMYS MILLERI Mottaz.

1907. Neomys milleri ^lottaz, Mem. Soc. Zool. de France, xx, p. 22,

September 20, 1907.
1910. Ncomijs milleri Trouessart, Faune Mamm. d'Europe, p. 58.

Type localiti/. — 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'^ and vf ; 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. Mamma; : a 1-1,
i 4-4 = 10.

Colour. — The colour is similar to that of typical specimens of
Neomys fodiens fodiens. Among the ^ins 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 nr. Teeth smaller throughout, a
difference especially noticeable in the anterior upper incisor.
Upper unicuspids with crowns longer and narrower than in
Neomys fodiens, the long posterior portion especially noticeable
in lateral view by comparison with height of cusp ; cingula
not so well developed as in the related species, and seldom, if
ever, forming a postero-external cusp.

Measurements. — External measurements of type : head and
body, 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
sjDecimens from Untervatz, Grisons, Switzerland : head and body,
79-3 (75-82); tail, 46-8(45-51); hind foot, 14-16 (14-15-4).
Two adult males from Locarno, Ticino, Switzerland : head and
body, 86 ; tail, 50 and 53 ; hind foot, 16 and 17. Average and
extremes of three adult females from 1' Hospitalet, Ariege,

NEOMYS 79

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.M.) ;
Chesieres, Alpes Vaudoises, 10 (B.^NI. and Mottaz) ; Meiringen, Bern, 2
(U.S.N.M.) ; Untervatz, Grisons, 6 (U.S.N.M.) ; Ziiberwangen, 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).

Austria-Hungary : Zuberei', Northern Hungary, 1.

Prance : L'Hospitalet, Ariege, 3 ; Luchon, Haute-Garonne, 1 ; Bareges,
Hautes-Pyreuees, 1.

Bemarhs. — Though superficially resembling Neomi/s 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 trvie 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 highh^ 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. i. 1-38, 140.

France.
?. Chesieres, Alpes Vau- C. Mottaz (p). 6. 2. 6. 4.

doises, 1200 m. Swit-
zerland.
?. Ziiberwangen,St.Galleu, 0. Thomas (p). 4. 4. 5. 52.

Switzerland.

{E. H. Zollikofer.)
2 i. Locarno, Ticino, Swit- 0. 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 (I'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, I\Ir. r^Iottaz
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
ac[uatic as those of N. fodiens.

80

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PACHYURA

81

NEOMYS ANOMALUS Cabrera.

1907. Ncomys anomalus Cabrera, Ann. and Mag. Nat. Hist., 7th ser., xx,

p. 214, September 1, 1907.
1907. Neomys anomalus Cabrera, Bol. Real Soc. Espafi. Hist. Nat., Madrid,

VII, p. 224. Published November, 1907.
1910. Neomys fodiens anomalus Trouessart, Faune Mamm. d'Europe, p. 57.

Type locality. — San Martin de la Vega, Madrid, Spain.

Geographical distribntion. — -Northern and central Spain.

Diagnosis. — Similar to Neomys milleri, but with longer tail
(ilsually 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.

Remarlcs. — The Spanish water-shrew is well differentiated
from Neomys milleri by its longer tail and larger hind foot. As
3'et 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 6,19. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 22-30.

2 al. La Granja, Segovia. M. dela Escalera (c). 8. 7. 30. 6-7.

(5,9. Barracas, Castellon. O. Thomas (p). 8.2.9.41-42.
[N. Gonzalez.)

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. u. 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 j"^, c ^J, pm -^^, m ^^ = 30.

Bemarlcs. — 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 etriiscus Savi, Nuovo Giom. de' Letterati, Pisa, i, p. GO (Pisa,

Italy).
1835. [Sorex] pachyurus Kiister, Isis, p. 77 (Cagliari, Sardinia).
1841. Pachyura etrusca Bonaparte, Iconogr. Faun. Ital., i, Indice distrib.
1857. Crocidiira suaveolens Blasius, Saugethiere Deutschlands, 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 Crocidura russula (head and
body, 35-40 mm. ; hind foot, 7*6-8 mm.), but with tail (:^o-
30 mm.) relatively longer, its ratio to head and body about
70 ; skull excessively small, its condylobasal length only about
1 2 • 8 mm. , the dorsal profile nearly straight from front of nasal
to back of parietals, usually a little concave in interorbital
region, the brain-case narrow but even more flattened than in
Crocidura leucodon and G. mimula ; teeth not conspicuously
difierent from those of the smaller European species of Crocidura,
except for the presence of the fourth upper unicuspid, and their
much smaller size (maxillary tooth-row about 6 mm.).

External characters. — ^In general external form 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 C. russula, and the head and feet are equally small
in proportion, a character by which Pachyura etrusca mav be
easily distinguished from young individuals of the larger animal.
Mammte, 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.

Slctdl. — 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. — 0. T.

G 2

tWdj^

84 INSECTIVORA

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
^^^T^ which to condylobasal length is about 46,

slightly less than in Crocidura mimula. Ante-
orbital foramen relatively large, its position
normal. Lachrymal foramen over posterior
half of m^. Palate relatively short and wide.
Mesopterygoid space relativel}^ longer than in
the European forms of Crocidura, its anterior
border on line with posterior edge of third
Fig. 16. molar instead of distinctly behind it, its lateral

Nat. size. borders nearly parallel, though slightly con-

verging posteriorly ; hamular excessively deli-
cate, bowed outward and upward, its length relatively greater
than in C. russiila, and about equal to greatest breadth of fossa.
Mandible not pecuKar 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 Crocidurse 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 Crocidurn, 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

PACHYTJRA

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86

INSECTIVORA

Genoa, Italy : head and body, 41 and -Ir'i ; tail, 26 and 28 ; hind
foot, 7 • 6 and 8 * 0. Adult female from vSassari, 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) ; Parti, Fiiialborgo, 2 (Genoa) ; near
Genoa, 5 (U.S.N.M. and Genoa) ; Pisa, 5 (B.M. and Mottaz) ; Florence, 6
(U.S.N.M. and Mottaz) ; Tuscany, 2 ; Rome, 1 (Genoa) ; no exact locality, 2.

Sicily : No exact locality, 2.

Sardinia: Assuni, 1 (U.S.N.M.). No exact locality, 1.

Greece : Lamia, 1.

lal.

Pisa, Italy.

Marquis G. Doria (p).

69. 3. 4. 1.'

2al.

Tuscany.

(No history).

1 al.

Italy.

Zoological Society's
Collection.

55. 12. 2G. 295.

Siieleton.

Italy.

Purchased.

58. 10. 21. 13.

6, ? al.

Sicily.

Purchased (Parzu-
daki).

52. 2. 26. 22-23,

1 al.

Sardinia.

Hon. N. C. Roths-
chUd (p).

11. 10. 1. 1.

lal.*

Lamia, Greece.

E. MacDonell (p).

8. 7. 22. 1.

Genus CROCIDURA Wagler.

1832. Crocidiira Wagler, Isis, p. 275 (Sorex leucodon Hermann).

1857. Crocidiira Blasius, Saugethiere Deutschlands, p. 137.

1869. Leucodon Patio, Faune Vert. Suisse, i, p. 132 (substitute for

Crocidura).
1897. Paurodus Schulze, Mamm. Europ., in Helios, Abhandl. u. Vortrage

Gesammtb. Naturwiss., xiv, p. 90 {Sorex leucoclon Hermann and

S. arancus Schreber = S. russulus Hermann).

Tijpe species. — Sorex leucodon Hermann.

GeograpMcal 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 :
^ lE'i' ^ 111' J"'^ i5i' "^ |E| = 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 Sore.v 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 lather heavy ; ear rising noticeabh^

* (5, 2 ?. Marismas, Lower Guadal- A. Chapman (c & p). 11.12.19.1-3.
quivir, Spain.
Received December, 1911. — 0. T.

CROCIDURA

87

above fur, the meatus closed by two valves as in Nt'omijs ; tail
somewhat thickened, its surface covered with short hairs, among
which are sprinkled numerous longer ones ; habits terrestrial.

Bemarks. — Though not highly modified in external pecu-
liarities the genus Groddura is the least primitive group of
shrews occurring in Europe. This is indicated by the reduced
number of up^ier 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 Soriddae the species of Groddura 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 OF CROCIDURA.

Tail decidedly more than half as long as head and
body, the ratio normally varying from 70 to 80.

Hind foot about 14 mm. (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 unicuspid 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. mimiua, p. 94.

Colour tending to be pale and greyish (Basque

Provinces, Spain) C. m. caatabra, p. 99.

Colour tending to be dark and brownish.

Brain-case not decidedly flattened (Charente,

France) C. m. icidisma, p. 98.

Brain-case decidedly flattened (entire range

of species, western portion excepted)... C. m. mimula, 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) C. leitcodon, p. 88.

Animal obscurely bicolor ; condylobasal length

of skull usually 18 to 19 mm. (Sicily) C. sicula, p. 108.

Brain-case more than half as high as wide C. riissula, p. 99.

Condylobasal length of skull usually 19 to
20-4 mm. (Central Europe, south into

Italy) C. r. russvla, p. 101.

Condylobasal length of skull usually 18 to 19 mm.

Colour a dark, coppery-brown (Portugal) C. r. cintrx, p. 108.

Colour a pale drab-brown (Spain and south-
western France) C. r. pulchra, p. 103.

88 INSECTIVORA

CROCIDURA LEUCODON Hermann.

1780. Sorcx leucodon Hermanu in Zimmermann, Geogr. Gesch., ii,

p. 382 (vicinity of Strassburg, Germany).

1781. Sorex leucodon Hermann in Schreber, Siiugthiere, pi. clix.

1782. Sor[t'.r] 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, Siiugethiere Deutschlands, p. 140.
1869. Leucodon microuriis Fatio, Faune Vert. Suisse, i, p. 173 (Substitute

for leucodon).
1897. [Crocidura] leucodus Scbulze, ]\Iammalia Europsea, p. 18 (Substitute

for leucodon).
1910. Crocidura russula leucodon Trouessart, Faune Mamm. d'Europe, p. 14.

Tijpe locality. — Vicinity of Strassburg, Germany.

Geograjjhical 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, 11 to 13 mm. ; condylobasal length of skull, 18 ■ 4 to 20 mm. ;
upper tooth-row, 8*8 to 9*0 mm. ; tail short, its actual length
28 to 38 mm., its ratio to head and body usually ranging from
38 to 43 ; skull with brain-case noticeably depressed, its height
less than half its width ; large upper premolar with antero-
external cusp rather large, its height usually greater than that
of first unicuspid ; colour of underparts whitish, strongly con-
trasted with rather slaty brown of back, the line of demarcation
along sides well defined.

External characters. — Fvir 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 (h) loosely spreading hairs about 5 mm. long, rather
thickly sprinkled among the others ; annulation ill-defined,
about 35 to the centimeter at middle. Mammae, i 3 — 3 = 6.

CROCIDURA

89

Colour. — UpjDer 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 eftect 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 suff'used 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.

SJculI. — 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 consjiicuously
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 earl}^ in life. Brain-case
slighth^ longer than broad, its posterior out- /^ " — -~~^
line rounded but broken by the slightly U^s^i^-i.^'^^^^r'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 tig. 20, page 100). Dorsal profile
with a slight concavity at front of brain-case
and slight convexity over middle of rostrum.
Nares squarely truncate posteriorly, the crotiav^aieucodon.
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'^. 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

INSECTIVOKA

Fig. 18.

Crocidura leucodon. Anterior
teeth in pioflle. 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,
occupjnng about one-third area of
crown. Second and third unicus-
pids aj)proximately 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 difieren-
tiated than in first.
Second smaller than
third, its crown aboiit
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

i'lG. 19.
Crocidura leucodon. Teeth X 10.

CROCIDURA 91

Neomys, 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 jjoint 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 nir^
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, tSwitzerland : head and body, 80 (77-81); tail,
32 • 2 (31-36) ; hind foot, 12 • 5 (1 1 • 8-13). Average and extremes
of three specimens from Florence, Italy (in alcohol, bodies
contracted): head and body, 63-3 (63-64); tail, 34-3 (32-36) ;
hind foot, 12 '3 (11 •6-13). For cranial measurements see Table,
p. 92.

Specimens examined. — Sixty-six, from the following localities : —

Belgium: Esneux, Liege, 2; Waremme, Li6ge, 1 (U.S.N.M.).

France : 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.) ; Ziiber-
wangeu, St. Gallen, 1.

Italy : Porlezza, Como, 1 (^Mottaz) ; Boccadassa, Genoa, 1 (Genoa) ;
Pisa, 1 (Mottaz) ; Florence, .3 (U.S.N.M. and Mottaz) ; Borne, 1.

Bemarks. — This species is recognizable among the European
members of the genus by its sharply bicolor pattern of coloura-
tion and by the low, flattened brain-case. From Crocidura
riissnla, the only species with which it is likely to be confused,
it is further distinguished by the peculiarities of the large upper
premolar, a tooth whose higher cutting edge, better developed
paracone and more prominent cingulum indicate a degree of
etticiency superior to that of the corresponding tooth in the related
animal.

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94 INSECTIVORA

2 9, 1. Esneux, Liege, Belgium Lord Lilford (p). U5. 1. 1. 1-2.

(if. Gronvold.)
6. Dinan, Brittany, France. G. Barrett-Hamilton 11. 1. 2. 158.

6. Manonville,Meurthe-et- Lord Lilford (p). 8.9.8.1.

Moselle.
5 (^, 4 ?. Etupes, Doubs, 350 m. O. Thomas (p). 8. 8. 10. .33-41.

(C. Mottaz.)
2 9. IMarxheim, Bavaria, Lord Lilford (p). 8. 9. 8. 2-3.

Germany.
1. Frankfort, Hessen- Dr. Diefienbach (p). 47.1.8.50.

Nassau.
6. Nussberg, Brunswick. G.Barrett-Hamilton 11. 1. 2. 90.

1 al. Germany. Tomes Collection. 7. 1. 1. 45.

9. Hatszeg, Transylvania, C. G. Danford (c). 3. 2. 2. 16.

1500 ft. Hungary.
3(5,4 9. Untervatz, Grisons, O. Thomas (p). 4.4.5.21-22.

Switzerland. 10. 8. 16. 9-18.

{E. H. Zollikofcr.)
9 sk. Grisons, 550 m. O. Thomas (p). 4. 4. 5. 20.

{E. H. Zollikofcr.)
9. Lugano, Ticino. 0. Thomas (p). 10. S. 16. 15.

{E. H. Zollikofcr.)
9. Ziiberwangen,St.G'allen. 0. Thomas (p). 10. 8. IC. 14.

[E. H. Zollikofer.)
6. Rome. (C. Coll.) G.Barrett-Hamilton 11.12.103.

CROCIDURA MIMULA Miller.
(Synonymy under subspecies.)

Geographical distribution. — Central Europe from north-
western Spain through France, Switzerland, central Germany
and northern Hungary to Roumania and Bulgaria, south into
Italy and Greece.

Diagnosis. — Size less than in Crocidura russtda and C. leucodon
(hind foot, 10 to 12 mm. ; condylobasal length of skull, 16 to
17 "6 mm.) ; skull and teeth resembling those of C. leucodon, the
brain-case nearly as much depressed, its height seldom more than
half width, the third unicuspid similarly low as compared with
small anterior cusp of large premolar ; colour not very different
from that of C. aranea, the upper and lower surfaces of body not
strongly contrasted, and no line of demarcation along sides.

Colour. — ^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 butty whitish, usually with
some dark clouding. Tail greyish or brow^nish, very obscurely
bicolor.

Skull and teeth. — Except for their noticeably smaller size the
skull and teeth closely resemble those of Crocidura leucodon,
though the brain-case is less constantly flattened, sometimes
assuming a form essentially like that in C. ritssida. Plate forming

CROCIDURA 95

outer wall of anteorbital canal narrower than in C. lencodon, its
width scarcely twice that of lachr^-mal foramen. Large upper
premolar with high cutting blade and well developed paracone,
the form and size of which relatively to main cusp and to third
unicuspid are exactly as in C. leucodon.

Measurements. — According to measurements made by various
collectors, the head and body varies from 55 to 72 mm., tail
from 28 to 40 mm., and hind foot from 10 to 12 mm. The
condylobasal length of skull ranges between 16 '0 and 17*6 mm.,
and length of upper tooth-row between 7 '2 and 8"0 mm.

lie marks.— Grocidnra 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 G. russula, though
not sufficiently to cause any confusion. In addition to the
typical form two geograj^hical races have been described, one from
south-western France, the other from northern Spain, the
status of neither of which is clearly understood.

Crociduea mimula mimula Miller.

1839. ?? ICrocidura arancaA, var. minor de S^lys-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 antipx Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 228, November, 1901 (Siulnita and Barza,
Roumania).

1902. [Crocic/«)Tf] viinuta Lydekker, Zool. Record, xxxviii (1901), Mamni.,

p. 27 (Accidental renaming of mimula).
1910. Crocidura mimula and C. antipai Trouessart, Faune Mamm. d'Europe,
pp. 46, 48.

Tjipc locallti/. — Ziiberwangen, St. Gallen, Switzerland.

GeugrajjJiical distribution. — Range of the species from the
Rhone Valley eastward.

Diagnosis. — Size maximum for the species ; skull with brain-
case tending to be strictly of the flattened type ; colour usually
dark.

Measurements. — External measurements of type : head and
body, 72; tail, 33; hind foot, 11. A male and female from
Untervatz, Grisons, Switzerland : head and body, 71 and 65 ;
tail, 35 and 35 ; hind foot, 10 and 11. A male and female from
Marxheim, Bavaria : head and body, 71 and 58 ; tail, 36 and 30 ;
hind foot, 10 and 10. Average and extremes of four specimens
from Haida, Arva, Bohemia : head and body, 59 • 7 (55-65) ;
tail, 30 (28-32) ; hind foot, 11-5 (1 1-1 2). A male from Gageni,
Roumania, and female from Bustenari, Roumania : head and
body, 64 and 66 ; tail, 31 and 34; hind foot, 11 and 11. Two
adult males from Agay, Var, France : head and body, 64 and 66 ;
tail, 35 and 34; hind foot, 11 and 11. Adult female from
Italy : head and body, 70 ; tail, 38 ; hind foot, 11 "6.

96

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98 INSECTIVOKA

Adult male and female from Corfu, Greece : head and bod}-, 74
and 75 ; tail, 44 and 44 ; hind foot, 11 "8 and 11 • 4. For cranial
measurements see Table, p. 96.

Specimens examined. — Thirty-seven, from the following localities : —

Prance : Abbeville, Somme, 1 ; Agay, Var, 2.

Germany : Marxheim, Bavaria, 2.

Austria-Hungary : Haida, Arva, Bohemia, 4 ; Hatszeg, Hunyad,
Transylvania, 1; Tatra Mts., Hungary, 1.

RouMANiA : Gageni, Prahova, 1 ; Busteiiari, Prahova, 1.

Bulgaria: Sofia, 1 (Andersen); Varna, 1 (Andersen).

Switzerland: Ziiberwangen, St. Gallen, 3 (U.S.N.M. and INIottaz) ;
Untervatz, Grisons, 2 ; Faido, Ticino, 1 ; Santa Margherita, Ticino, 3
<Mottaz) ; Davesco, Ticino, 1 (U.S.N.M.) ; Lugano, Ticino, 2 (B.M. and
Mottaz) ; Locarno, Ticino, 1.

Italy : Porlezza, Como, 4 (Mottaz) ; Viareggio, Lucca, 2 : Rome, 1.

Greece : Corfu, 2.

1. Abbeville, Somme, Baillon Collection. 56. b.

France.
2 6. Agay, Var. G. S. MiUer (c). 8. 8. 4. 162-163.

6. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.4.

Germany. (Wolter-

storff.)
6,9. Haida, Bohemia, Lord Lilford (p). 8. 9. 8. 5-6.

Austria.
9. Hatszeg, Hunyad, Tran- C.G.Danford (c&p). 3.2.2.12.

sylvania, Hungary.
1 al. Tatra Mc-mtains. Dr. R. Collett (p). 91. 1. 21. 2.

?. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 11.

mania. {W. Dodson.)
6. Bustenari, Prahova, Lord Lilford (p). 4. 4. 6. 12.

840 m. {W. Dodson.)
?. Faido, Ticino, Switzer- 0. Thomas (c & p). 5. 8. 2. 18.

land.

5. Locarno, Ticino. 0. Thomas (c & p). 5. 8. 2. 1.
(5,9. Untervatz, Grisons. O. Thomas (p). 4.4.5.21-22.

(E. H. Zollikofer.)
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. Rome. (Coli.) G.Barrett-Hamilton 11.1.2.97.

(P)-
6. Corfu, 50 m. Greece. J. I. S.Whitaker (p). 8.10.1.8.
(C. Mottaz.)

Crocidura mimula iculisma Mottaz.

1908. Crocidura tnimula 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'Europc, p. 47.

Type locality. — Lignieres-Sonneville, Charente, France.

Geograiiliical 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. russida.

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).

BemarJcs. — 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.

Crocidura mimula cantabra Cabrera.

1908. Crocidura cantabra Cabrera, Bol. Real Soc. Espaii. Hist. Nat., viii,

p. 239, May, 1908. Type in Madrid Museum.
1910. Crocidura cantabra Trouessart, Faune IMamm. d'Europe, p. 46.

Tjipe locality. — Basque Provinces, Spain, exact locality not
known.

Geographical (Jistrihution. — 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.)

Remarhs. — I have not seen this animal, but from the original
description, as well as from information received from Mr. Cabrera,
it appeal's to be paler than the typical form, to which it bears
much the same relation as C. russula pulclira to true riissula.

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 unicusjaid ;
colour of underparts not strongl}^ 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 bufly 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 bufiy grey or
light drab. Tail obscurely bicolor, like back above, like belly
below.

SJctdl 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

Pig. 20. Fig. 21.

Posterior view of skull of Crocidura russula. Anterior

Crocidura leucodon (upper teeth in profile. X 5.

figure), and C. russula (lower
figure). X 1*.

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.

Meastirements. — 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 ihoracicus
and Dehne's S. chrysotliorax.

CROCIDURA 101

Crocidura eussula russula Hermann.

1777. Sorex araneus Schreber, Saugthiere, iii, p. 573 (not of LiniiEeus, 1758).
1780. Sorex rassulus Hermann in Zinimermanu, Geogr. Gesch., ii, p. 382

(vicinity of Strassburg, Germany).
1780. ?? Sorex constrictus Hermann in Zimmermann, Geogr. Gescb., ii,

p. 383 (vicinity of Strassburg, Germany). Based on yomig in nest.

1792. ?? Sorex unicolor Kerr, Auim. Kingd., p. 208 (Strassburg, Germany.

Based on Pennant's account of Sorex constrictus).

1793. Sorex musaraneus Cuvier, Tabl. 15lem. de I'Hist. Nat. des Anim.,

p. 109 (France).

1800. ?? Sorex leucurus Sbaw, Gen. Zool., i, pt. 2, p. 538 (Strassburg,

Germany. Based on Scbreber, pi. clixc, S. constrictus Hermann).

1801. S[prcx'\ a\_raneHs'] cincrcus Bechstein, Gemeinn. Naturgescb. Deutscb-

lands, I, 2iid ed., p. 867, misprinted 863 (Tbiiringen, Germany).
ISOl. S[orcx\ a[raiieu,s'] candiclus'Bechstem,Geraeinn. Naturgescb. Deutscb-

lands, I, 2nd ed., p. 867, misprinted 863 (Tbiiringen, Germany).
1832. Sorex fimbriatus Wagler, Isis, p. 54 (Bavaria, Germany).
1832. Croc[^idiira\ moschata Wagler, Isis, p. 275 (Substitute for Sorex

fimbriatus).
1832. Cl^rocidura] major Wagler, Isis, p. 1218 (Bavaria, Germany).
1832. Crocidura rufa Wagler, Isis, p. 1218 (banks of tbe Rbine, Germany).
1832. Crocidura poliogaster Wagler, Isis, p. 1218 (banks of tbe Rbine,

Germany).
1832. Sorex tJwracicus Savi, Nuovo Giorn. de' Letterati, Pisa, xxiv, p. 52

(near Pisa, Italy).
1839. ? Sorex inodorus de Selys-Longcbamps, Etudes de Micromamm.,

p. 34 (Savi cited as autbority, but name apparently publisbed berc

for tbe first time as synonym of aranea. (= russula)).
1839. ? [Crocidura aranea?\ var. minor de S61ys-Longcbamps, Etudes de

Micromamm., p. 35 (Silesia).
1839. [Crocidura arujiecC] var. albivcntris de S61ys-Longcbamps, Etudes de

Micromamm., p. 36. (No locality given.)
1839. ? Crocidura- hydrunfina Costa, Fauna del Eagno di Napoli, Mamm.,

p. 6 (Otranto, Calabria, Italy).
1855. Sorex chrysothorax Debne, Allg. deutscbe Naturbist. Zeitung, Neue

Folge, I, p. 241 (Wilsdurf, near Dresden, Germany).
1857. Crocidura araneus Blasius, Siiugetbiere, Dentscblands, p. 144.
1895. Crocidura russula Tbomas, Tbe 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, 11 '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 bave seen five Sardinian specimens in tbe Genoa Museum
resembling tbe typical form of Crocidura russula and differing widely
from tbe Corsican C. cyrnensis (see pp. 111-112).

101

INSECTIVORA

extremes of nine specimens from Esneux, Liege, Belgium : head
and body, 77 (72-85); tail, 35-6 (33-38); hind foot, 12*4
(11* 7-13). Average of ten specimens from Pas-de-Calais, France :
head and body, 74 • 5 (71-80) ; tail, 40 • 9 (38-46) ; hind foot, 1 3 • 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 huudred 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 ; Liguieres,
Charente, 1 (Mottaz) ; Nancy, Meurthe-et-Moselle, 1 (Merriam) ; Etupes,
Doubs, 9 (Mottaz); MontaulDan, Haute-Savoie, 4; Valescure, Var, 2; Ax-
les-Thermes, Ariege, 1 ; Luchon, Haute-Garonne, 2.

Germany : Ummerstadt, Thiiringen, 2 ; Ingelheim, Rheinhessen, 2 ;
Strassburg, 1.

Switzerland : Geneva, 10 (U.S.N.M. and Mottaz) ; St. Cergues, Vaud, 9
(U.S.N.M. and Mottaz) ; Grosjean, Vaud, 1 (Mottaz) ; Chesieres, Vaud, 1
(IMottaz) ; Lucerne, 1 ; A^itznau, Lake of Lucerne, 3 ; Thurgau, Roggwil, 1 ;
St. Gallen, 3 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 (B.M. and U.S.N.M.) ;
Degersheim, St. Gallen, 5 (B.M. and U.S.N.M.) ; Engelberg, Unterwalden, 1.

Italy: Ceresole d'Alba, Turin, 2 (Turin).

3 (J, 1 9. Oosterbeek, Guelderland, 0. Thomas (c & p). 98. 2. 1. 9-12.

6,9.

4c5,4?,l,

1.
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1.
2.

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9.

2 9.

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1 al.

50 m. Holland

Oosterbeek, Guelderland.
(0. Thomas.)

Esneux, Liege, Belgium.

Belgium. (H. Grdnvold.)

Guernsey, Channel Is-
lands. {R.H. Bunting.)

Alderney.

Boulogne, Pas-de-Calais,
10 m. France.

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.Motta

Vitznau, Lake
Lucerne, 500
Switzerland.

Engelberg, 3300
Switzerland.

of
m.

Miller Collection.

Lord Lilford (p).
Tomes Collection.
0. Thomas (p).

W. Eagle Clarke (p).
0. Thomas (c & p).

0. Thomas (c & p).
Baillon Collection.
W. M. Daly (c & p).
A. Robert (c &p).

0. Thomas (p).

G. S. Miller (c).
V. Builles (c & p).
O. 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.

ft. Dr. J. Anderson (p). 99.7.17.3.

CROCIDURA 103

Crocidura russula pulchra Cabrera.

1907. Crocidura russula pulclira Cabrera, Ann. and Mag. Nat. Hist.,
7th ser., xx, p. -2,1'd, September 1, 1907. Type in Cabrera col-
lection.

1907. Crocidura russula pulchra Cabrera, Bol. Real Soc. Espafi. Hist.
Nat., Madrid, vii, p. 223, October, 1907. (For date see Cabrera,
Ann. and Mag. Nat. Hist., Sth ser., i, p. 189, February, 1908.)

1910. Crocidura russula pulchra Trouessart, Faune Mamm. d'Europe, p. 45,

Type locaUtjj. — 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 1.3, 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, 11*9 (10* 8- 12* 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 : —

France: 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 cintrai).

?. St. Genies, Gard, 102 m. 0. Thomas (p). 8. 8. 10. 42.
France. (C. Mottaz.)

4 i. Silos, Burgos, 980 m. G. S. Miller (c). 8. 8. 4. 38-41.
Spain.

?. Dehesa de Valencia. 0. Thomas (p). 8. 2. 9. 46.
{N. Gonzalez.)

2 <5, 1 ?. Alcoy, Alicante. 0. Thomas (p). 8. 2. 9. 4:3-45.
{N. Gonzalez.)

6,9. Venta del Baul, Granada. G. S. Miller (c). 8.8.4.31-32.

4 (J, 1 9. Granada. G. S. Miller (c). 8. 8. 4. :33-37.

(Jal. Sierra deGerez.Portugal. Dr. H. Gadow 87.3.28.1.

(c & p).

* Intermediate hetvieen jjulchra and true russula.

104

INSECTIVORA

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,, slightly worn.

,, not worn.

,, slightly worn.

,, not worn.
,, slightly worn.

,, moderately worn.
,, slightly worn.

,, much worn.

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108 INSKOTIVOKA

Crocidura russula ciNTRiE Miller.

1907. Crocidura russula cintrx Miller, Anu. and Mag. Nat. Hist., 7th ser.,

xs, p. 390, November, 1907. Type in British Museum.
1910. Crocidura russula cintrie Trouessart, Faune IMamm. d'Europe, p. 45.

Type locality. — Cintra, near Lisbon, Portugal.

Geograpltical distribution. — At present known only from the
type locality.

Diagnosis. — Size as in C. russula pulehra (hind foot, 11*4 to
12-7, condylobasal length of skull, 18 to 19 "2), but colour fully
as dark as in true russula, the back between the mars-brown and
russet of 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, 11-4. Average and extremes of
ten specimens from the type locality : head and body, 67 • 6
(64-72); tail, 37-7 (33-42); hind foot, 11-9 (11 -4-12 -7). For
cranial measurements see Table, j). 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, 5 ?. Cintra, Estremadura, 0. Thomas (c & p). 98. 2. 2. 10-20.
300-350 m. Portugal. (Tijpe of subspecies 98. 2. 2. 11.)

CROCIDURA SICULA Miller.

1879. ? Crocidura sicula Giglioli, Wiegmann's Archiv fiir Naturgesch.,
1879, I, p. 96. Nomen nudum : " Crocidura sicula (Giglioli MSS.
sp. nov. ?) 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
pulehra (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.

Shdl and teeth. — Except for its distinctly flattened brain-case

CKOCIDURA 109

the skvill i-esembles that of the small races of Crocidura russnla.
Teeth as in C. russiila, but upper molars with crown area
somewhat reduced, a character readily appreciable on comparison,
and lartje 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 Sicily. Exact localities :
Palermo, 5 (B.M. and U.S.N.M.) ; Marsala, 2 ; San Giuglielmo, Castel-
buono, 6 ; Ficuzza, 1.

2 ?. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 2-2*.

1. Marsala. {A. Eobert.) O. Thomas (p). 6. 8. 4. 26.

3 <5, 1 9. San Giuglielmo, Castel- 0. Thomas (p). 8. 9. 1. G-9.
buono. {A. Eohert.y

CROCIDURA CANEiE Miller.

1909. Crocichtra caneai Miller, Ann. and ]\Iag. Nat. Hist., 8th ser., iii,

p. 418, May, 1909. Type in British ]\Iuseum.

1910. Crocidura cancie Trouessart, Faune Mamm. d'Europe, p. 48.

Type locality. — Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Size and general appeai'ance as in the smaller
forms of Crocidura russula, and skull with similarly deep cranium ;
but second upper premolar as large as third, and entire anterior
portion of upper tooth-row unusvially 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 unicusj^ids
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 unicvispid 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 sufiiciently
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, Avdiile 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 tyjae (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. Eor 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.)
6. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 1.

(C. H. B. Ch-ant.)

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, Fauna Mamm. d'Europe, p. 49.

Type locality. — Palermo, Sicily.

GeograpMcal distribution. — Sicily.

Diagnosis. — Size about as in large specimens of C. russnla
(hind foot, 14 mm.). Tail very long, its ratio to head 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 difl:er in external characters from
C. russula. The tail is so long that when laid forward over back
it extends to between ears. It is distinctl}^ 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. russida. 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.

CROCIDURA 111

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.

Bemarlcs. — 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 IMiller, Anu. and ilag. Nat. Hist., 7th ser., xx,

p. 390, November, 1907. Type in British Museum.
1910. Crocidura cyrnensis Trouessart, Faune Mamm. d'Europe, p. 49.

TyjK' locality. — Bastia, Corsica.

Geographical distribution. — Corsica.

Diagnosis. — Smaller than Crocidura candata (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 ; hind foot,
12. For cranial measurements see Table, p. 113.

Specimens examined. — Three, all from Corsica.

Memarhs. — Crocidura cyrnensis is nearly related to C. caudata,
though readily distinguishable by its smaller size and less
thickened tail. So far as known it is the only shrew inhabiting
Corsica. Whether an animal of this type occurs in Sardinia
is a matter of doubt. Five Sardinian specimens (three from

112 INSECTIVORA

Ovile Seardu and two from Zinnigas) in the Genoa museum
appear to be strictly of the russida type, though without com-
parison of the skulls it is impossible to say whether they are most
nearly related to true russida or to skida. Their average and
extreme measurements (from sj^irit 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 & r). 93. 9. 15. 3,
Corsica.
S al. Bastia, Corsica. Mrs. Southwell (c & p). 6. 3. 14. 1.

{Type of species.)
? al. Bastia, Corsica. Mrs. Southwell (c & p). 9. 6. 14. 1.

CROCIDURA BALEARICA Miller.

1901. Crocidura russida Thomas, Proc. Zool. Soc. London, p. 39.

1907. Crocidura 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'Enrope, p. 49.

Type locality. — San Cristobal, Minorca, Balearic Islands.

Geographical distribution. — Balearic Islands, Spain.

Diagnosis. — In general similar to Crocichira 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.

Sliull 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. Extei'nal 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.

6,29 San Cristobal, Minorca; 0. Thomas and R.I. Pocock 0. 7. 1. 40-42.
Balearic Islands. (c & p).

(0. 7. 1. 42. Type of species.)

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11.4 INSECTIVORA

Family ERINACEIDiE.

1821. Ermaccidx Gray, London Med. Eepos., 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 ; aviditory process of basi-
sphenoid large, sometimes forming an evident half-bulla ; glenoid
surface directed downward (normal) ; a large external jjterygoid
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, stifi" spines.

EemarJi-s. — At present this family is usually regarded as
containing the single genus Erinaceiis. It is very probable,
however, that several genera are represented among the members
of the group.

Genus ERINACEUS Linnajus.

1758. Erinaceus Linnaeus, Syst. Nat., i, 10th ed., p. 52 {E. europseus).

1857. Erinaceus Blasius, Saugethiere Deutschlands, p. 152.

1868. Herinaceus Mina-Palumbo, Ann. Agric. Sicil., 2nd ser., sii, p. 37.

Type species. — EriJiaceus europseiis Linnaeus.

Geographical distribution. — Essentially coincident with that
of the family.

Characters. — Skull rather short and broad, the zygomatic
breadth distinctly more than half greatest length ; posterior
palatal region conspicuously fenestrate ; auditory process of basi-
sphenoid well developed, concave, sometimes forming a half-bulla ;
external pterygoid plate rather larger than internal pterygoid
plate, formed about equally of ectopterygoid and a broad horizontal
outgrowth from palatine ; dental formula : i ^"^, c 1^, pm |5|^
m 3I3 = 36 ; canines not difierentiated by form from the contiguous
teeth ; third upper molar consisting of a large i^rotocone 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.

ERINACEUS 115

Remarks. — About' twenty-five species are currently referred
to this genus, four of them occurring in Europe.

KEY TO THE EUROPEAN FORMS OF ERINACEUS.

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 southerji France)... E. alginis, p. 130.

Underparts clouded with brown, at least in inter-

ramial and intercrural regions (Spain and

southern France) 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. europxus, 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. Ms^xmicus, 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. italkus, p. 123.

Colour of head and shoulders darker than

that of contiguous spiny area (Sicily) E. e. consolei, p. 126.

ERINACEUS EUROPvEUS Linnaeus.

(Synonymy under subspecies.)

Geographical distribution. — Europe from the Mediterranean
coast to Scotland and sovithern 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 au 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 I'ostral
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, seconil 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 \vrist,
the outer the larger ; skin between pads w^-inkled ; hind foot
much like fore foot but longer ; second, third and fourth digits
sub-equal and longest, but their claws noticeably graduated from
second to fourth, fifth digit extending to base of fourth, first not
reaching base of second ; sole naked, the tubercles as on palm
but more crowded, the two posterior sub-equal. Tail short and
thick, its length much less than that of hiud foot. Ear simple,
rounded, shorter than bristles on crown, the meatus without
valves. Muzzle moderately produced, somewhat pointed, the
muzzle-pad well developed, naked, its surface marked w ith 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. Mammfe : p 1-1, a 2-2,
i 2-2 = 10.

Colour. — Furred portions of bodj- varying from dull brown to
dirty whitish, the under fur usually a dusky hair-brown, the
longer hairs lighter and more bufty. 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

* Rarely the bristles are replaced by coarse hair like that on under-
parts. See Natural Science, xiii, p. 156, pi. ii, September, 1898.

ERINACEUS

117

a slaty area of variable width, followed by a narrow but sharply
defined buffy annulation and an obscurely darker tip. The
general eftect 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.

Skidl. — General form of skull rather short, heavy and deep,
the zygomatic breadth about f upper length, the brain-case not

Fig. 22.
Erinaceus europxvs. 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 INSECTIVOEA

Ventral profile straight, the dorsal profile essentially parallel to
it from lambda nearly to front of interorbital region, then sloping
forward at a slight angle (about lo") ; 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, esjiecially in fourth unicuspid, to form a very
rudimentary secondary cusplet. First and second incisors invari-
ably single-rooted. Third incisorsingle-rooted, but root occasionally
showing trace of longitudinal furrow on outer side. Canine
usually single-rooted, the root with or without longitudinal
furrow ; but in a small series of specimens every stage may be
observed from this condition to a completely two-rooted tooth,
each root with a distinct alveolus. First premolar single-rooted,
the root 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

riG. 23.

Erinaceus eui-ojixus. Teetli.

120 INSECTIVORA

absent.* A small but evident metaconule. Second molar like
first but smaller, its crown area about equal to that of large
premolar, its greatest diameter anterior instead of posterior.
Cusps essentially as in first molar, except that paracone is larger
than metacone and hypocone, and metaconule is barely indicated.
Third molar reduced to a protocone nearly as large as in the
other teeth, and a rudimentary paracone, the two connected by
a cutting edge sloping obliquely outward, forward and uj^ward.
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 europ^us europ^us Linnaeus.

1758. [Eri7iaceus'} europseus Linnaeus, Systema Naturse, i, 10th ed., p. 52

(Sweden).
1779. \_Hystrix\ erinaceus Blumenbach, Handbuch d. Naturgesch., p. 72

(Germany).
1803. Erinaceus suillus Geoffroy, Catal. Mammif. du Mus. Nat. d'Hist.

Nat., p. 67 (France).
1803. Erinaceus caninus Geoffrey, Catal. Mammif. du Mus. Nat. d'Hist.

Nat., p. 68 (France).

1857. Erinaceus eurox^ams 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. u. Vortr. Gesammtb. Naturw. iv,

No. 10, p. 19 (Substitute for europaiiis).
1900. Erinaceus europa^us Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., v, p. 362, April, 1900.
1900. Erinaceus eiiropseus occidentalis Barrett-Hamilton, Ann. and Mag.

Nat. Hist., 7th ser., v, p. 362, April, 1900 (Haddingtonshire,

Scotland). Type in British Museum.
1910. Erinaceus europgeus and E. europmus occidentalis Trouessart, Faune

Mamm. d'Europe, p. 38.

Tijpe 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 (cond^'lo-
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.

EPaNACEUS 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; Innerwick,
Haddingtonshire, 1; Glendoc, Inverness, 1 (Wilson); Lanarkshire, 1.

England: Kelnsea, Spurn, Yorkshire,!; Leeds, Yorkshire,! (U.S.N.M.);
Somersetshire, 4 ; Shrewsbury, Shropshire, 3 ; Oundle, Northampton, 2 ;
Graftonbury, Herefordshire, ! ; Saffron Walden, Essex, 2 ; Banstead,
Surrey, !; Elstead, near Godalming, Surrey, !; Ockley, Surrey, !;
Wandsworth Common, Surrey, 1 ; Hampshire, 1 (U.S.N.M.).

Wales : Cardiff, 2.

Ireland : Castle Hamilton, ! ; Nenagh, Tipperary, 2 ; Ennis, Co.
Clare, 2; Glenmore, Co. Donegal, 1 (U.S.N.M.) ; Kilmanock, Wexford, 2.

Norway : Asker, near Christiania, !.

Sweden : Upsala, 5 (U.S.N.M.) ; Upland, !.

Denmark: Copenhagen, 5 (Andersen).

Holland : No exact locality, !.

Belgium : Maredsous, Namur, 2.

France : Foret de Bouconne, Gers, ! ; Cranves-Sales, Haute-Savoie, !.

Germany: Brunswick, 1 (U.S.N.M.) ; Heidelberg, ! (U.S.N.M.) ; Strass,
near Burgheim, Bavaria, 3; Ingelheim, Rheinhessen, 1.

Switzerland : Geneva, 2 (Mottaz) ; St. Gallen, 7 (B.M. and U.S.N.M.) ;
Uzwil, St. Gallen, ! (U.S.N.M.) ; Heresau, St. Gallen, ! (U.S.N.M.) ; Wolf-
halden, Appenzell, ! (U.S.N.M.); Thurgau, 2.

Bemarjcs. — The typical race of Erinaceus europseus is dis-
tinguishable from the forms occurring in the Mediterranean
region by its combination of large size with dark colour. It is
more readily confused with the dark E. roumanicus, whose range
adjoins it on the east, and from which it cannot be distinguished
with certainty except by comparison of the skull and teeth. The
cranial character supposed to distinguish British specimens from
the Continental form appears to be too inconstant to warrant the
recognition of an insular race.*

6, 9. Gordonstown, Elginshire, W. B. Ogilvie-Grant !1. !. 3. 379-

Scotland. (c & p). 380.

6. Innerwick, Haddington- W. Eagle Clarke (c & 0. 3. !3. !.

shire. p).

(Type of E. c. occidentalis B.-Ham.)

9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 76.

S, 9. Oundle, Northampton, Hon. N. C. Roths- !!. !. 3. 38!-

England. child (c & p). 382.

6. 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

IXSECTIVOEA

2 6. Saffron Walden, Essex.
{Wright.)
6. Banstead, Surrey.
i. Godalming, Surrey.
6. Ockloy, Surrey.

3juv.al. Shrewsbury, Shropshire.
6, 9. Somerset.
6, 9 al. Somerset.

9. Cardiff, Glamorganshire,

Wales.
6. Nenagh, Tipperar}', Ire-
land. {W. Smithivick.)
6 al. 6. Ennis, Clare.
2 6. Kilmanbck, Wexford.

1. Asker, Christiauia, Kor-
■way.
6 juv. Upland, Sweden. (G.
EoUhoff.)

1 al. Holland." {Seba Coll.)

2 6. Maredsous, Namur, Bel-

gium.
6. Foret de Bouconne, Gers,
250 m. France. (.-1.
Robert.)
9. Cranves - Sales, Haute-
Savoie. (A. Robert.)
6, 9. Burgheim, Bavaria, Ger-
many. (Woltcrstorff.)
6. Ingelheim, Rheinhessen.
2 6,9 juv. St. Gallen, 500 m.
Switzerland. {E. H.
Zollikofcr.)
6, 9. Thurgau, 400 m. {E. H.
Zollikofer.)

G. Barrett-Hamilton 11. 1. 2. 91-92.

C. H.B. Grant (c&p). 11. 1. 3. 386.

W.T.Blanford(c&p). 11. 1. 3. 384.

Hon. Ella Scarlett 11. 1. 3. 385.

(c & p).

H. E. Forrest (c & p). 0. 9. 23. 1-3.

Dr. J. Anderson (p). 93. 7. 31. 1-2.

Dr. J. Anderson (p). 93. 7. 31. 3-4.

R. Drane (c & p). 11. 1. 3. 387.

G. Barrett-Hamilton 11. 1. 2. 95.

J. W. Scott (c & p). 93. 10. 30. 1-2.

G. Barrett-Hamilton 11. 1. 2. 93-94.

(c & p).

Christiania Museum 93. 3. 1. 7.

(E).

Lord Lilford (p). 11. 1. 1. 151.

Lidth de Jeude Coll. 67. 4. 12. 555.

Rev. G. Fournier (c 1. 6. 2. 1-2.

& p).

0. Thomas (p). 0. 4. 1. 10.

0. Thomas (p).

Lord Lilford (p)

C. Hilgert (c).
0. Thomas (p).

0. Thomas (p).

6. 4. 2. 1.

11. 1. 1. 92, 150.

8. 11. 2. 5.
4. 4. 5. 27-29.

4. 4. 5. 30-31.

Erinaceus EUROPiEUS HisPANicus Barrett-Hamilton.

19C0. 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 axid 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 an}^

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 :
Arrechavaleta, Vitoria, 1; Pajares, Leon, 7; Burgos, 5; Palacios de la
Sierra, Burgos, 1; Bejar, Salamanca, 2; Seville, 4.

RemavTis. — The Spanish hedgehog is a moderately well
differentiated form. Extreme specimens are easily distinguish-
able from typical europseus ; but in general the difference between
the two races must be regarded as an average one.

6. Arrechavaleta, Vitoria, 0. Thomas (p). 8. 2. 9. 9.

Spain. (iV. Gonzalez.)

c5, 2 9, 9 juv. Pajares, Leon. [N. Gon- 0. Thomas (p). 8. 2. 9. 2-6.

zalez.)

6, 9, 9 juv. Burgos. G. S. Miller (c). 8. 8. 4. 17-19.

2 6. Burgos. {N. Gonzalez.) 0. Thomas (p). 8. 2. 9. 10-11.

2 6 juv. Bejar, Salamanca. {N. 0. Thomas (p). 8. 2. 9. 7-8.

Gonzalez.)

6, 2 9, 1. Seville. (Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 1-4.

{Type of subspecies 95. 3. 3. 2.)

Erinaceus eueop^us italicus Barrett-Hamilton.

1857. Erinaceus europseus Blasius, Saugethiere Deutschlands, p. 153 (part).
1900. Erinaceus europaus italicus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 364, April, 1900. Type in British Museum.
1910. Erinaceus europseus italicus Trouessart, Fauna Mamm. d'Europe,

p. 39.

Type locality. — Siena, Italy.

Geographical distribution. — Italian Switzerland, Italy and
Sardinia ; Corsica ?

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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126 INSECTIVORA

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.

Specime?is examined. — ^Seventeen, from the following localities : —
Switzerland : Bigorio, Ticino, 1 (U.S.N.M.) ; Curoggio, Ticino, 1
(U.S.N.M.); Gentilino, Ticino, 1 (U.S.N.M.).

Italy : Empoli, Florence, 1 ; Siena, 4 ; Ostia, Rome, 1 ; Rome, 4.
Sardinia: Su Cramu, 1; Bare, 1; Marusei, 1; Trecorgia, 1.

BemarJcs. — ^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.

9. Empoli, Florence, Italy. A. H. Savage Landor 97. 3. 7. 1.

(c & p).
2 (5, 2 9. Siena. (S. Brogi.) Dr. E. Hamilton (p). 98. 10. 2. 5-8.

{TyjJe of subspecies 98. 10. 2. 5.)
1. Ostia, Rome. Dr.L. Sambon(c&p). 1. 1, 2. 7.

3 6. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 3-4. 96.

6. Su Cramu, Sardinia. {W. 0. Thomas (p1. 0. 12. 3. 5.

Wolterstorff.)

S. Bare. {W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 3.

9. Marusei. (W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 6.

9 juv. Trecorgia. {'W. Wolterstorff.) 0. Thomas (p). 0. 12. 3. 4.

Erinaceus europ.eus 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 E. europseus italicns (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, si3i'inkled
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 jaroducing a slight
though evident darker median dorsal area. Feet so injured that
colour cannot be determined.

Shull 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 consolci 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 curopmus roumanicus Barrett-Hamilton, Ann. and Mag.

Nat. Hist., 7th ser., v, p. 365, April, 1900. (Gageni, Roimaauia.)
Type in British Museum.

1901. Erinaceus clanubicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 229, December, 1901. (Prundu, Roumania.)
1910. Eri7iaccus europxus roumanicus and E. europams dayiabicus Troues-
sart, Faune Mamm. d'Europe, pp. 40-41.

Type locality. — Gageni, Prahova, Rovimania.

Geoc/rapMcal 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.

shull 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

INSECTIVORA

transverse palatal ridge usually lower and less developed,
particularly at middle, where there is often a slight angle.
Ridge at margin of orbit in lachrymal region not so long as in
E. 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 Erinaceus
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.).

KRINACEUS

129

?.

Haida, Bohemia.

6, ?.

Gageni, Prahova, Roumania.

(ir. Dodson.)

6.

Potamos, Corfu, Greece.

(C. Mottaz.)

9.

Argostoli, Cephalonia.

(C. Mottaz.)

6.

Tatoi, Athens.

AusTRiA-HuxGARY : Haida, Arva, Boliemia, 1; Vasoiir, Eisenburg,
Hungary, 1 (U.S.N.M.).

Roumania : Gageni, Prahova, 2.

Greece: Corfu, 2; Cephalonia, 1; Tatoi, near Athens, 1.

Remarks. — Though at first sight very similar to Erinaceiis
europsens, 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.

Lord Lilford (p). 97. S. 14. 1.

Lord Lilford (p). 4. 4. 6. 15-lG.

(Type of species 4. 4. G. 16.)
J. I. S. Whitaker (p). S. 10. 1. G.

J. L S. Whitaker (p). 8. 10. 1. 7.

C. Mottaz (c). 8. 11. 8. 8.

ERINACEUS NESIOTES Bate.

I'JOG. Erinnceus curopfeus ncsiotes Bate, Proc. Zool. Soc. London, 1905, ir,
p. 316, April 5, 1906. Type in British Museum.

1910. Erinaceus eiiropievs nesiotes Trouessart, Paune Mamm. d'Europe,
p. 40.

Type locality. — Near Gonia, western Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Similar to Erinaceus nmmanicus 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. roumatticu.^. 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.

Remarks. — The Cretan hedgehog is nearly related to Erinaceus
roumanicus, though well differentiated by its smaller size and by
the relatively large fourth unicuspid tooth.

6, 9. Gouia, 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. 1.3.

* Type.

130

INSKCTIVOEA

ERINACEUS ALGIRUS Duveriioy and Lereboullet.

(Synonymy under subspecies.)

GeograpMcal distribution. — Northern Africa ; also in southern
Spain, southern France, and the Balearic Islands.

Diac/nosis. — Smaller than Erinaceus europseus (hind foot less
than 40 mm.) ; sjDines 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. 2b b).

External characters. — Externally Erinaceus alyiriis 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.

Coloxir.- — The colour resembles in general that of the paler
races of E. euroj.seus.

SlxulL- — The form of the skull is essentially as in E. < uropseus
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 visually 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
wtj and m.^ is completely destroyed. In the
upper jaw the third incisor, canine, and
a h first premolar are two-rooted, apparently

Fig. 2.5. without exception.

Large lower premolar of Measurements. — Head and body about

and'^.''aJ^(>Ms°(iO.''*x3*. 200 to 250 ; tail, 25 to 40 ; hind foot, 32 to
37 ; condylobasal length of skull, 54 to 59 mm.
Memarhs. — Thisspecies is readily distinguishable fromJG/•/J^ace^^s
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 southeini 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 aLgiuus algirus Duvernoy and Lereboullet.

1840. Erinaceus algirus Duvernoy and Lereboullet, M6m. Soc. Mus.

d'Hist. Nat. Strasbourg, in, fasc. 2, p. 4.
1898. Erinaceus algirus de Winton, Proc. Zool. Soc. Loudon, 1897, p. 955.

Tyjje local iiy. — Oran, Algeria.

Geographical distribution. — Northern Africa ; also in southern
Spain and south-eastern France,

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 buflfy white to base of hairs, except on
muzzle, cheeks, inten-amial 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, 20 ; 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 "0; lower tooth-
row, 22-0.

K 2

132

INSKCTIVOKA

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KRINACEUS

133

1. Audalucia, Spain. Lord Lilford (p). 94. 6. 11. 5.

6 juv. Elche, Alicante. G. S. Miller (c). 8. 8. 4. IG.

juv. al. Lecques, Var, France. Dr. P. Siepi (p). 98. 8. 25. 1.

Erixaceus aloirus vagans Thomas.

1901. Erinaceus alginis vagans Thomas, Proc. Zool. Soc. London, p. 38.

Type in British Museum.
1910. Erinaceus algirus vagans Trouessart, Faune Mamm. d'Europe, p. 39.

Type locality. — San Cristobal, Minorca, Balearic Islands.

Geographical distrihufion. — 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.

SjJecinicns examined. — One from ^lajorca and five from Minorca,
Balearic Islands.

BemarJcs. — 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 6, ?. San Cristobal, :Minorca. O. Thomas A- R. I. Pocock 0. 7. 1. 35-39.

(c & p).

(0. 7. 1. 36. Type of subspecies.)

134

CHIROPTER\

Order CHIROPTERA.

1779. Chiroptcra Blumeubach, Handbuch dei' Naturgeschiclitc, p. 74.

Geographical distrihution. — 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.

Memarhs. — ^The order Chiroptera, containing the only living
vertebrates, except birds, capable of true flight, is the most
sharply circumscribed of the main groups of mammals. Not
only are its living members invariably distinct from all other
recent forms, but the fossils also are, so far as known, 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 Slammals, p. IG,

November, 1872.
1875. Microchiroptera Dobson, Ann. and Mag. Nat. Hist., 4tli 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 distrihution. — The same as that of the order.
In Europe north to the limits of tree growth, west to Ireland
and the Azores.

MICROCHIROPTERA

135

Characters. — Anterior limb 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
Dfismodontidse) ; margin of ear not forming a ring ; tragus usually
present. Mammae in all European genera, j^ 1-1 = 2.

JRemarks. — The sub-order Microcluroptera is essentially cos-
mopolitan in distribution. Though more highly modified than
the Megacliiroiitera 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 MICROCHIROPTERA.

Tragus absent ; muzzle with leaf-like outgrowths ;
premaxillaries represented by palatal branch
only, not fused with surrounding parts (often

lost in prepared specimens) Rliinolophidai, 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 Molossidx, 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 Vcspcrtilionidee, p. 165.

Presternum with median lobe nruch smaller
than body of bone ; coracoid curved out-
ward ; second phalanx of third finger not

specially elongated Vespertilion'um, p. 165.

Presternum with median lobe lai-ger than
body of bone ; coracoid straight, directed
inward ; second phalanx of third finger
nearly three times as long as first Minioptennie, p. 26S.

136

CHIROPTERA

KEY TO THE GENERA OF EUROPEAN BATS.

(A wholly artificial key based primarily on external characters.)

Muzzle with leaf-like outgrowths RJnnolophus, p. 137.

^luzzle without leaf -like outgrowths.

Tail projecting conspicuously beyond membraue... Nijctinomus, p. 27G.
Tail not projecting conspicuously beyond membrane.
Ears joined.

Ear longer than head Plccotus, p. 256.

Ear shorter than head Barbastella, p. 263.

Ears separate.

Second phalanx of third finger nearly three

tiines as long as first Minioptcrus, 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 Nydalus, p. 242.

Fifth finger much longer than metacarpal of
fourth or third.
Ear wider than high, its lower margin
forming a small pocket near angle of

mouth Vesperfilio, p. 238.

Ear higher than wide, its lower margin
not forming pocket near angle of
mouth.

Upper cheek-teeth 6-6 Mijotis, p. 166.

Upper cheek-teeth less than 6-6.

Upper cheek-teeth 5-5 Pipistrcllus, p. 202.

Upper cheek-teeth 4-4 Eptesicus, p. 224.

Family RHINOLOPHID^.

1827. Rhinolophina Lesson, Man. de Mammalogie, p. 81 (part).
1857. Phyllostomata Blasius, Siuigethiere Deutschlauds, p. 26.
1866. Bhinolopliidm Gray, Proc. Zool. Soc. London, p. 81 (part).
1878. Bhinolophidie. Dobson, Catal. Chiropt. Brit. Mus., p. 100 (part).
1907. Rliinolopliidx IMiller, 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 Phihppine Islands.
Solomon Islands and north-eastern Australia. In Europe north
to northern England and the Baltic coast of Germany.

Characters. — Ear without tragus ; muzzle w ith 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.

Bemarks. — The Rhinolophidse are the most widely distributed

KHINOLOPHUS

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.

(ienus RHINOLOPHUS Lacepede.

1799. Rhinoloplius Lacepede, Tabl. des div. sousdiv. ordres et genres des

Mammiferes, p. 15 (ferrjim-equiniini).
1836. Bhinocrepis Gervais, Diet. Pittoresque d'Hist. Nat. iv, pt. 2, p. GIT

(attributed to GeofEroy and Cuvier, Mag. Encyclopedique, 1795,

but the name does not occur in the paper alluded to).
1847. Aquias Gray, Proc. Zool. Soc. London, p. 15 {luetics and trifoliatiis).
1857. Rhinolopluis Blasius, Saugethiere Deutschlands, p. 26.
1866. Phyllotis Gray, Proc. Zool. Soc. London, p. 81 {'phiUjjjx-nsis) not

Phyllotis Waterhouse, 1837.
1866. Coslophijllus Gray, Proc. Zool. Soc. London, p. 427 (ccelopJiyllus).
1878. Rhinolophus Dobson, Catal. Chiropt. Brit. ]\Ius., p. 100.
1901. Euryalus INIatschie, Sitz.-Ber. Gesellsch. naturforsch. Freunde,

Berlin, p. 225 {meliclyi).
1904. Euryalus Matschie and Andersen, Sitz.-Ber. Gesellsch. naturforsch.

Freunde, Berlin, p. 71 (curyale group).
1907. Rhhiolopihus Miller, Families and Genera of Bats, p. 108, June 29,

1907.

Type species. — Vespert'dio ferriim-equinum tSchreber.

Geographical distribution. — Same as that of family (p. 136).

Characters. — Dental formula : / ^^^7, ) '' \E\ j 1^'"^ ^', ) "' |55 — •^'-^
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 (jjm^) and
middle lower premolar (pm^) small, functionless, usually crowded
(juite out of tooth-row. Other teeth showing no special peculiari-
ties ; m^ and m'" without hypocone, m^ with live cusps and
three commissures (in many species a rudimentary fourth com-
missure), the crown area much more than half that of m^ or
m'^. Skull with large brain-case and much shortened, globularly
inflated rostrum, beyond which the maxillaries, bearing the large
canines, conspicuously project ; palate so deeply emarginated
both anteriorly and posteriorly that its median length is less
than least distance between tooth-rows. Tail well developed,
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 (tig. 26).

Remarks. — Among European bats the members of the genus
Rhinolophus are at once recognizable by the presence of the nose-

138

CHIROPTERA

leaf and absence of tragus. Tlie skull differs from that of all
other members of the fauna in tlie short, globularly inflated
rostrum and long, projecting maxillaries, between which lie the

M

c ^ d

Fig. 26.

Noseleaf of Rh'uwloiihv>i ferrfim-eqninum (a), R. hipposideros (W)., R. euryale (c),
and R. blasii (d). JTat. 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 FORMS OF RHINOLOPHUS.

Noseleaf with connecting process broadly
rounded above ; skull with nasal sweUiugs
long, rising gradually above line of fore-
head.
Forearm over 50 mm. ; condylobasal length
of skull over 20 mm. ; sella pandurate ;
large upper premolar in contact with

canine (Greater Horseshoe) B. ferrum-equimim, ]^. 139.

Wing relatively long ; forearm 54-58 mm.,
longest finger 84 to 92 mm. (Southern

and central Continental Europe) H. f. ferrum-equinum, p. 142.

Wing relatively shorter ; forearm 52 to
55 mm., longest finger 83 to 88 mm.
(England) R. f. insulanus, 'p. 147.

* See Andersen, Ann. and Mag. Nat. Hist., 7th ser., xvi, pp. 618-662.

RHINOLOPHUS 139

Forearm under 43 mm. ; coudylobasal length
of skull under IG mm. ; sella cuneate ;
large upper premolar not in contact

with canine (Lesser Horseshoe) B. hlpposidcros, p. 147.

Greatest length of skull 14 -5 to 15' 5 mm.

(Mediterranean region) jt. li. muiimus, p. 151.

Greatest length of skull more than
15 "5 mm.
Forearm 36 -.3 to 39 mm. (England and

Ireland) /?. /,. minutus, p. 154.

Forearm 89 to 41 -7 mm. (Central Europe) R.h. 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 88) ; point of lancet gradually

narrowed, never linear R. euryalc, p. 155.

Size larger, forearm 48 '6 to 51-4, upper
tooth-row about 7 nnn. ; gradation
between phalanges of fourth finger
less abrupt (ratio of first to second
about 44) ; point of lancet linear R. mchclyi, p. 159.

RHINOLOPHUS FERRUM-EQUINUM Hchreber.

(Sjmonymy under subspecies.)

GeograpJiical distrihuiion. — From southern Japan and China,
through the Hiniahiyas, the Mediterranean sub-region (exclusive
of Egypt), and central Europe to southern England (Andersen).

Diagnosis. — 8ize 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 bufiy brown resembling the wood-brown of Eidgway. While
intermediate shades occur the extremes are more often met with,
evidently representing two dichromatic phases. Underparts
usually rather lighter than back but neA^er approaching whitish.
Skull. — General outline of skull long and narrow, the breadth
of brain-case much less than twice that across canines, and con-
tained about 2^ 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
bulke 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 jjostero-
internal angle of )*('- ; palate with an
evident emargination on each side ex-
tending between m^ and mesopterygoid space ; anterior palatal
emargination extending back to line joining protocones of
anterior molars ; premaxillary ligulate, somewhat wider pos-
teriorly than anteriorly, the 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

'r^\r:^eEa^

Rhinolophusferrum-equinum.
Nat. size.

RHINOLOPHUS

141

bottom of jDalatal emarginatioii 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 ipiddle 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 ^^remolars and upper incisors the
teeth are robust and heavy I'elatively 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 i^remaxillar}^, 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^ and m- there is a low-lying
heel (best developed in m^), but no indication of a true hypocone :
paracone and metacone well developed, the latter slightly the
higher in m^ and m-, this reversed in m^ ; styles and commis-
sures in m^ and m- well developed, forming a normal VV-pattern ;
in m^ the metastyle and fourth commissure are absent, and the
third commissure is much reduced in length ; crown area of m'

U:

OHIROPTERA

about two-thirds that of vr ; lower molars with protoconid
noticeably higher than the other cusps, the posterior triangle
slightly the wider of the two in m^ and m.^, a little the narrower
in m^ ; 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 mai'gin of the connecting process of sella.
In R. mchehji, 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.

Rhinolopiius ferrum-equinum ferrum-equinum Schreber.

1774. Vespertilio ferrum-equimim Schreber, Saugthiere, i, pi. lxii, upper

figures ; description, i, p. 174 under name : Die Hufeisennase

(part). France ; based primarily on Daubenton.
1776. Vespertilio equinus P. L. S. ?\Iuller. Natursyst. Suppl. u. Kegist.-

Band, p. 20 (part), France.
1779. [Fesper^iHo] j3e?'S2.iiciH«f'UsBlumeubach, Handb. d. Naturgesch., p. 75,

(part : included the leaf-nosed bats of Europe and South America).
1785. [}^es'pertilio'] ungula Boddaert, Elenchus Animalium, i, p. 71

(Burgundy) .
1792. Vesp\ertilid] fcr\_rum']-cqii\j,num'] major Kerr, Anim. Kingd., p. 99

(not y. molossus major Kerr, I.e., p. 97), France.
1798. Ves2)e7-tilio hippocrepls Schrank, Fauna Boica, i, p. 64 (Renaming

of ferrum-equinum Schreber).
1803. Bhinolophus major Geofiroy, Catal. Mamm. Mus. Xat. d'Hist. Nat.,

Paris, p. 56 (Burgundy).
1813. Bhinolophus unihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris,

XX, 10. 257 (France).
1829. '> Bhinolophus unifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ.

Thierwelt, i, ]0. 104 (nomen nudum).
1857. Bhinolophus ferrum-equimim Blasius, Saugetliiere Deutschlands,

p. 81.
1863. [BhinolopMis fcrrnm-eguinum'] a var. germanicus Koch, Jahrb. des

Vereins fiir Naturkunde im Herzogthum Nassau, xviii, p. 522

(Wiesbaden, Hessen-Nassau, Germany).
1863. [^Bhinolophus ferrum-equinum^ B var. italicus Koch, Jahrb. des

Vereins fiir Naturkunde im Herzogthum Nassau, xviii, p. 523

(Italy).
1878. Bhiiwlophus fciruni-eijuiitum Dobson, Catal. Chiropt. Brit. ]\Ius.,

p. 119 (part).

1885. Bhinolophus unihastatus, var. homorodahnasiensis Daday, Orvos-

Termeszettudomanyi Erteseto, Kolozsvar, x, p. 274 (Homorod-
Almas cave, Hungary).

1886. Bhinolophus imihastatus var. homorodalmasiensis Daday, Verhandl.

u. Mittheiluugen des Siebenbiirgischen Vereins fiir Naturwissensch.
in Hermannstadt, xxxvi, j). 79.

1887. Bhinolophus ferrum-equinum var. homorodensls Daday, Ertekezesek a

Term^szettudomanyok K6r6bi'il, Budapest, xi, pt. 7, p. 13
(Renaming of homorodalmasiensis).

KHINOLOPHUS

143

1904. Rhinoloplius ferrum-eqiunniii, obsciirtts Cabrera, Mem. Soc. Espau.

Hist. Nat. Madrid, ii, p. 257 (Valencia, Spain).

1905. Bhinolophus fcrrum-ecininuvi typicits Andersen, Proc. Zool. Soc.

London, 1905, ii, p. 113, October 17, 1905.

1905. Bhinolophus fm-um-equintmi obscurus Andersen, Proc. Zool. Soc.
London, 1905, ii, p. 116, October 17, 1905.

1910. BhiiwlopJms ferrmn-cquinum, B. fcrncm-equinum obscurus and B.
ferrum-cqiiinum homorodcnsis Trouessart, Faune Mamm. d'Europe,
pp. 4-5.

Ty^pe 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.

3Ieasurements. — 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, France : tibia.,
24-1 (23 -4-24 -6); foot, 12-1 (11 •8-12-8); forearm, 55-4
(54-65) ; third finger, 87-3 (84-91) ; fifth finger, 71-5 (69-74).
Female from Marseilles, France : forearm, 57. Male and female
from near Genoa, Italy : forearm, 57 and 57. Two females from
Rimini, Itah* : 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
Tiibingen, Wiirtemberg, Germany : head and body, 63 ; tail, 37 ;
tibia, 24 • 4 ; foot, 1 1 ; 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 cxamlnecl. — One hundred and forty-one, froni the following
localities : —

Portugal: Cintra, 1.

Spain : Silos, Burgos, 22 ; Granada, 4 ; Elche, Alicante, 3 ; San
Cristobal, Minorca, Balearic Islands, 4.

144

CHIROPTEHA

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CHIROPTKRA

Fkance : Troubate, llautes-Pyren^es, G ; St. Genies, Gard, 48 (INIottaz) ;
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, Ligiiria, 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,!;
Sicily, 2.

Corsica : Commune di Barbaggio, 1.

Sardinia : Sassari, 1.

Geeece : Parnassus region, 1 (U.S.N.M.) ; Island of Syra, Cvclades,

1 (U.S.N.M.) ; Mt. Goria Monastery, Crete, 8.

Germany : Tiibingen, Wiirtembej'g, 1.

Switzerland : Geneva, 6 (U.S.N.M. and Mottaz) ; Boudry, Neuchatel,

2 (JMottaz) ; Tremona, Ticino, 1 ; Mendrisio, Ticino, 2 ; Lugano, Ticino, 3.

.\ 1"STRIA-Hungary : Herkulesbad, Hungary, 6 ; Ofener Mts., Hungary, 1.

HemarJcs. — Spanish specimens of Ithinolophua ferruia-equinutii
have been regarded by both Cabrera and Andersen at i-epresent-
ing a pecuHar race, B. f. ohscurus, 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.

6 al. Silos, Burgos, Spain.

2 6,2 9. Silos, Burgos.

6, 9. Granada, 2250 ft.

2 6, 9. Elche, Alicante, 20 m.

<5, 2 9. San Cristobal, Minorca, Ba-
learic Islands.

S al. San Cristobal, INIinorca.

G 6. Trovibate, Hautes-Pyrenees,

France. {A. Robert.)
1 al. IMeounes, Var.

9 al. Turin, Italy.
9 al. Isoverde, Genoa.
Skeleton. Italy. {Prince Bonaparte.)
2 c5 al. Sicily.

Barbaggio, Corsica.

<^al.
(5al.

Sassari, Sardinia.

.6, 2 9. Mt. Goria Monastery, Crete.
9 al. Tubingen, Wiirtembcrg, Ger-
many.
9. Tremona, Ticino, Switzerland.

{E. H. Zollikofer.)
2 9. Mendrisio, Ticino. {E. H.

Zollikofer.)
8. Lugano, Ticino. [E. H. Zolli-

kofer.)
•2 6, 4 9. Herkulesbad, Hungary.

9 al. Ofener Mts., Budapest.

0. Thomas (c&p).
N. & S. Gonzalez

(c).
G. S. Miller (c).
G. S. Miller (c).
G. S. Miller (c).
0. Thomas and

R. I. Pocock

(c & p).
0. Thomas and

R. I. Pocock

(c & p).
0. Thomas (p).

Dr. K. Jordan

(c & p).
Prof. Bonelli (p).
O.Thomas (c & p).
Tomes Collection.

Dr. C. I. Forsyth

Major (c & p).
Marquis G. Doria

Miss D. Bate (c).
Dr. A. Giinther

0. Thomas (p).

0. Thomas (p).

0. Thomas (p).

Hon. N. C. Roths-
child (p).
Budapest Museum

(E).

'J8. 2. 2. 1.
S. 7. 7. 38-48.

8. 8. 4. 1-4.
8. 8. 4. 8-9.
8. 8. 4. 5-7.
0. 7. 1. 24-2G.

0. 7. 1. G8,

G. 4. 1. 1-6.
8. 3. 15. 1.

88. 12. 7. 1.
7. 1. 1. 727.

G. 4. 14. 1.

6. 12. 1. 11.

5. 12. 2. 1-3.
G6. 2. 1. 1.

2. 8. 4. 1.

2. 8. 4. 2-3.

4. 4. 5. 1-8.

7. 'J. 16. 1-6.

94. 7. 18. 5.

EHINOLOPHUS

u;

Rhinolophus ferrum-equinum insulanus Barrett-Hamilton.

1910. ' BJiUiolophits ferrum-eqiiinum insulatizos Barrett-Hamilton, Ann. and
Mag. Nat. Hist., 8tli ser., v, p. 292, March, 1910. Type in British
IMuseum.

1910. Rhinoloplms fernmi-eqidnum insulanus Trouessart, Faune Mamm.
d'Eurojpe, p. 273.

Ti/pe localiti/. — Cheddar, Somersetshire, England.

(TCtujraplncal distribution. — Central and southern England.

DItK/nosis. — ^Wing relatively short ; forearm, 52 to ^)b ; third
linger, 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, ?. Cheddar, Somerset, J. A. Coward (c & p). 7. 1. 10. 1-2.

England.

G al. Cheddar. J. A. Coward (c & p). 7. 1. 10. 8-10.

(5. 2 9al. Wells. Hon. N.C.Rothschild 5.1.23.1-3.

3 c5, c5 juv., 9 St. Wells. Hon. N.C.Rothschild /I. 9. 3. 1-4.

(p). 12.9.6.1.

6, ?. Wells. Hon. N.C.Rothschild 11. 1. 3. 3-4.

(P)-
2 9. Dorset. W. M. Hardy (c & p). 11. 1. 3. 1-2.

1. Bonchurch, Isle of Rev.C. A.Bury (c& p). 11.1.3.388.

Wight.

RHINOLOPHUS HIPPOSIDEROS Bechstein.

(Synonymy under subspecies.)

Geoyrapliical 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 fc) with bluntly cuneate sella
.and low, broadly rounded connecting process ; fourth finger with

L 2

148 CHIROPTERA

tirst 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 Wiinolophus ferrmii-equinum. General outline of
noseleaf narrower than in R. ferrum-equinum, the width of horse-
shoe less than distance from Hat area at base of sella to tip of
lancet ; sella narrowly rounded at tip, the sides straight, slightly
convergent above, the residting 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.
AVings and feet essentially as in _R. fcrrum-equinam.

Colour. — The colour does not differ appreciably from that of
Wiinolophus ferrum-equinum.

Skull. — The skull differs from that of Hhinoloplms ferrum-
equinum chiefl}^ 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 maxillar}'
region, the bi'eadth of former being about twice that of latter.
As the breadth of brain-case is contained distinctly less than
2 1 times in greatest length it is probable that the 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 jB. ferrum-equinum, particularly
at side of nares, but dorsal outline not rising abruptly above
level of interorbital region. Mesopterygoid fossa so conspicu-
ously widened anteriorly that it occupies almost entire extent of
palate between posterior molars, leaving no space for lateral
palatal emarginations, a peculiarity by which the skull may be
distinguished from that of all other European members of the
genus.

Teeth. — Though in general resembling those of Bhinolophus
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
fattened posteriorly, and onl}^ 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 B. ferrum-
equinum. Large upper premolar
and upper molars essentially as in
_B. ferrnm-eqninum except that in
)iv^ there is a distinct meta-
style and fourth commissure,
and area of tooth is nearly
equal to that of m"^. Lower
molars as in Rhmolophus
ferriim-eqwinnm, except that
in m^, 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.jCondylobasal length
of skull, 13 "8 to 15 mm.
(greatest length 14 • 5 to 16 • 2).
Details under subspecies.

BemarJcs. — Three imperfectly differentiated forms of Bhiiw-
lophus liipposideros occur in Europe : a larger central race, a
smaller Mediterranean race, and an intermediate form pecviliar
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.*

Rliiiwloplti

Fig. 28.

' hi'p2}Osideros.

Teeth X 10

Rhinolophus hipposideros iiipposidekos Bechstein

1789. Vespertilio ferrum-equinum B., Die kleine Hufeisennase, Bechsteiu.

Gemeinn. Naturgesch. Deutschlands, i, 1st ed., p. 186.
1792. Vesp[crtilio] fer[rum']-equ[imi»i'] minor Kerr, Anim. Kingd., p. 99

(not V. violossHS minor Kerr, I.e., p. 97) (France).
ISOO. Vespertilio hipposideros Bechstein, Thomas Pennant's Allgemeinc

Uebersicht der vierfiissigen Thiere, ii, p. 629.
1803. FlJiinolojjJms minor Geoftroy, Catal. JIamm. IMus. Nat. d'Hist. Nat.,

Paris, p. 57 (Neighbourhood of Paris).
1S13. Rhinolophus bihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xx,

p. 259 (Neighbourhood of Paris).
181G. ? 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.

150 CHIROPTERA

1S29. ? lihinolophus bifcr Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ,

Thierwelt, i, p. 104 (nomeu nudum).
1857. ? Rhinolophiis hipposideros Blasius, Saugethiere Deutschlands, p. 29.
1863. \_Rhinoloplms hipposidcros] a var. typus Koch, Jahrb. des Vereins fur

Naturkunde im Herzogthum Nassau, xviii, p. 530 (Wiesbaden).
1863. 'iRhinolophus hipposideros'\ |8 var. alpinus Koch, Jahrb. des Vereins

fiir Naturkunde im Herzogthum Nassau, xviii, p. 530 (Alps).
1870. Rlihwlophus cggenhoffner Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Slath.-Naturwiss. Classe, lxi, Abth. i, p. 151

(MS. synonym of bihastatus).
1878. RhinolopJius hipposideros Dobson, Catal. Chiropt. Brit. Mus., p. 117.

1885. Rliinolophus bihastatus var. kisnyiresiensis Daday, Orvos-Termesz-

ettudomanyi Ertesito, Kolozsvar, x, p. 274 (Kis-Nyires, Szolnok-
Dobaka, Hungary).

1886. Rhinolophus bihastatus var. kisnyiresiensis Daday, Verhandl. u.

Mittheilungen des Siebenbiirgischen Vereins fiir Naturwissensch.
in Hermannstadt, xxxvi, p. 80.

1887. Pthinolophus hipposideros var. troglopJiilus Daday, Ertekezesek a

Term6szettudomd,nyok Korebol, Budapest, xvi, pt. 7, p. 8 (Re-
naming of kisnyiresiensis).

1904. Rhinolophus ejiryale helvetica Bretscher, Vierteljahrsschrift der

Naturforsch. Gesellsch. in Ziirich, xlix, p. 256. See Mottaz,
Bull. Soc. ZooL, Geneve, i., p. 172, 1908 (Baar, Zug, Switzerland).

1905. PJtinolophus hipposiderus typicus Andersen, Proc. Zool. Soc. London,

1905, II, p. 141, October 17, 1905.
1910. Rhinolophus hipposiderus Trouessart, Faune Mamm. d'Europe, p. 9.

Type locality. — France.

Geoijrn/pMcal 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 : iifth 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) : IST. Bulgaria (1), 39 ; Eoumania
(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 col lection), J
38-5 to 40-6. Four males from the same locality (U.8.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 From back of occiput to front of canine.

X Measured by Chas. Mottaz (base of metacarpals included) and verified
by Gerrit S. ^filler.

RHIN^OLOPHUS 151

Specimens examined. — One huudred and seventeen, from the following
localities : —

France : Dions, Gard, 1 (Mottaz) ; St. Cergues, Haute-Savoie, 1.

Germany : Strass, near Burgheim, Bavaria, 3 ; JMainz, 1 (Strassliurg) ;
Bitsch, Alsace, 1 (Strassburg).

AusTRiA-HiNGARY : Hatszeg, Hunyad, Hungary, 1.

Switzerland: Near Geneva, 94 (B.M., U.S.N.M. and Mottaz); St.
^NToritz, 1; Thayngen, Schaffhausen, :2 (U.S.N.M.) ; Eoggwil, Thurgau,
7 (B.M. and U.S.N.M.); Canton Thurgau, 5 (B.M. and U.S.N.M.).

6. St. Cergues, Haute-Savoie, A. Roi^ert (c & p). 5. 4. 9. 1.

France.
2 9. Burgheim, Bavaria, 375 m. Lord Lilford (p). 11. 1. 1. 117-

Germany. 118.

6. Hatszeg, Hunyad, Tran- C.G.Danford (c). 3.11.8.1.

sylvania.
3d. Grand Pre, Geneva, Swit- C. Mottaz (c & r). G. 2. G. 1-3.

zerland.
1. St. Moritz, Grisons. Leon O. Galliard 75. 9. 20. 3.

(c & p).
3 S, 2 9, imm. Roggwil, Thurgau. {E. H. 0. Thomas (p). 2. 8. 4. 4-9.
Zollikofer.)
9al. Thnrgau^ {E.H. Zollikofer.) 0. Thomas (p). 2.8.4.55.

Rhixolophus hipposideros minimus Heiiglin.

18G1. lihinolophns minimus Heuglin, Nov. Act. Acad. Caes. Leop. -Carol.,

XXIX, Abliandl. viii, p. 6 (articles separately paged) (Keren,

Abyssinia).
1863. [RhinolopJtus liippoaideros"] y -va.r. jviUidus Koch, Jahrb. des Vereins

fiir Naturkunde im Herzogthum Nassau, xviii, p. 531 (^Jlediter-

ranean region).
1904. llliinolophus pliasma Cabrera, ]\Iem. Soc. Espafi. Hist. Nat., Madrid,

II, p. 252 (Madrid, Spain).

1904. FJi[inolop]i.us] h\^ipposiderus] minimus Andersen, Ann. and ^lag. Nat.

Hist., 7th ser., xiv, p. 456, December, 1904.

1905. Rhinoloplius hipposiderus minimus Andersen, Proc. Zool. Soc.

London, 1905, ii, p. 140, October 17, 1905.
1910. PJiinolopJius hipposiderus minimus Trouessart, Faune Mamm.
d'Europe, p. 10.

Type locality. — Keren, Abys.sinia.

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 ( 9 )? 35 • 2 ; Rome (^),
36-4; Siena (3 ^), 36-8 to 38; Marseilles, France (9), 38;
Dions, Gard, France (<^), 34-8, (9)? 36-6 Two females from
Ax-les-Thermes, Ariege, France, 36 "6 and 38*4. Two males
from Silos, Burgos, Spain, 36 and 37. Female from the same
locality, 38. Five males from Ticino, Switzerland, 37 "0
(36*2-38). Mottaz collection: St. Genies, Gard, France (15,

152

CHIROPTERA

Observations.

Teeth slightly worn.
,, not worn.

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,, slightly worn.

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Germany : Strass, near Burg
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R. hipposideros minut

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both sexes), 36 • i to 38 • 2) ; Sardinia (2 9 ), 37 • 4 to 37 • 8. For
cranial measurements see Table, p. 153.

Specimens examined. — Eighty-one, from the following localities : —

Spain : Inca, Majorca, Balearic Islands. 1 ; San Cristobal, Minorca,
Balearic Islands, 3; Elche, Alicante, 2; Silos, Burgos, 5; Seville, 1.

Portugal: Cintra, 1.

France : As-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.); Faenza, 1
(U.S.N.M.) ; Eome, 3 ; Ostia, Rome, 2 ; Ficuzza, Sicily, 6.

Sardinia : No exact locality, 2 (Mottaz) ; Zimmigas, Siliqua, 2.

Corsica: Diutorni di Patrimonio, 3; no exact locality, 1.

Malta: 7.

?. Inca, Majorca, Balearic O. 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 & p^

?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 11.

6. Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 12.

<5 al. Cintra, 500 m. Portugal. 0. Thomas (c & p). 98. 2. 2. 57.

S, ?. Ax-les -Thermes, Ariege, G. S. Miller (c). 8. 8. 4. 124-

2400 ft. France. 126.

6. Rome, 37 m. (C. Coli.) G.Barrett-Hamilton 11.1.2.42.

2. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 5-6.

6?. Ficuzza, Sicily. {A. Robert.) 0. Thomas (p). I s' 9 1 1-2

6, 9 al. Zimmigas, Siliqua, Sardinia. Marquis G. Doria (p). 6. 12. 1. 20-21.

(R. Meloni.)
3 6,2 9. Malta. (-/. Micalleff.) Lord Lilford (p). 11. 1. 1. 119-

123.
2. Malta. Lord Lilford (p). 95. 3. 2. 1-2.

Rhinolophus hipposideros iiiNUTUs Montagu.

1808. Vespertilio minutns Montagu, Trans. Linn. Soc. London, ix, p. 163.
1905. Rhinolophus hipposiderus mimitus Andersen, Proc. Zool. Soc.

London, 1905, ii, p. 142, October 17, 1905.
1910. Rhiiiolopluis hipposiderus juimitus Trouessart, Faune Mamm.

d' Europe, p. 10.

Type locality. — Wiltshire, England.

Geographical distrihutiou. — 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.

RHINOLOPHUS

155

3 6. St. Asaph, Denbighshire, Charles Oldham (c & v). 11. 1. 3. 5-7.
Wales.

1. Conway, Carnarvonshire. Sir W. Jardine (c \- p). GO. 9. 17. 1.

9. 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.)

6 al. Hope End, Herefordshire. N. C. Hewitt (c it p).

6 al. Wells, Somerset. S. Lewis (c & p). 5. 1. 24. 1.

1. Devizes, Wiltshire. J. E. Harting (c & p). 87. 2. 21. 1.

6, ? al. Zeals, Wiltshire. F. Norgate (c & p). 4. 11. 6. 1-2.

■1 ?. Devonshire. Osley Grabham (c & p). 11. 1. 3. 8-9.

3. Eagley House, Warwick- Tomes Collection. 7. 1. 1. 289-
shire. 291.

RHINOLOPHUS EURYALE Basius.

1853. Bhinoloplms curijalc Blasius, Wiegmann's Archiv fiir Naturgesch.,

1853, I, p. 49 (Milan, Italy).
1857. Bhinolophus euryale Blasius, Siiugethiere Deutschlauds, p. 35.
1878. Bhinolophus euryale Dobson, Catal. Chiropt. Brit. ]\Ius., p. 116.
1904. E\iiryalus] 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\jiryalus] cabrene Andersen and ilatschie, 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. — ^Milau, Italy.

Geographical distribution. — Southern Europe from Portugal
to Greece, north to Hungary and central France.

Diagnosis. — Size medium, forearm, 44*6 to 49, condylobasal
length of skull, 16 '4 to 18, mandible, 12 to 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 Mhinolophxis ferrum-equimim ; sella parallel
sided, rounded off above, connecting process sharply linear-
pointed, rising conspicuously above sella ; lancet Avith slight con-
striction above middle, beyond which the tip narrows graduall}' 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 Ithinolophns ferrum-equinmn

156

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and Ji. hijjjjosUleros ; antitragal lobe about half as high 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, 66 mm.; second phalanx, 17-4 mm.;
ratio of jSrst to second, 38 +). Foot averaging slightly more
than half as long as tibia.

Colour. — While essentially as in Ttldnoloplim ferrum-equinum
and B. hipjiosideros, 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
i?. mehelyi.

SJmll. — In both size (greatest length about 19 mm.) and form
the skull is somewhat intermediate between that of Bhimloiylim
ferrum-equumm 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 cochlea; less narrowed than in the preceding
species. Nasal region slightly less inflated than in B. Mpposideros,
and inflated area relatively shorter, its posterior border rising
abruptly above interorbital level.

Teeih. — In all respects the teeth closely resemble those of
B III noloph us 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.

Fbanck : St. Paterne, Indre-et-Loire, d (B.M. and U.S.N.M.) ; St. Genies,
Gard, about 50 (ilottaz) ; Gapeau River, Var, 12.

Italy: Near Genoa, 33 (B.M., U. S.N. :M., Genoa, and Mottaz) ; Monte
Pisanino, 2 ; Siena, 3 (U.S.N.M.) ; Rome, 2 ; Velletri, Rome, 5 (U.S.N.M.) ;
Nicotera, Calabria, 1 ; Marsala, Sicily, 2.

Sardinia : Mount Gennargeutu, 3 (U.S.N.M.).

Austria-Hungary: Ofener Mountains, 2; Orsova, 1.

Dalmatia : Zara, 1.

Geef.ce : Missolungi, Acarnania, 2 (U.S.N.M.).

Bemarks. — Bhinolophus euri/ale 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 bv the form of the lancet and

EHINOLOPHUS

159

aiititi'agus, together with its rather smaller size and' usually
darker colour. Witli the material at hand I am unable tt)
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, 500 m. Portugal. O. Thomas (c & p). 98.2.2.2-3.

2 6,2 9 al. Cintra, 500 m. O. Thomas (c & p). 98. 2. 2. 5-3-50.

9 al. Villalba, Lugo, N.W. Dr. V. L. Seoane (p). 94. 1. 1. 1.
Spain.

6, 9 al. Madrid. A. Cabrera (p). 5. 2. .3. 1-2.

1 al. St. Paterne, Indre-et- Royal Army Medical 9. 1. 4. 9.

Loire, France. College (p).

9 al. St. Paterne, ludre-et- G. E. Dobson (p). 80. 12. 14. 3.

Loire.

9 al. Gapeau River, Var. Dr. K. Jordan (c & p). 8. 3. 15. 2-10.

3 J, 9 al. Finalborgo, Liguria, Marquis G. Doria (p). 6.12.1.14-17.
Italy. {A. Gagcro.)

2 al. Monte Pisanino, Liguria. Lord Lilford (p). 73.1.8.6.

2 6. Rome. (C. Coll.) G. Barrett-Hamilton 11. 1. 2. 40-41.

6 al. Nicotera, Calabria. Florence Museum (e). 85. 7. 6. 1.

2 6. Marsala, Sicily. (.1. 0. Thomas (p). G. 8. 4. 10-11.
Robert.)

i, 9 al. Ofener Mts., Budapest. Budapest Museum (e). 94. 7. IS. 2-3.

9. Orsova, Hungary. Hon. W. Rothschild 7. 9. 16. 7.

9. Zara, Dalmatia, 50 m. Lord Lilford (p). 11. 1. 1. 128.

(Kolomhatovic.)
1 al. S. Europe. Purchased (Parreys). 47. 5. 27. 44.

RHINOLOPHUS MEHELYI Matschie.

1901. Rhinolophus mehclyi Matschie, Sitz.-Ber. Gesellsch. Naturforscli.

Freunde, Berlin, p. 225 (Bucharest, Roumania).
1904. Rhinolophus carpctantis Cabrera, Mem. Soc. Espafi. Hist. Nat., ii,

p. 254 (Madrid, Spain).
1910. Rhinolophus euryalc mehclyi and R. carpetamis Trouessart, Faune

Mamm. d'Europe, pp. 7-8.

Type locality. — Bucharest, Roumania.

Geograpliical distribution. — Roumania, southern France
(Gard), Sardinia, central Spain. Details of distribution not
known.

Diagnosis. — Like Bhinolophis euryale but larger (forearm,
■18 '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 j^halanx
decidedly more than one-third as long as second (ratio about 44) :
colour usually paler than in the related animal.

:i External characters. — Slightly larger and more robust than
Bhinoloplais euryale, a difference especially noticeable in freshlv
killed individuals. Noseleaf as in R. euryale, except that the
lancet is very abruptly narrowed above middle to a distinctlv
linear tip. Ear as in B. enryale but broader, the antitragal lobe

160

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162 CHIROPTERA

scarcely half as high as conch, its width slightly greater 'than
height. Wing difiering from that of the related animal in the
less degree of shortening of the first phalanx of fourth linger as
compared with second (average of ten specimens from Gard,
France, first phalanx, 8' 1 ; second phalanx, 18*0 ; ratio of first
to second, 44 +)• Foot as in It. euryale.

Colour. — Though not invariably distinguishable the colour is
usually paler than that of Bhinolophus euryale, a difference
especially noticeable in the region between ears, on sides of face
and neck, on chin and throat, and along median portion of chest
and belly, all of which are frequently a very pale almost whitish
drab-grey.*

Skull and teeth. — Except for its slightly greater average size
(greatest length about 20 mm.) the skull agrees with that of
Mhinoloplius 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) .

France: Near St. Genies, Gard, about 50, skins and in flesh (Llottaz).

Sardinia: Sassari, 3 (B.M. and U.S.N.M.).

Koumania: Bucharest, 1 (U.S.N.M.) ; Dobrudscha, 1 (Mottaz).

Bemarks. — At first sight this species appears very similar to
Mhinolophus 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 Bhinolophus 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 E. carpetanus Cabrera.)
2 6 al. Sassari, Sardinia. Marquis G. Doria(p). 6. 12. 1. 18-19.

RHINOLOPHUS BLASII Peters.

1857. PJiinoloplms clivosus Blasius, Saugethiere Deutschlands, p. 33. Not
of Riippell, 1824 (Italy, Sicily, Istria and Dalmatia).

1866. Ilhinoloplncs blasii Peters, Monatsber. k. Akad. Wissensch. Berlin,
p. 17 (Renaming of clivosus Blasius).

1878. Rhinolophus blasii Dobson, Catal. Chiropt. Brit. Mus., p. 117.

1910. Rhiiwlophus hlasiusi Trouessart, Faune 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 Hhinolophus euryale ; nose-

* When seen by candle-light flying in caverns these bats are said to
appear entirely white.

RHINOLOPHUS

163

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Number.

Mottaz
122133

6. 12. 1. 18

6. 12. 1. 19
86536

Hu. 143 Mottaz
„ 144
M 145
„ 149
„ 151
,, 415
„ 444
» 146
„ 152
„ 433
5.2.3.1*

94. 12. 1. 1

94. 12. 1. 4

37508

Locality.

R. mehelyi.

Eoumania: Dobrudscha .
Bucharest .
Sardinia : Sassari .

France : near Nimes, Gard

Spain : near Madrid

R. blasii.
Cyprus ....

Greece : Nauplia

164 CHIROPTERA

leaf with cuiieate sella and very high, sliari)ly pointed cunnecting
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 1 have examined
is in bad condition. It indicates that the colour is not essentially
different from that of Rhinolophus euryale.

Shiill. — In general the skull resembles that of Bhinolojjhns
euri/ale, 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 oft' the occipital region from rest of brain-cast>.

Teeth. — Incisors, canines and molars as in Hhinolojjhns 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 dift'ering from those of all the other European members
of the genus in the approximately equal crown areas of the two
larger teeth, the antei'ior subterete, the posterior with trapezi-
form section ; shaft of posterior tooth with diametei' in axis of
tooth-row much less than transverse diametei', the cusp when
viewed from the side appearing to rise from middle of crown with
noticeable flat area before and behind it.

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 crown,
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 localitv, 4.
Geeece : Nauplia, 1 (U.S.N.IM.).

ReiiiarJis. — This species is so readily distinguished from the
othei- 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.

vespertilionin.e 165

Family VESPERTILIONID^.

1821. Ves2yeiiilio)>idui Gray, London Med. Repos., xv, p. 299, April 1, 1821

(part).
1857. ^^rsjicrtiliones Blasius, Saugetliiere Deutscblands, p. 37.
1878. Ycspertilionidx Dobson, Catal. Chiropt. Brit. Mus., p. 167 (except

the genera Natalus and Thyroptcra).
1907. Vcspcrfilionidx ^liller, Families and Genera of Bats, p. 195, June 29,

1907.

(jreiH/raphical 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.

GJiaracters. — Ear with tragus ; muzzle without distinct leaf-
like outgrowths ; skull with premaxillaries represented bv nasal
branches only, the two bones very early fused with surrounding
parts ; median length of palate greater than least distance between
tooth-rows ; auditoiy bulla not emarginated on inner side ; shoidder
girdle normal, without fusion of its elements ; secondary
articulation of humerus w^ith scapula better developed than in
the RJiinoJopliidse ; tibula very slender, not adding appreciably to
strength of leg ; foot normal, the toes slender ; tail not project-
ing conspicuously beyond membrane.

Remarhs. — This family is the most widely distributed grouj.)
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 VESPERTILIONINiE.

1878. Yi'spcrtiliones Dobson, Catal. Chiropt. Brit. Mus., p. 168 (except

genera Kcrivoula and Harpiocephalus) .
1907. Vcspertilioniiix Miller, Families and Grenera of Bats, p. 197, June 29,

1907.

Gi'ograpliical distribution. — Same as that of the family
Vespet-tilionidse.

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 Ronaldshay, Orkney Islands, but the
occurrence seems open to question (see Wolley, The Zoologist, vii, p. 2343,
1849; VIII, pp. 2095-96, 2813-14, 1850; Barrett-Hamilton, Hist. Brit.
Mamm., i, pp. 222-224, March, 1911).

A specimen of another North American member of this genus
{N. horealis), bearing the label: " Yilleveque (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 CHIROPTERA

dorsal ; scapula with coracoid curved outward ; nostrils simple ;
lower incisors in all known genera, 3-3.

Bemarks. — The sub-family Vespertilioninae contains all but
eight of the known genera of Vespertilionidse, and all but one,
3Iinio_pterus, of the eight found in Europe. It is the central,
least specialized portion of the family.

KEY TO THE EUROPEAN GENERA OF VESPERTILIONID^.

Cheek teeth 5i? Myotis, p. 166.

Cheek teeth less than ^.
Upper premolars 1-1.
Rostrum noticeably concave on each side of middle
line ; nares extending about halfway to inter-
orbital constriction ; palatal emargination

broader than deep 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 Piinstrellus, 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 [viyotis).

1829. Nystactcs Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,

p. 108 {bechsteinii).

1830. Leuconoe Boie, Isis, p. 256 (daubentonii).

1839. Yespertilio Keyserling and Blasius, Wiegmann's Archiv fiir Natur-
gesch., 1839, i, p. 306 (Not Vespertilio Linnaeus, 1758).

1841. Sclysius 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 {Jiasseltii, macellus = advcrsus and blepotis). •

MYOTIS 167

1849. Tralatitus Gervais, Diet. Univ. d'Hist. Nat., xiii, p. 213 (Modifica-
tion of Trilatitus).

1856. Bruchyotus Koleuati, Allgem. deutsch. Naturhist. Zeitung, Dresden,
neue Polge, ii, p. 131 {mystacinus, daiihentonii, and dasycneme).
Not Brachyotus Gould, 1837.

1856. Isotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, neue

Folge, II, p. 131 [nattereri and emarginatus).

1857. Vc&pertilio Blasius, Saugethiere Deutschlands, p. 78. Not Vespertilio

Linnseus, 1758.

1866. Tralatitius Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 90,

February 1866 (Modification of Trilatitus).

1867. Pternopterus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin,

p. 706 (sub-genus of Vespertilio = Myotis, type lobipes = muricola).

1870. Exochurus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,
Math.-Naturwiss. Classe, lxii, p. 75 (macrodactylus, horsfieldii =
adversus and ruacrotarsus).

1870. Aeorestes Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wiea,
Math.-Naturwiss. Classe, lxii, p. 427 [villosissimus, albescens, and
nigricans).

1870. Comastes Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.-
Naturwiss. Classe, lxii, p. 565 {capaccinii, megapodius, dasycneme,
and limnophilus).

1878. Vespertilio Dobson, Catal. Chiropt. Brit. Mus., p. 284. Not Vespertilio
Linnffius, 1758.

1897. Mijotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 382,
October, 1897.

1899. Euvespertilio Acloque, Fauue de France, Mammiferes, p. 38 (emargin-
atus, mystacinus, ynuriniis = myotis, nattereri, and beclisteinii) .

1907. Myotis Miller, Families and Genera of Bats, p. 201, June 29, 1907.

«

Type sjjecies. — Vespertilio myotis Borkhausen.

Geographical distribution. — Entire mainland of Eastern and
AVestern hemisphere.s to limits of tree growth ; also the Malay
Archipelago, New Guinea, Australia and Samoa, and in America
the Lesser Antilles.

Characters. — Dental formula : i ||, c Jfi, pm ||) ^ H = 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 CHIROPTEKA

KEY TO THE EUROPEAN SPECIES OF MYOTIS.

Size large (forearm 53 to 64 mm., condylobasal
length of .JiuU 18-6 to 23 "(5 mm., upjjer tooth-
row 8'2 to 10'6 mm.) ; middle upper premolar
normally crowded inward frour axis of tooth-
row ; third lower molar with second triangle
much smaller than first and noticeably different
from it in form.
Condylobasal length of skull 22 to 23*6 mm.;
mandible 17 "8 to 19 mm. ; maxillary tooth-
row 9"8 to 10 "6 mm. (Central and southern

Europe) M. mijotis, p. 192.

Condylobasal length of skull 18-6 to 21-4 mm. ;
mandible 15 ■ 2 to 17 • 2 mm. ; maxillary tooth-
row 8'2 to 9' 4 mm. (Mediterranean region) M. ox-ygnathus, p. 199.
Size smaller medium (forearm .34 to 47 mm., condy-
lobasal length of skull 12-4 to 16 "8 mm., upper
tooth-row 5 to 7 mm.) ; middle upper premolar
not crowded inward from axis of tooth-row ;
third lower molar with second triangle nearly
as large as first and essentially like it in
form.
Foot relatively large, obviously more than half
as long as tibia ; calcar about twice as long
as free border of interfemoral membrane ;
skull broad, the width of brain-case more
than half greatest length ; crown area of
molars relatively small ; upper molars with
evident protoconule.
Forearm about 47 mm. ; condylobasal length
of skull about 16 mm.. (Central and

southern Europe) M. dasi/ciicme, p. 189.

Forearm less than 45 mm. ; condylobasal length
of skull never more than 15 mm.
Tibia and adjacent membrane densely furred ;
forearm about 42 mm. ; condylobasal
length of skull 14-0 to 14-8 mm. (Medi-
terranean region) M. ccqxiccinii, p. 187.

Tibia and adjacent membrane not furred ;
forearm about 35 mm. ; condylobasal
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. ; condylobasal 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. ; condylobasal length
of skull 12-6 to 13-2 mm.; posterior
border of ear conch with shallow, in- ,

conspicuous emargination M. mystacinus, -g. IGO. sj

Ear elongated, extending conspicuously beyond
nostril when laid forward.

ilYOTIS

169

Combined length of tibia and foot less than

25 mm. ; condylobasal length of skull

14 • 0 to 14 • 6 mm. ; ear narrow, its width

about 10 mm. ; tragus conspicuously

more than half as high as conch M. nattercri, p. 174.

Combined length of tibia and foot about

30 mm. ; condylobasal length of skull

16 to 17 mm. ; ear broad, its width about

14 mm. ; tragus scarcely half as high as

conch -1/. Ijcclistcinii, p. 17'J.

MYOTIS MYSTACINUS Kuhl.

1S19. Vespcrtilio mystacinus Kuhl, Ann. Wotterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 2, p. 202.
1821. yespcrtilio coUaris Schinz, Das Thierreich von Cuvier, i, p. 177

(Mt. Blanc, Haute-Savoie, France).
1834. Vespcrtilio humeralis Baillon, Mem. Soc. Eoyale d'Emulation

d' Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1837. Vespcrtilio schinzii Brehm, Ornis, Heft iii, p. 27 (Renthendorf,

Thiiringen, Germany).
1843. V[espcrtilio'\ scUrankii Wagner, Wiegmann's Archiv fiir Naturgesch.,

IX, Bd. II, p. 25 (Munich, Germany? See Fitzinger, Sitzungsber.

kais, Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 219, 1871).
1857. Vespcrtilio mystacinus Blasius, Saugethiere Deutschlands, p. 9G.
1863. [Brachyottcs ■mystacinus'] var. nigricans Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden,

Nassau, Germany).
1863. [Brachyotus mystacinus] var. rnfofnscus Koch, Jahrb. des Vereins

fiir Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden.

Nassau, Germany).
1863. [BracJiyotus mystacinus] var. aureus Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 445 (Breisgau,

Cxermany).
1869. [Vesjxrtilio mystacinus] var. nigricans Fatio, Faune Vert. Suisse,

I, p. 92 (Switzerland). Not of Koch, 1863.
1869. [Vespcrtilio] lugubris Fatio, Faune Vert. Suisse, i, p. 93 (Alternative

for nigricans Fatio).
1871. Vespertilio mystacinus, nigro-fuscus Fitzinger, Sitzungsber. kais.

Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 217 (Renaming of V. schinzii Brehm).
1878. Vespertilio ntysfacinus Dobson, Catal. Chiropt. Brit. Mus., p. 314.
1900. Myotis mystacinus M6hely, IMonogr. Chiropt. Hungarise, p. 200.
1910. Myotis mystacinus Trouessart, Faune Mamm. d'Europe, p. 38.

Gcograplilcal disirihution. — Entire Continent of Europe north
to about the hiuits of tree growth ; west to Ireland ; east into
Asia.

Diagnosis. — Smallest species of European Myotis (forearm
about 34 mm., longest finger about 60 mm., condylobasal 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 uropatagium 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
prodvTcing a general darkening effect ; underparts paler and more
buffy, usually not forming any decided contrast with back, but
occasionally almost whitish, especially on
chest ; muzzle and cheeks dusky ; ears and
membrane blackish.

SJmll. — 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-

Mi/oti^mystacums. ^^^^^ 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 gi^eat. Lower incisors
slightly but evidently imbricated,
forming a continuous, broadly
V-shaped row between
canines ; crown of i^ sub-
terete, its area more than
half that of canine and
about equal to that of pm^,
its blunt main cusp, the
highest in the incisor series,
situated at outer side, the
three smaller cusps (third
obsolete) on inner margin ;
ij and in, sub-equal, slightly
lower than i^, 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

Fig. 30.
Myotis mystacimts. Teeth x 10.

172 CHIKOPTKRA

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 cusjjs ; 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 slightl}^ greater than that of first ; anterior
and posterior borders slightly concave, inner border narrowly
rounded, especially in m^, the antero-internal and postero-internal
outlines often flattened or even a little concave ; protocone large,
its base occupying entire inner border of tooth, its cusp a little
in front of middle, its anterior commissure simple, extending
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 ujjper molar with crown area
equal to about three-quarters that of m^, the protocone as in the
other teeth but smaller, paracone larger than metacone, the first
outer commissure longer than in m'^ and nr, 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^ ; protoconid decidedly higher than
Iwpoconid in all three teeth ; inner cusps of about equal height
thx'oughout (slightly more than half as high as protoconid) ; a
distinct cingulum cus^j behind entoconid.

Measurements. — Adult female from Skane, Sweden : tibia,
15 ; foot, 7*6; forearm, 32 ; thumb, 6 • 2 ; third finger, 49 ; fifth
finger, 38 ; ear from meatus, 1 2 ; width of ear, 8. Adult male from
Madrano, Tyrol: head and body, 38; tail, 38; tibia, 16; foot,
8 ; forearm, 35 ; thumb, 6 • 2 ; third finger, 56 ; fifth finger, 47 ;
ear from meatus, 13; width of ear, 9. Adult female from the
Carpathian Mountains : head and body, 44 ; tail, 40 ; tibia, 15 ■ 4 ;
foot, 8 ; forearm, 34 ; thumb, 7 ; third finger, 54 ; fifth finger,

MYOTIS 173

■45 ; ear from meatus, l-4'2 ; 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,

Speciiiiois examined. — Thirty-five, from the following localities: —

England: Newby Bridge, Laice Windermere, Cumberland, 1; iVberia,
Merionethshire, 2 ; Colwyn, Denbighshire, 1 ; Cheadle, Staffordshire, 1 ;
Manchester, Lancashire, 1; Ragley House, 1; Welford, Herefordshire, 2 ;
^lacclesfield, 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, Li^ge, 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.); Csallokoz-Somorja, Pressburg, 2; Tatra Mts., Hungary, 1.

Switzerland : Geneva, 1 (Mottaz) ; Grotte de Vallorbe, Vaud, 2
(Mottaz); Stein, Appenzell, 1 (U.S.N.M.); St. Gallen, 1 (U.S.N.M.); no
exact locality, 1 (Geneva: type of luguhris Fatio).

liemarhs. — Myotix 'mijsiacinun, 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 9. Aberia, ^Merionethshire,

Wales.
9 st. Colwyn, Denbighshire.

1 al. Newby Bridge, Cumber-

land, England.

?. Cheadle, Staffordshire.

6. Manchester, Lancashire.

6. Ragley House, Warwick-

shire.

2 9. Welford, Herefordshire.
6. Pewsev, Wiltshire.

(P. S.'Hembly.)
9. Dover, Kent. {B. Hesse.)

1. Hastings, Sussex.

6 st. Bath, Somerset.
1. Skullno, Sweden.

9. Haida, Bohemia. (U"u/-

terstorff.)
1. Csallokoz- Somorja,

Pressburg, Hungary.
1 al. Tatra j\Ioun tains,

HunRarv.

Ct. H. Caton Haigh
(c & p).

Hon. N. C. Roths-
child (p).

T. Paul (c & p).

E. Blagg (c & r).
C. Oldham (p).
Tomes Collection.

Tomes Collection.
C. H. B. errant (p).

C. H. B. Grant (p).
MissI.Eoods (c & p).
G. Dalgleish (c & p).
Stockholm Museum

(E.)

Lord Lilford (p).

Budapest Museum

(e).
Dr. li. Collett (p).

IL 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. 8. 12.
49. 1. 16. 1.
4. 10. 13. 3.
46. 1. 2. 22.

11. 1. 1. G.

94. 3. 1. 20-21.

91. 1. 21. 1.

174 CHIROPTERA

MYOTIS NATTERERI Kulll.

1818. Vespertilio nattcreri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 33.
1857. Vespertilio nattcreri Blasius, Saugethiere Deutschlands, p. 88.
1863. [Isotiis nattcreri'] var. ti/jnis Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 430 (Wiesbaden, Nassau,

Germany).
1863. [Isotus nattcreri'] var. spelieus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 430 (Erdbach, Nassau,

Germany).
1878. Vespertilio nattcreri Dobson, Catal. Chiropt. Brit. Mus., p. 88.
1900. Myotis nattcreri Mehely, Monogr. Chiropt. Hungarife, p. 179.
1904. Myotis cscalerai Cabrera, Mem. Sec. Espan. Hist. Nat., ii, p. 279

(Bellver, L6rida, Spain).
1910. Myotis nattcreri and AI. cscalerai Trouessart, Fauna Tilamm. d'Europe,

l^p. 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, 14-0 to- 14-6 mm.); ear elongated, extending about
5 mm. beyond tip of muzzle when laid forward, the conch narrow
(about 10 mm.), obscurely emarginate on upper half of posterior
border, the tragus relatively longer than in any other European
species, its height distinctly more than half that of conch ; foot
about half as long as tibia ; wing membrane extending to base of
outer toe ; edge of interfemoral membrane fringed.

External characters. — General form essentially as in Myotis
mystacinus, but difiering in the following particulars : ear much
longer and relatively narrower, extending conspicuously beyond
nostril when laid forward, its extremity more broadly rounded
off (owing to more uniform convexity of anterior border), and
concavity on posterior border even less evident ; antitragus
about as large as in 31. mystacinus but less well defined ; tragus
relatively longer and more attenuate than in any other European
bat, its height conspicuously more than half that of conch, its
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. inystacinus, 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 cscalerai
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
1 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 dvisky,
but not so dark as in 31. 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 difier, 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 ci"owns,
the cingulum of the third forming a very low, sometimes obsolete
antero-internal cusp ; molars, especially m^, with narrower
crowns.

Measurements. — External measurements of adult male from
Colpin, Brandenburg, Germany, and adult male fi'om Magdeburg,
Germany: head and body, 50, 44; tail, 41, 40; tibia, 17, 16;
foot, 8•L^ 9-0; forearm, 39-2, 40-2; thumb, 7-0, 6-2; third
finger, 71,71 ; fifth finger, 56, 56 ; ear from meatus, 18, 18 ; tragus,
11 ■ 0, 11*2. Two adult females from Spain (No. 94.1.1.8, Seville,
and No. 8. 7. 23. 4 from Bellver, Lerida, paratype oiM. escalerai) :
head and body, 46, 46 ; tail, 41, 43 ; tibia, 15, 17 ; foot, 7 '6, 9*2 ;

17G

CHIROPTEEA

forearm, oG-4, 40-6 ; thumb, 7-2, 8-4 ; third finger, 69, 72 ;
fifth finger, 52, 56; 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.) ; Bradtleld, 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 ; ^lagdeburg,
Saxony, 2; Moritzburg, Saxony, 3 (U.S.N.M.).

Austria-Hungary: Haida, Arva, Bohemia, 1.

Switzerland: Canton Thurgau, 3 (U.S.N.M.): St. Gallen, 9 (B.JM.
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.IM. and Genoa ; paratypes of escalerai
Cabrera) ; Seville, 1.

Bemarks. — This species is easily recognized b}" 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.

1 al.

Inverary, Argyllshire,

Scotland.
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. Germanv.
(ir. Baer.)

6 al. Colpin, Brandenburg,
Prussia.

6 al. ^Magdeburg, Saxony.

Magdeburg, Saxony.
(WoUerstorff.)

Haida, Bohemia. ( Wol-
tcrstoif.)

St. Gallen, GOO m. Swit-
zerland. (E. H. Zolli-
kofer.)

Duke of Argyll (p).

Dr. G. E. Dob.son

(c & p).
R. F. Hibbert (p).
.J. Backhouse (p).

Tomes Collection.

Tomes Collection.
Lord Lilford (c & p).

N. H. Joy (c & p).
J. G. Millais (c & p).
Heatley Noble (c & p).
R. H. Read (p).
Dr. J. E. Gray (p).
Lord Lilford (p).

58. 8. 16. 1.

76. 11. 3. 2.

11. 1. 8. 20.
11. 1. 3. 16-18.

7.1.1.488-493.

7.1.1.736-743.
72. 8. 21. 3-5.
72. 11. 12. 13.
11. 1. 3. 19.

I. IL 2. 1.
0. 8. 28. 1.

II. 1. 3. 21.
5L 1. 29. 12.
99. 1. 9. 6.

Dr. H. Gadow (c & p). 82. 7. 31. 2.

3 6, 9.

Dr. \N. Wolterstorff
Lord Lilford (p).
Lord Lilford (p).
0. Thomas (p).

92. 12. 1. 1.

11. 1. 1. 8-

11. 1. 1.

4. 4. 5. 4-8.

MYOTIS 177

2 al. Arezzo, Italy. Florence Museum (e). 85.7.6.4-5.

? 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.

(c & p).

MYOTIS EMARGINATUS Geoffroy.

1806. Vcs2o[crtilio'] cmarginatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris,
Yiii, p. 198 (Charlemont, Givet, Ardennes, France).

1844. Vesplertilio] rufescens Crespou, Faune IMeridionale, i, p. 20 (near
Nimes, Gard, France). Type in Nimes Museum.

1853. V[esp{;rtilio] ciliatus Blasius, Wiegmann's Archiv fiir Naturgesch.,
1853, I, p. 287 (near Cologne, Germany).

1856. Vles2)crtilio] schranJdi Kolenati, AUgem. deutscbe Naturhist. Zeitung,
Dresden, ueue Folge, ir, p. 178. A nomen nudum (ex Kocb and
Giebel) cited as synonym of emarginatus. Not of Wagner, 1843.

1858. Vespertilio ciliatus Blasius, Saugetbiere Deutscblauds, p. 91.

1878. Vespertilio cmarginatus Dobson, Catal. Cbiropt. Brit. Mus., p. 303.

1880. Myotis ciliata var. budapestiensis Margo, " IMagyar orv. es termes-
zetvizsg. xx,nagygyui. munk, p. 255 " (Budapest, Austria-Hungary).
See M^bely, Monogr. Cbiropt. Hungarise, p. 170.

1890. Vespertilio neglectus Fatio, Arcb. Sci. Pbys. et Nat., Geneve, 3rd ser.,
XXIV, p. 512, November 15, 1890 (Valavran, near Gieneva, Swit-
zerland). Type in Geneva Museum.

1900. Myotis emarginatus M6bely, Monogr. Cbiropt. Hungarise, p. 170.

1910. Myotis cmarginattis Trouessart, Faune Mamm. d'Europe, p. 28.

Type locality. — Charlemont, Givet, Ardennes, France.

Geographical distribution. — Central and southern Continental
Europe.

Diagnosis. — Size essentially as in 3fyotis 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 boi-der with a deep almost
angular emargination slightly above middle ; foot about half as
long as tibia ; wing membrane extending to base of outer toe ; free
margin of interfemoral membrane sometimes fringed ; fur some-
what woolly in texture, the hairs of back tricolored, drab at
base, yellowish at middle and dark brown at extreme tij:).

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,
Vjoth its margins essentially straight from level of anterior base

N

178

CHIROl'TERA

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 31.
mystacinus, extending to between ears when laid forward, only
the minute cartilaginous tip free from membrane.

Fiir and colour. — Fur shorter and more dense than in M.
mystacinus and M. nattereri, and of a slightly woolly texture
unique among the Euroj^ean 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 similar!}^ low as compared with that
of 31. mystacinus. Mandible slightly more robust than in 31.
nattereri, but of similar form.

Teeth.- — The teeth are larger than those of 3Iyotis 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^ and i.,
alike in form and distinctly 4-cusped, that of ?\ not thickened
posteriorly and with no trace of postero-internal cusp. Canines
about equal to those of 31. nattereri in size, but shaft of upper
tooth with evident postero-external longitudinal groove. First
and second upper premolars even more strongly contrasted in
size than those of 31. mystacinus, the cusp of second only a little
exceeding cingulum of first ; the two teeth less crowded between
canine and large premolar than in 31. mystacinus and 31. nattereri ;
lower premolars essentially as in 31. nattereri, but less closely
crowded, and second relatively larger. Upper molars as in
31. nattereri, but crowns less narrowed ; a rudimentaiy commissure
extending outward from base of hypocone and another extending
inward from base of paracone • lower molars not peculiar.

Measurements. — 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 8' 6; forearm, 40 and 41 ; thumb, 8 and 7 '8
third finger, 67 and 70 ; fifth finger, 56 and 57 ; ear from meatus
16*6 and 17 ; width of ear, 11 "4 and 12. For cranial measure
ments see Table, p. 183.

Specimens examined. — Six, from the following localities : —
• Holland : Maastricht, 1 (U.S.N.M.).

MYOTIS 179

Prance: Near Nirnes, Gard, 1 (Nimes : type of rufescens Crespon).
Austria-Hungary : Herkulesbad, 2.

Switzerland : Valavrau, near Geneva, 1 (Geneva : type of neqlectus
Fatio).

Italy : Florence, 2 (U.S.N.M.) ; no exact locality, 1.

Remarks. — The peculiar form of the ear, the short, somewhat
woolly fur, and the yellowish colour are highly characteristic of
this well-defined species.

•2 ?. Herkulesbad, Hungary. Hon. N. C. Eoths- 7. 9. 16. 9-10

child (p).
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 733.

without parte.)

skull.

MYOTIS BECHSTEINII Kuhl.

1818. Vesjyertilio bcchstelnii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 1, p. 30 (Hanau, Hessen, Germany).

1857. Vesjpertilio beclisteinii Blasius, Saugethiere Deutschlands, p. 85.

1878. Vespertilio bechsteinii Dobson, Catal. Chiropt. Brit. Mus., p. .308.

1900. Myotis bechsteinii M6hely, Monogr. Chiropt. Hungarise, p. 184.

1905. Yesp\_ertilio\ bechst[einii] ghidinii vel Ves2J[ertilio'] ghidinii Fatio,
Arch. Sci. Phys. et Nat., Geneve, 4th ser., xix, p. 511, May 15,
1905 (Lugano, Ticino, Switzerland). Tj^pe in Geneva Museum.

190G. Myotis bechsteinii favonicus Thomas, Ann. and Mag. Nat. Hist.,
7th ser., xviii, p. 220, September, 1906 (La Granja, Segovia,
Spain). Type in British iluseum.

1910. 2Iyotis bechsteini and M. bechsteini favonicus Trouessart, Faune
Mamm. d'Europe, pp. 80-31.

Type localitii. — 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
forw^xrd, 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 auritns ; form
of ear essentially as in M. mystacinus, the upper half of posterior
border faintly concave ; inner surface of conch wath about eight
rather ill-detined transverse ridges near posterior border ; anti-
tragus about 2 • 5 mm. in length along ba.se, rather abruptly

N 2

1(^0 CIIIROPTEEA

rounded ofl' above, its inner margin not continuous with posterior
border of conch. Tragus about half as high as conch, its form
essentially as in 31. mystacinu,s though a little more slender
(greatest width contained nearly three times in height of
anterior border) ; basal lobe small bvit well developed. Wing
essentially as in 31. mystaciims ; 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 31. mystadnns. 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 31yoHi<
mystacinus and M. natter eri. Upper parts uniform wood-brown
Avith a slight tinge of umber, the exact shade almost impossible
to describe exactly ; vmderparts a buffy grey in strong contrast,,
irregularly clouded by the slaty brown of underfur ; ears and
membranes an indefinite dai'k brovrn.

ShuJl. — In general aspect the skull resembles that of 3Iyotii<
myotis, due allowance being made for its much smaller size, since
it is much more slender than in any of the other small species ;
breadth of brain-case equal to about one-half distance from
lambda to posterior border of narial emargination. Dorsal
profile rising gradually above rostrum and forming a strong
convexity over anterior portion of brain-case, behind w^iich it is
essentially fiat 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 obsciu-ely 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 oS" 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 3lyotis myotis, the coronoid j^rocess higher and
with more oblique posterior boi-der than in the other small species.

Teeth. — The teeth are rather small relatively to size of skull..

MYOTIS 181

Upper incisors essentially as in 31. mystacinus but more slender.
Lower incisors slightly imbricated, the row as a whole U-shaped
or A^ery broadly V -shaped, the form of the individual teeth not
l>eculiar. 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. natter eri. Upper molars with no
special peculiarities except that m^ is more reduced than in the
other small species, its tranverse diameter through metacone
noticeably less than half length of anterior border. Lower
molars normal, but second triangle of m^ a little more reduced
than in the other small species, though much less so than in
31. myotis and M. oxygnatlius.

Measurements. — Adult female from Sweden : head and body,
46; tail, 38; tibia, 19-6; foot, 9-8; forearm, 42 ; thumb, 9-6;
third finger, 67 ; fifth finger, 56 ; ear from meatus, 22 ; ear
from crown, 19; width of ear, 13-6; tragus, 10. Two adults
from the New Forest, Hampshire, England : tibia, 18*8 and 20 ;
foot, 9 • 8 and 10 • 6 ; forearm, 39 and 42 • 4 ; thumb, 8 • 4 and 10 ;
third finger, 64 and 68 ; fifth finger, 53 and 57. Adult male
from Lugano, Switzerland (Geneva : type of gJudinii Fatio) :
tibia, 19-4; foot, 10; forearm, 41*2; thumb, 10-4; third
finger, 65; fifth finger, §3 ; 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
measui'ements see Table, p. 183.

Specimens examined. — Seventeen, from the following localities : —
England : Henley-on-Thames, Oxfordshire, 1 ; New Forest, Hamp-
shire, 6.

Sweden: Sktlne, 1 (U.S.N.M.).

182

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184

CHIROPTERA

Feance : Etupes, Doubs, 1 (Mottaz) ; Ste. Baume, Var, 1 (Genoa) ;
Cadillac, Gironde, 1 (Lataste).

Switzerland: Lugano, Ticino, 1 (Geneva: type of ghidinii Fatio).
Spain : La Grauja, Segovia, 1 (type oi favoniciis Thomas).
Austria-Hungary: Zay-Ugrocz, Treucsen, 4 (B.M. and U.S.N.M.).

Remarhs. — 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.

i. 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.
Sal. La Granja, Segovia, M. de la Escalera (c). 6.11.4.1.

Spain. (Type of M. favonictis Thos.)

6, 9 al. Zay-Ugrocz, Trencsen, Budapest IMuseum 6. 6. 20. 1-2.
Hungary. (e).

1. Europe. " Leyden Museum. 37. 4. 28. 23.

MYOTIS DAUBENTONII Kuhl. "

1819. Vcspertilio daubentonii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 2, p. 195 (Hanau, Hessen, Germany).
1839. Ves2xrtilio mdilis Jenyns, Ann. Nat. Hist., iii, p. 73, April, 1839

(Aukland St. Andrew, Durham, England).
1844. V\cspertilid] lanatiis Grespon, Faune M^ridionale, i, p. 15 (South of

Nimes, Gard, France). Type in Nimes I\Iuseum.
1857. Vespertilio daubentonii Blasius, Saugethiere Deutschlands, p. 98.
1871. Vespej-tilio capucinellus • ' Koch, Bayr. Fauna," Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Glasse, LXii, pt. 1,

p. 206 (Bavaria?).
1871. Vespertilio minutelliis " Koch, Bayr. Fauna," Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 206 (Bavaria?).
1871. Vespertilio dmibentonii, albus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, p. 210

(Renaming of V. mdilis Jenyns).
1878. Vespertilio daubentonii Dobson, Catal. Chiropt. Brit. Mus., p. 297.
1890. Vespertilio staufferi Fatio, Faune Vert. Suisse, v, 3""= suppl. aux

Mamm., p. 6 (Lucerne, Switzerland). IMS. synonym; repudiated.
1900. Myotis daubentonii Mehely, i\Ionogr. Chiropt. Hungarias, p. 164.
1910. Jlyotis daubentoni Trouessart, Faune Mamm. d'Europe, p. 27.

Tjipe locality. — Hanau, Hessen-Nassau, Germany.

GeograjjJiical distribution. — Europe from the Mediteri-anean
north to central Norway, west to Ireland, east into Asia.

Diagnosis. — Size nearly as small as in Myotis mystacinus
(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 m.m. 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.
iiiystacinKK, 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 conca\T.ty above ; antitragus small and ill-defined ; tragus
about half as high as conch, its greatest width contained three
times in length of anterior border, the posterior border
moderately and evenly convex, the anterior border nearly
straight, the tip rather blunt ; posterior basal lobe relatively
large and well defined. Wing slightly broader than in 31. rnysta-
cinus, the three main metacarpals evidently graduated from
third to fifth, the third very slightly shorter than forearm ;
membr«,ne 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. naftereri,
though usually inclining more definitely toward wood-brown ;
underparts butly grey usually less contrasted than in M. nattereri,
though sometimes pale enough to produce a distinct line of
demarcation along sides of neck. Muzzle and cheeks dusky.
Membranes and ears an indefinite brown.

Skull. — The skull of Myotis clauhentonii 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 rnysta-
^■inus, a difference particularly noticeable in the crown areas of
the first and second upper molars. Upper incisors as in mystacinus,
Ijut with cingulum less developed. Lower incisors very slightly
imbricated, the form of the row vacillating between U-shaped
and broadly V- shaped, the cusps as in M. mystacinus but less

186

CHIROPTKRA

FIG. 31.
Ml/of is dauhentonii. Teeth x 10.

developed. Canines both above and below weaker and less
trenchant than in M. mystacinus, their form essentially as in

M. nattereri. Premolars
and lower molars not
obviously different from
those of 31. mystadnns.
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 secondar}-
commissure extending to
base of jDaracone ; m^ and
VI- 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.

Specimetifi 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, ; Eppiug, Essex, 2 ;
Northamptonshire, 1 ; Henley-on-Thames, Oxfordshire, 2 (B.M. and
U.S.N.M.); Christchurch, Hampshire, 1 ; Devonshire,!.

Sweden: Upsala, 3 (B.M. and U.S. N.M.); upland,!; no exact locality, 2.

SwiTZEBLAND : Goncva, 1 (Mottaz).

Prance : Near Nimes, Gard, ! (Nlmes : tvpe of lanatus Crespon).

Italy : Lecco, Lombardy, 2! (U.S.N.M.) ; Pavia, 1 (U.S.N.M.);
Florence, 30 (Mottaz).

BemarJcs. — 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). !!. !. 3. 60.

land.

MYOTIS

im

?.

Knutsford, Cheshire.

T. A. Coward (c&p).

11. 1. 3. 22.

2.

Stratford-on-Avon, War-
wickshire.

Tomes Collection.

7.1.1.486-487.

(5al.

Hillingdon, Middlesex.

0. Thomas (c & p).

84. 1. 29. 1.

2al.

Epping, Essex.

H. Doubleday (c&p).

44. 10. 21. 1-2.

al.

Northampton.

Mrs. Jenyns (p).

9 St.

Henley, Oxfordshire.

J. G. Millais (c & p).

1. 11. 2. 2.

1 al.

Christchurch, Hamp-
shire.

Lord Lilford (f).

S7. 9. 1. 1.

6.

Upland, Sweden.

{G. Kolthoff.)

Lord Lilford (p).

11. 1. 1. 25.

2.

Sweden.

Stockholm Museum

46. 6. 2. 15.

(E).

48. 6. 28. 3.

MYOTIS CAPACCINII Bonaparte.

1837. Vcspertilio capaccinii Bonaparte, Iconogr. Faun. Ital., i, fasc. sx

(Sicily). Type in British Museum.
1839. Vcspertilio megapodius Temminck, IMonogr. de Mamm., ii, p. 189

(Sardinia).
1841. Vcsp[ertilio'] clasypus de Selys-Longchamps, Atti della seconda

Riunione degli Scienziati Italiani, Torino, 1840, p. 249 (Published

as synonym of capaccinii).
1844. Ycsp^crtilio] pellucens Crespon, Faune JMeridionale, i, p. 16 (Cave

near Pont-du-Gard, Gard, France).
1857. Vesijertilio capaccinii Blasius, Siiugethiere Deutschlands, p. 101.

1877. Vcspertilio hlasii Major, Atti Soc. Tosc. Sci. Nat., Pisa, iii, p. 108

(New name for the capaccinii of Blasius should it prove to be
different from that of Bonaparte).

1878. Vcspertilio majori Ninni, Atti Reale Instit. Veneto, 5th ser., iv,

pt. 1, p. 721 (Substitute for blasii Major).
1878. Vespertilio capaccinii Dobson, Catal. Chiropt. Brit. IMus., p. 293.
1901. Myotis capaccinii Thomas, Proc. Zool. Soc. London, p. 37.
1910. Myotis {Leuconoe) capaccinii Trouessart, Faune Mamm. d'Europe, p. 26.

Type locality. — Sicily.

Geographical distribution. — Mediterranean region, north to
Italian Switzerland, east into Asia.

Diagnosis. — Not so small as Myotis daubentonli (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. dauhentonii. The foot, however, is relatively
larger, and the wing membrane is inserted at the ankle.

Fur and colour. — Fur rather dense and short, the hairs at
middle of back about 6 mm. in length ; distribution peculiar
among the European members of the genus in its tendency to
spread on membranes, forming a 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 CHIROPTERA

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 butty 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.

Shull. — In all respects the skull so closely resembles that of
Myotls dauhenfnnii that it is only distinguishable by its larger
size. From the skulls of M. nattereri and M. emarginatus it
differs in its greater breadth, a character perhaps most readily
appreciable in the form of the post-palatal region.

Teeth. — Except for their larger size the teeth resemble those
of Myotis dauhentonii. The crown area of upper molars is
i-elatively less than in M. nattereri and M. emarginatus, though
m^ retains the broader outline characteristic of M. dauhentonii
and M. mystacinus. Upper molars with protoconule and its
accessory small commissure, and m^ and m~ 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 Corleone, Sicily :
head and body, 49 ; tail, 38 ; tibia, 16 ; foot, 10 ; forearm, 41 ;
thumb, 8-2 ; third finger, 68 ; fifth finger, 55 : ear from meatus,
15; width of ear, 10*4. Two adult males from Sassari, Sardinia :
head and body, 50 abd 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) ; Pavia, 1 (U.S.M.M.) ; Ostia,
Rome, 2 ; Marsala, Sicily, 1 ; Corleone, Sicily, 1 (U.S.N.M.) ; Sicily, no
exact locality, 1 (type).

Sardinia: Cagliari, 4 (B.M. and U.S.N.M.); Sassari, 5 (U.S.N.M.);
Grotte de Sardale, 2.

France : Marseilles, 1 (U.S.N.M.).

Spain : Inca, Majorca, Balearic Islands, 1 ; Elche, Alicante, 1.

Bemarlis. — In general appearance this .species resembles
Myotis dauhentonii; but it is immediately recognizable by its
even larger foot, and by the densely pubescent upper surface of
leg, the latter character unique among European bats.

4 6, 9. Near Lugano, Ticino, 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 Italj-. (Prince Bona- Tomes Collection. 7. 1. 1. 734.

without parte.) {Tyxx' of species.)

skull.

?. Marsala, Sicily. 0. Tliomas (p). 6. 8. 4. 21.

(.4. Robert.)

2 al. Cagliari, Sardinia. Florence Museum (e). 85. 7. 6. 6-7.

2 al. Grotte de Sardale, Sar- O. Thomas (p). 0. 12. 3. 1-2.

dinia.

c?. Inca, jMajorca, Balearic O. Thomas and R. I. 0. 7. 1. 3.

Islands. Pocock (c & p).

6. Elche, Alicante, Spain. G. S. Miller (c). S. 8. 4. 13.

MYOTIS DASYCNEME Boie.

1823. Vcspcrtilio viijstacimis Boie, Isis, p. 965. Not V. mystacinus Kuhl

(Jutland, Denmark).
1825. Vespe7-tilio dasycncme Boie, Isis, p. 1200 (Renaming of mystacinus

Boie).
1889. Vespcrtilio limnopliilus Temminck, Monogr. de ]\Iamm., ii, p. 176

(Holland).
1857. Vespcrtilio dasycneme Blasius, Siiugethiere Deutschlauds, p. 103.
1878. Vespertilio dasycneme Dobson, Catal. Chiropt. Brit. Mus., p. 295.
1904. Myotis dasycneme Trouessart, Catal. ]\Iamm. Tam viv. quam foss.,

suppl., p. 88.
1910. Myotis dasycneme Trouessart, Faune INIamm. d'Europe, p. 27.

Type locality. — Chalk quarries at Dagbierg, near Wiborg,
Jutland, Denmark.

Geographical distrihition. — Central and southern Europe,
west to the Atlantic coast,* north to Sweden, east into Asia.

Diagnosis. — Form and proportions essentially as in Myotis
dauhentonii 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 dauhentonii ;
quality not peculiar, the longest hairs on middle of back about
8 mm. in length. Colour of upper parts a light yellowish wood-
brown ; underparts strongly contrasted greyish white with a
tinge of bufi', 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

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192

CHIROPTERA

Skull. — The skull is large, slightly exceeding that of Myotis
hechsteinii 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 Myotic dauhentonii, but the rostrum is relatively
shorter and the brain-case broader and more depressed. Sagittal
crest barely indicated ; auditory bulke 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 Myotic-
dauhentonii ; molars decidedly more robust than those of M.
hechsteinii, the width of the crown diminishing much less rapidly
toward inner border ; small premolars much ci'owded, the second
barely or not visible from outer side, the diameter of its crown,
however, not much less than that of anterior tooth ; upper
molars as in M. dauhentonii and M. capaccinii ; mandibular teeth,
with no special peculiarities, the premolars and outer cusps of
molars more slender than in M. hechsteinii and M. dauhentonii.

Measurements. — Two adult males from Maastricht, Holland :
head and body, 57 and 58 ; tail, 49 and 51 ; tibia, 19 "8 and
20-2 ; foot, 11-8 and 12 ; forearm, 43 6 and 44 ; thumb, 9-6
and 9 ; third finger, 72 and 77 ; iifth 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 exammcd. — Six, from the following localities : —
Holland : Leyden, 1 ; Maastricht, 4 (U.S.N.M.).
Belgium : Near Naniiir, 1.

1. Leyden, Holland. Tomes Collection. 7. 1. 1. 501.

{H. Schlegel.)
6 al. Namur, Belgium. Rev. D. B. Lebbe 9. 1. 11. 1.

(c & p).

MYOTIS MYOTIS Borkhausen.

1775. Vespertilio murinus Schreber, Stiugthiere, i, p. 165 (Not of Linnaeus,

1758).
1797. Vesyertilio myotis Borkhausen, Deutsche Fauna, i, p. 80 (Germany).
1797. Vespertilio myosotis B[orkhause]n, Der Zoologe (Compendiose

Bibliothek gemeinniitzigsten Kenntnisse f iir alle Stiinde, pt. sxi),

Heft v-viii, 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 iii, p. 24 (Renthendorf,

Thiiringen, Germany).
1844. V{espertilio\ latipinnis Crespon, Fauna M^ridionale, i, p. 17 (Near

Nimes, Gard, France).
1857. Vespertilio murinus Blasius, Saugethiere Deutschlands, p. 82 (Not of

Linnseus, 1758).
1863. [Myotus murinus'] var. typus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 415 (Wiesbaden, Nassau,

Germany).
1863. [Myotus murinus'] var. aljnnus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 415 (St. Gothard, Uri,

Switzerland).
1878. Vespertilio nmrimis Dobson, Catal. Chiropt. Brit. IMus., p. 309 (Not

of Linnseus, 1758).
1886. Myotis tnurina var. spelxa Bielz, Verhandl. u. Mittheilungen des

Siebenbiirgischen Vereins fiir Naturwissensch. in Hermannstadt,

XXXVI, p. 83 (Homorod- Almas cave, Hungai'y).
1897. Myotis myotis Miller, Ann. and IMag. Nat. Hist., 6th ser., xx, p. 383,

October, 1897.
■1909. Myotis myosotis Miller, Ann. Mus. Zool. E. Univ. Napoli, N.S., iii,

No. 3, p. 1, April 26, 1909.
1910. Myotis myotis and M. m,. spelxa Trouessart, Faune Mamm. d'Europe,

p. 32.

Type locality. — ^Thiiringen, Germany.

Geographical distrihution. — Central and southern Continental
Europe, vv^est 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 31. 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 posteiiorly 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 . . .
Las hitherto only been taken in the gardens of the British Museum."

o

194

CHIROPTERA

conch, its greatest width (slightly above level of anterior base;
contained about 2^ times in length of anterior margin, the
posterior border conA^ex 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 })eculiarities 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. natter eri. 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 deej?
and interorbital concavity relatively shallow ; occipital region

riG. 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. mystacinns) ;
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 fi'om 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
«\ and ir^ trifid, but decidedly
oblique owing to reduction
in size of outer cusp ; a
minute cingulum cusp usu-
ally present at extreme outer
edge ; i^ with small postero-
internal tubercle ; i^ sub-
terete, 4-tuberculate, the me-
dian outer tubercle largest.
Canines relatively smaller
than in M. mystacinns and
with less developed posterior
cutting edge, their form
essentially as in M. nattereri
and M. einarginatus. 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 crow^n 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^ and m^ barely
indicated ; m^ 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

Fig. 33.
Myotis mrjotis. Teetli x 10.

196

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ill the small species, except that cingulum does not form a postero-
internal cusp behind entoconid ; m^ with hypoconid and ento-
conid more reduced than in the small species, the hyj)oconid
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 Tiigerwilen, 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.) ; Nlmes, 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.

Austria-Hungary : Herkulesbad, 1 ; Fiinfkirchen, S.W. Hungary, 2.

Roumania: Bustenari, 3; Sinaia, 1 (U.S.N.M.).

Switzerland : Geneva, 7 (Alottaz) ; Grotte de Vallorbe, Vaud, 1
(Mottaz) ; Boudry, Neuchatel, 2 (Mottaz) ; St. Moritz, 1 ; Thayugen, Schafl-
hausen, 2 (U.S.N.M.); Canton Thurgau, 5; Tiigerwilen, Thurgau, 10
(U.S.N.M.); Andermatt, Uri, 1 (U.S.N.M.); St. Gothard, Uri, 3 (B.M.
and U.S.N.M.) ; 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, Rome, 2 ; Marsala, Sicily, 6.

Sardinia : Oristauo, Cagliari, 10.

Spain : Seville, 2.

Portugal : Cintra, 2.

RemarJcs. — Mijotis myotis differs strikingly from the other
European members of the genus, 31. oxygnathus excepted, in its
much larger size. From the large Vespertilionidse 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. oxygnatlms ,• but it is the only large
species known to occur north of the Alps.

1 al. Nimes, Gard, France. G. E. Dobson (e). 80. 12. 14. 2.

1. Hamburg, Germany. Dr. J. E. Gray (p).

2 (J, 8?, Niesky, Silesia. {Dr. Dr. E. Hamilton (p). 97.12.4.7-17.
? juv. W. Baer.)

6 9. Strass, Burgheim, Ba- Lord Lilford (p). 11. 1. 1. 10-15.

varia. {Korbitz.)
?. Strass, Burgheim, Ba- Lord Lilford (p). 11. 1. 1. 124.

varia. [Korbitz.) 11. 1. 1. 131.

MY OTIS

199

?. Heidelberg, Badeu. Hon. N. C. Eoths- 10. 5. 29. 1.

child (p).

6 juv. Herkulesbad, Hungary. Hon. W. Rothschild 7. 9. 16. 8.

(F. J. Cox.) (p).

2 al. Fiintkirchen. Budapest Museum (e). 94.7. 18.11-12.

2 6,9. Bustenari, Prabova, Lord Lilford (p). 4.4.6.8-10.

840 m. Roumania.
(W. Dodson.)

1. St. Moritz.Grisons, Swit- Leon 0. Galliard (p). 75. 9. 20. 4.

5 9, Hhnxg&u.' [E. H. Zolli- 0. Thomas (p). 4.4.5.9-13.

Icofcr.)
2 9. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 26-27.

2. Ostia, Rome. Dr.L. Sambon (c&p). 1.1.2.1-2.

6 9. Marsala, Sicily. (.4. 0. Thomas (p). 6. 8. 4. 17-20.

Eohert.) 8. 9. 1. 3-4.

10 al. Oristano, Cagiiari, Sar- Hon. N. C. Roths- 7. 5. 24. 1-10.

dinia. child (p).

2 i al. Seville, Spain. {Dr. A. Lord Lilford (p). 73. 1. 8. 1-2.

liuiz.)

2 6. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 4-5.

MYOTIS OXYGNATHUS Monticelli.

1885. Vcspertilio oxygnathus Monticelli, Ann. Accad. O. Costa de Aspir.
Nat., I, p. 82. Type in Naples Museum.

1909. Myotis oxygnathus Miller, Ann. Mus. Zool. R. Qniv. Napoli, N.S.,

III, No. 3, p. 1, April 26, 1909.

1910. Myotis myotis oxygnathus Trouessart, Faune Mamm. d'Europe, p. 32.

Type locality. — Matera, Basilicata, Italy.

Geographical distribution. — Mediterranean region from .S2:)ain
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
tinger, 98 ; tifth finger, 76 ; ear from meatus, 23 ; width of ear,
13-6; tragus, 10*8. Two adult males from Velletri, Rome,
Italy : head and body, 60 and 62 ; tail, 58 and 58 ; tibia, 23
and 24 • 6 ; foot, 12-8 and 13 ; forearm, 53 • 6 and 57 ; thumb, 12
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

CHIROPTERA

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202

CHIROPTERA

14 and 14 ; forearm, 59 and 60 ; thumb, 11-4 and 11 -G ; 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.

Switzerland : Lugano, Ticino, 1 (U.S.N.M.).

Austria-Hungary: Bozen, Tirol, 3 (U.S.N.M.).

Italy : Finalborgo, Liguria, 2 (Genoa) ; Isoverde, 1 (Genoa) : Yallom-
brosa, Florence, 1 (U.S. KM.) ; Rome, 4; Velletri, Rome, 15 (U.S.N.M.) ;
INIatera, Basilicata, 1 (Naples, type).

Sardinia: Cagliari, 5 (U.S.N.M.); Monte Gennargentu, 3 (U.S.N.M.) ;
no exact locality, 3 (U.S.N.M.).

Malta : El Ghain, 5 ; Rubato, 3 ; no exact locality, 4.

Montenegro : Beri, 4.

Greece: Patras, 2; Corinth, 7 (U.S.N.M.); Nauplia, 1 (U.S.N.M.);
Lamia, Thessaly, 1 (U.S.N.M.)

Crete : Labyrinth, 4.

Tunis : No exact locality, 3 (U.S.N.M.).

Bemarks. — In general appearance Myotis oxygnathus resembles
31. 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 oxygnathus is, so far
as known, strictly Mediterranean, probably coincident with that
of 21. capaccinii and Pipistrellus kuhlii. North of the Alps
31. myotis occurs alone.

9. Burgos, Spain. G. S. Miller (c). 8. 8. 4. 10.

4 6. Rome, Italy. {Coll.) G. Barrett-Hamilton 11. 1. 2. 22-25.

3 6,2 9. El Ghain, Malta. Lord Lilford (p.) 11. 1. 1. 16^20.

(Micallef.)

3 6, ?. Rubato, Malta. ' Lord Lilford (p). 11. 1. 1. 21-24.

(MicaUcf.)

4. JIalta. (MicaUef.) Lord Lilford (p). 95. 3. 2. 3-6.

2 <J, 2 ?. Beri, 50m. Montenegro. O. Thomas (p). 5. 8. 4. 1-4.

{L. Filhrer.)

2 $. Patras, Greece. Hon. N. C. Roths- 8. 10. 2. 15-16.
(C. Mottaz.) child (p).

3 6, 9. Labyrinth, Crete. Miss D. Bate (c). 5. 12. 2. 5-8.

Genus PIPISTRELLUS Kaup.

1829. Pipistrellus Kaup, Entw.-Gesch. u. Natiiii. Syst. Europ. Thierwelt,

I, p. 98 [pipistrellus).
1887. Pipistrellus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx.

1838. Bomicia Gray, Mag. Zool. and Bot., ii, p. 495, February, 1838

{calcarata = kuhlii).

1839. Vesperugo Keyserliug and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 312 (part).
1856. H]jpsugo Kolenati, AUgem. deutsche Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 (m,aurus and krascheninikoivii).
V

PIPISTRELLUS

203

1856. Nannugo Kolenati, AUgem. deutsche Naturhist. Zeitung, Dresden,

neue Folge, ii, p. 131 {iiathusH, pipisti-ellus, and kuhlii).

1857. Vesperugo Blasius, Saugethiere Deutschlands, p. 49 (part).
1878. Vcsperugo Dobson, Catal. Chiropt. Brit. Mus., p. 183 (part).

1897. Pipistrellus Miller, Ann. and INlag. Nat. Hist., 6th ser., xx, p. 384,

October, 1897.
1899. Euvespcrugo Acloque, Faune de Prauce, Mammiferes, p. 35 (part,

included noctula, leisleri, maurus, kuhlii, pipistrellus, and

ahramus).
1907. Pipistrelkis IMiller, Families and Genera of Bats, p. 204, June 29,

1907.

Type specie.^. — Vespertilio pipisti-ellus Sclireber.

Geograpliical 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 Ejjtesicus (p. 224), but with 2 — 2 upper
j^remolars ; dental formula : i '^, c ^—\ pm l~?, m ;^,' = 34.

Remarks. — 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 ti-agus. 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 vesper tilionine 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 j)remolar ; 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 ; largo 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- kulilii, p. 215.

204

CHIKOPTERA

Outer upper incisor more thau half as high as

inner ; large upper premolar separated from

canine hy 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 ; condylobasal length of

skull 11 to 12 mm. ; thumb short, its length

about equal to width of wrist ; fifth finger

about 40 mm P. pipistrellus, p. 204.

Lower canine slender, the length of base along

cingulum slightly more than half length

of anterior border of shaft ; condylobasal

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. (France, based

primarily on Daubenton.)
1776. Vespertilio pipistrelle P. L. S. !Miiller, Natursyst. Suppl. u. Regist.-

Band, p. 16 (France, based on Schreber).
1825. Vespertilio pygmmus Leach, Zool. Journ., i, p. 560, January, 1825

(Dartmoor, Devonshire, England).
1834. Vespertilio brachyotos Baillon, M6m. Soc. Royale d'Emulatiou

d'Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1839. ? [Vespertilio pipistrellus'] var. nigra de Selys-Longchamps, Etudes

de Micromamm., p. 140 (nomen nudum).

1839. ? [Vespertilio pipistrellus] var. rufescens de S61ys-Longchamps, Etudes

de Micromamm., p. 140 (nomen nudum).

1840. Vlesp)ertilio'] pztsillus Schinz, Europ. Fauna, i, p. 9 (Synonym of

pipistrellus ; Brehm cited as authority).

1840. V\espertilio] vielanopterus Schinz, Europ. Fauna, i, p. 9. Brehm
cited as authority, but name apparently published here for first
time (Rhentendorf, Thiiringen, Germany).

1840. V[cs2xrtilio] stenotus Schinz, Europ. Fauna, i, p. 9. Brehm cited as
authority, but name apparently published here for first time
(Rhentendorf, Thiiringen, Germany).

1840. Vespertilio minutissimus Schinz, Europ. Fauna, i, p. 9 (Ziirich,
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. Vesper ugo pipistrellus Blasius, Saugethiere Deutschlands, p. Gl.

1862. Vesperugo pipistrellus var. macropterus Jeitteles, Verhandl. der k. k.

Zool. Bot. Gesellsch., Wien, xii, p. 250 (Kaschau, Hungary).

1863. [Nannugo pipistrellus'] var. typus Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 490. Not of
Bonaparte, 1845 (Wiesbaden, Hessen-Nassau, Germany).

PIPISTRELLUS

205

1863. [Nannngo pijnstrellus] var. flavescens Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 491 (Nassau,

Germany).
1863. [Nannugo inpistrellus] var. nigricans Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 491. Not of

Bonaparte, 1845 (Nassau, Germany).
1863. [Nannugo inpistrellus] var. lirnbatus Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassa,u, xviii, p. 491 (Siegen, Nassau,

Germany) .

1878. Vcsperugo pipistrellus Dobson, Catal. Chiropt. Brit. Mus., p. 223.
1897. Pipistrellus pipistrcllus Miller, Ann. and Mag. Nat. Hist., 6th ser.,

XX, p. 384, October, 1897.
1904. Pipistrellus pipistrellus mediterrancus Cabrera, I\Iem. Soc. Espaii.

Hist. Nat., II, p. 273 (Valencia, Spain).
1910. Pipistrellus pipistrellus and P . pipistrellus mediterraneus Trouessart,

Faune Mamm. d'Europe, pp. 14-15.

Type locality. — France.

Geographical distribution. — Europe from the Mediterranean
north to Scotland and Scandinavia, west to Ireland and the
Hebrides, east into Asia.

Diagnosis.- — Smallest European member of the genus (forearm,
27*6 to 32 mm. ; condylobasal length of skull, 11 to 12 ram.);
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

CHIROPTERA

the others) and extending nearly to point of elbow ; fifth finger
extending beyond elbow to a distance equal to less than one-
third length of forearm ; thumb short, its length about equal to
width of wrist ; membrane inserted at base of outer toe. Foot
about half as long as the short, robust tibia ; calcar considerably
longer than free border of interfemoral membrane, robust at base,
but tajjering rapidly and terminating without lobe, its keel well
developed, with evenly convex margin. Tail about as long as
body without head and 2^ times as long as tibia, the short
terminal vertebra free from membrane.

Far 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 Mijotis. 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

Pipistreilus 'pipis- though evidently more than half greatest length
treiius. 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
bull* 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

PIPISTRELLUS

207

inflated, the inflated region with a median angle suggesting
a rudimentary postorbital process ; rostrum short and broad,
narrowing gradual!}^ in front, a sHglit 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
emarginatiou 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 I'ather 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 developed. The main cusps of the two teeth lie
in line of general curve of anterior portion of tooth-row. Space
between outer incisor and canine about equal to greatest diameter
of incisor. Lower incisors forming a continuous, broadly IJ'shaped
row between canines, their crowns very slightly imbricated ;
crowns much longer than high, trifid, that of /\ nari'owest,
longest and lowest, that of i.^ and {3 widened posteriorly but
without additional cusps or tubercles. Upper canine robust,
the greatest diameter of its crown about three-quarters length
of anterior border of shaft, the cross section of shaft broadly
triangular with longest side formed by nearly flat postero-internal
surface ; a sharply defined antero-exteral longitudinal groove, and
less definite postero-external concavity ; anterior edge narrow
but not strictly trenchant ; posterior edge trenchant, with well
marked angle slightly below middle, this angle frequently becom-
ing a distinct secondary cusp ; cingulum well developed but not

208 CHIROPTERA

forming true basal cusps. Mandibular canine low and heavy,
its apex scarcely rising above level of highest molar cusps, its
greatest diameter measured along cingulum nearly or quite equal
to length of anterior border of shaft ; cingulum well developed,
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
narrow and flattened-concave, with evident
elevated rim, the posterior border strongly
concave, the anterior border usually convex
= but occasionally a little concave ; height of
main cusp slightly greater than that of highest
Fig. 35. molar cusps and about equal to length of

Anterior\eelh X 5!*^ 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 conspicuovisly
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^'ttern normal ; m^ with crown area about
two-thirds that of m^, 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^ and m.,, but with less in m^ ; cingulum
well developed, forming a slight postero-internal cusp behind
entoconid.

Measurements. — Adult male from Henley-on-Thames, Oxford-

PIPISTKELLUS

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 j ear from meatus, 11 -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 • 0 ; third
finger, 49-53 ; fifth finger, 34 • 6-40. For cranial measurements
see Table, p. 210.

Specimens exmnined. — Two hundred and seventy -uine, 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.

Denmaek : Hillerod, Zealand, 1.

Prance : Boulogne -sur-Mer, Pas-de-Calais, 2 ; Etupes, Doubs, 1
(Mottaz) ; Nimes, Gard, 3 (Mottaz and Nimes, the last wrongly marked
type of nigrans Crespon) ; St. Genies, Gard, 1 (Mottaz) ; Marseilles, 1
(U.S.N.M.)

Germany : Bonn, 4 ; Ingelheim, Rheinhessen, 1 ; Magdeburg, Saxony, 1 ;
Berlin, 1 ; Rudolstadt, Bavaria, 3 ; Niesky, Silesia, 1.

Austria-Hungary : Transylvania, 1 ; Zara, Dalmatia, 1.

Switzerland : Geneva, 15 (Mottaz) ; Buchillon, Vaud, 4 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Neuchatel, 1 (Mottaz) ; Cortivallo, Ticino,
1 (U.S.N.M.) ; Mt. San Salvatore, Ticino, 13 (U.S.N.M.).

Italy: Campiglio, Tirol, 1; near 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.).

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 Real, Madrid, 2 ; Granada, 1 ; Seville, 1 ; Alcala, near
Seville, 1.

Remarks. — Pipistrellus plpistrellus is the smallest as well as
one of the commonest and most generally distributed of European

p

210

CHIROPTERA

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212

CHIROPTERA

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 Pij^istreUus 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.

6 al. Co. Longford, Ireland.
? St. Co. Antrim.
9. Alnwick, Northum-

berland, England.
2 6,2 9. Tring, Hertfordshire.
9. Lilford, Northampton-

shire.

2 9. Chelmsford, Essex.

9 juv. al. London.

6. Wimbledon, Surrey,

juv. New Forest, Hamp-
shire.
9 al. Netley, Hampshire.
9. Loddiswell, Devon-

shire.

3 9. Upsala, Sweden.

(Kolthoff.)

5. Hillerod, Zealand,

Denmark.

5, 9. Boulogne, Pas - de -

Calais, France.

c5, 9 al. Bonn, Rhineland,
Germany.
9. Ingelheim, Rhein-
hessen.

9 al. IMagdeburg, Saxony.

6 al. Berlin.

9 al. Transylvania, Hun-
gary.

6. Zara, Dalmatia.

2 al. Ciudad Real, Spain.
9. Burgos, Prov. Burgos.
9. Silos, Burgos.

1 al. Canipiglio, Tirol,
Italy.
6 juv. al. Sicily.
14 al. Aristano, Cagliari,
Sardinia. (C.
Kratisse.)
2 juv. al. Sardinia. (P. Bonomi.)

3 9. Tatoi, Athens, Greece.

(C. Mottaz.)

6, 3 9. Kephissia, Athens.

Dr. G. E. Dobson (p).
Hon. N. C. Rothschild (p).
W. E.de Winton (p).

Hon. N.C. Rothschild (p).
Lord Lilford (p).

M. Christy and E. L.

Thompson (p).
Dr. A. Giinther (p).
C. H. B. Grant (c & p).
Col.J.W.Yerbury(c&p).

Dr. G. E. Dobson (c & p).

Col.J.W.Yerbury(c&p).

Lord Lilford (p).

0. Thomas (c & p).

0. Thomas (c & p).

Dr. A. Giinther (p).

C. H. Hilgert (c).

Dr. W. Wolterstorff (p).

Dr. A. Giinther (p).

C. G. Danford and J. A.

Brown (c & p).
Lord Lilford (p).
A. Cabrera (p).
G. S. Miller (c).
G. S. Miller (c).
G. C. Champion (c & p).

76. 2. 12. 1.

I. 9. 3. 6.

II. 1. 3. 390.

9. 2. 19. 1-4.
11. 1. 1. 125.

11. 1. 3. 23-24.

74. 7. C. 1.
11. 1. 3. 27.
11. 1. 3. 389.

76. 11. 3. 1.
11. 1. 3. 25.

11. 1. 1. 26-28.

96. 6. 7. 1.

98. 1. 9. 1-2.

Hon. N. C. Rothschild (p).

E. N. Buxton (p).

Hon. N. C. Rothschild (p).

C. Mottaz (c).

8. 11. 2. 3-4.

92. 12. 1. 2.
66. 2. 1. 22.

74. 7. 4. 6.

11. 1. 1. 29.
8. 7. 23. 2-3.
8. 8. 4. 14.
8. 8. 4. 15.
96. 8. 7. 1.

46. 6. 15. P.
7. 5. 24. 11-24.

95. 4. 16. 4-5.
8. 10. 2. 19-21.

8. 11. 3. 4-7.

PIPISTRELLUS 213

PIPISTRELLUS NATHUSII Kejserling and Blasius.

1839. V[cspe7-tilio] natlmsii Keyserliug and Blasius, Wiegmann's Archiv

fiir Naturgesch., 1839, i, p. 820 (Berlin, Germany).
1857. Vesperiigo nathusii Blasius, Saugethiere Deutschlands, p. 58.
1878. Vesperugo abramus Dobson, Catal. Chiropt. Brit. Mus., p. 226 (Part:

not of Temminck).
1900. Pipistrelhcs nathusii Mehely, Monogr. Chiropt. Hungarise, p. 276.
1905. Ycsp[erugo'] nathusii var. imicolor Fatio, Arch. Sci. Phys. et Nat.,

Geneve, 4th ser., xix, p. 510, IMay, 1905 (Geneva, Switzerland).

Type in Geneva Museum.
1910. Pipistrcllus abramus Trouessart, Faune Mamm. d'Europe, p. 16.

Type locality. — Berlin, Germany.

Geograjjhical distribution. — Central and southern Continental
Europe ; exact limits of range not known.

Diagnosis. — Not so small as Pipistrelliis piinstrellus (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 lifth 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
PipistreUus pipistreUus. 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. pipistreUus and similarly blunt at tip, but with
posterior border more evidently convex ; posterior basal lobe
small, usually less well defined than in P. pipjistrellus. Wing
larger and relatively broader than in P. pipAstrellus, 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. pipistreUus.

Fur and colour. — Fur slightly more loose in texture than that
of P. pApistrellus, 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 PipistreUus pipistreUus, though usually distinguishable by a

214

CHIROPTERA

tendency away from the cinnamon and raw-umber tints toward
a clearer brown more resembling Eidgway's mars-bro^vn. Mem-
branes less blackish than in P. jjipistrdlus, the wing from foot
nearly to fifth finger with a noticeable jjale border about 1 mm.
in width, similar to that present in P. Jcnhlii, but less sharply
defined and less nearly white.

SJiiiU. — The skull is less diminutive than that of PijjistreUus
jpipistrellus, its general size about as 3It/oiis mystachms. General
form less robust than in P. pijristrellKS, 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 bj^ a slight transverse constriction.
Other details of form essentially as in P. pipistrellus.

Teeth. — As compared with those of Pijnstrellus pipistrellits 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 cusp ; outer
upper incisor distinctly larger than inner, its apex extending
noticeably beyond secondary cusp of inner tooth, its general form
essentially as in P. pipistrellus, but inner margin without evident
secondary cusp. Lower incisors less crowded than in P. pipis-
trellus, a slight space usually present in
median line, another between i^ and i.^, and
another between io and canine ; outer edge
of «\ slightly overlapping i.^ > ^^ 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

PipUtrelius nathusii. rising distinctly above that of highest molar
Anterior teeth x 5. cusps, its greatest diameter measui-ed along
cingulum equal to a little more than half
length of anterior border ; apex of anterior cingulum cusp not
rising above level of basal thii'd of posterior border. Upper
premolars as in P. pipisireUus, 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. pipisireUus.
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 exti'emes of six
adults from Buchillon, Vaud, Switzerland : tibia, 12 • 9 (12 • 6-14) ;
foot, 7-3 (6 -8-7 -6); forearm^ 03 (32 • 4-34 -G); thumb, 5*7

PIPISTRELLUS

215

(5-2-6); third finger, 61-3 (58-65); fifth finger, 44-6 (41-47).
Adult male and female from Florence, Italy : head and body,
46 and 47; tail, 40 and 38; tibia, 12-8 and 12-6; foot, 6-8
and 7 ; forearm, 35 and 33 ; thumb, 6 • 2 and 6*6; third finger,
63 and 62 ; fifth finger, 47 and 46 ; ear from meatus, 12 "6 and
12-6 ; width of ear, 11 and 11-4. For cranial measurements see
Table, p. 222.

Specimens examined. — Thirty-three, from the following localities : —

Fbance : St. Gilles, Gard, 1.

Germany : Berlin, 1 ; Bavaria, 1 (U.S.N.M.) ; Ingelheim, Rhein-
hessen, 1.

Switzerland: Geneva, 6 (Mottaz and Geneva, including type of
unicolor Fatio) ; Montreux, Vaud, 1 (Mottaz) ; Buchillon, Vaud, 6 (Mottaz) ;
Neuchatel, 1 (U.S.N.M.) ; Canton Uri, 1 ; St. Gothard, Uri, 1 (U.S.N.M.).

Austria-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.).

Bemarhs. — Though readily distinguishable from the other
European members of the genus by its cranial and dental
characters, Pipistrellus natlmsii is superficially much like P. ^jipis-
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, Rheinhessen. C. Hilgert (c). 8. 11. 2. 3.

2 al. Palics, Bacser, Hungary. Budapest jNIuseum (e). 0. 4. 9. 1-2.

6. Canton Uri, Switzerland. Tomes Collection. 7. 1. 1. 398.

6,9. 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 KUHLII Kuhl.

1819. Vespertilio kuhlii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 2, p. 199 (Triest).
1835. Vesp[ertilio'] alholimbatus Kiister, Isis, p. 75 (Cagliari, Sardinia).
1837. Vespertilio vispistrellus Bonaparte, Iconogr. Faun. Ital., i, fasc. xs

(near Rome, Italy). Type in British Museum.

1837. Vespertilio alcythoe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi

(Sicily.) Type in British Museum.

1838. Romicia calcarata Gray, Mag. Zool. and Bot., ii, p. 495 (locality

unknown). '
1811. Pipistrellus marginatus Bonaparte, Iconogr. Fauna. Ital., Indie.

distrib., nomencl. mod. (Substitute for albolimhatus).
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

CHIROPTERA

1840. Vlesjyertilio] ursula Waguer, Schreber's Saugthiere, Suppl., i, p. 505

(Morea, Greece).
1857. Vesperugo kulilii Blasius, Siiugethiere Deutschlands, p. 63.
1878. Vesperugo kiihlii Dobson, Catal. Chiropt. Brit. Mus., p. 230.
1886. [Vesperugo kulilii] var. albicans Monticelli, Atti Soc. Ital. Sci. Nat.,

Milauo, XXVII, p. 200, March, 1886 (Caivano, Naples, Italy).
1886. [Vesperugo kulilii] \SiV. pullatus Monticelli, Atti Soc. Ital. Sci. Nat.,

Milano, xxvii, p. 200, IMarch, 1886 (Bella Vista, near Portici,

Naples, Italy).
1900. Pipistrellus kulilii Mehely, Monogr. Chiropt. Hungarise, p. 261.
1910. Pipistrellus kulili Trouessart, Paune Mamm. d'Europe, p. 17.

Type locality. — Trieste, Austria-Hungary.

Geographical distribution. — Mediterranean region and eastward
into Asia.

Diagnosis. — Size about as in PipistreUus natJmsii (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. natJmsii ; tragus with greatest width less than
length of anterior border ; thumb short, its length about equal to
width of wrist ; posterior edge of wing membrane with sharply
defined whitish border.

External 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
diflferent 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. pninstrelhis 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
inpistrellus 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. ])ij)istrellus, 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. nathusii. Anterior .^ \i uu ijjj G>f-i
upper premolar crowded inward from tooth-
row and closely wedged between canine and
large premolar which are nearly or quite in Pij^Mi^kuMU.
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. pipistrelhis and P. nathusii, 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, 12 and 13 ; foot, 6 • 2 and 6 • 2 ; forearm, 33 • 4 and 3.5 ;
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

CHIROPTERA

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PIPISTRELLUS 219

Specimens exaviincd. — One hundred and eight, from the following
localities :^

Fbance: St. Gilles, Gard, 2; Mmes, Gard, 3 (Mottaz) ; St. Genies,
Gard, 4 (Mottaz) ; Marseilles, 1 (U.S.N.M.).

Switzerland: Coremmo, Ticino, 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 vispistrcllus Bonaparte) ; Sorrento, 3 (U.S.N.M.) ;
Catanzaro, Calabria, 6 (U.S.N.M.); Palermo, Sicily, 23 (U.S.N.M.);
Corleone, Sicily, 1 (U.S.N.M.); Ustica Island, Sicily, 4 (U.S.N.M.);
Sicily, 1 (type of alcytlioc Bonaparte).

Sardinia: Cagliari, 4 (U.S.N.M.) ;' Mt. Gennargentu, 7 (U.S.N.M.);
no exact locality, 2.

Gebece : Corfu, 2 ; Cepbalouia, 10 ; Patras, 3.

Spain: San Cristobal, Minorca, Balearic Islands, 2.

BemarJcs. — Pipistrellus kuhlii is easily recognizable by the
form and relative size of the upper incisors. Externally it naay
usually be known by the sharph^ defined whitish border to the
wing membrane, though too much reliance should not be placed
on this character alone. Many specimens from Sardinia are
Hghter in colour than those from the mainland. These represent
the albolimhatus of Kiister. Normally coloured examples also
occur ; and in the absence of adequate material it has seemed
preferable for the time being not to attempt to define the insular
form.

s.

St. Gilles, Gard,

G. S. Miller (c).

8. 8. 4. 127.

6.
3 6, ?.

France.
Siena, Italy.

{S. Brogi.)
Rome. (C. Coli.)

Dr. B. Hamilton (p).
G. Barrett-Hamilton

98. 10. 2. 2.
11. 1. 2. 16-1".

skeleton with- Sicily. (Prince Bona- Tomes Collection. 7.1.1.730.

out skull. parte.) [Type of V. alcythoe Bonaparte.)

skeleton with- Near Rome, Italy. Tomes Collection. 7. 1. 1. 729.

out skull. (Prince Bonaparte.) (Type oi V. vispistellus 'Bona.T^a.vte.)

6, ?. Corfu, Greece. J. I. S. Whitaker (p). 8. 10. 1. 4-5.

(C. Mottaz.)

3 6. Argostoli,Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 1-3.

(C. Mottaz.)

(J,?. Patras. (C. Mottaz.) Hon. N.C. Rothschild 8. 10. 2. 17-18.

2 6. San Cristobal, 0. Thomas and R. I. 0.7.1.29-30.

Minorca, Balearic Pocock (c & p).
Islands.

PIPISTRELLUS SAVII Bonaparte.

18.37. Vespc7-tilio savii Bonaparte, Iconogr. Faun. Ital., i, fasc. xs (Pisa).

Type in British Museum.
1837. Vespertilio aristippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi

(Sicily).
1837. Vesjwrtilio leucippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi

(Sicily). Type in British Museum.
1833. Vespertilio honapartii Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvii,

p. 226 (Tuscany).

220

CHIROPTERA

18i4. Vesplertilid] nigrans Crespon, Faune IMeridionale, i, p. 24 (Nimes,

Gard, France).
1853. Vesper ago maxirus Blasius, Wieginaun's Archiv fiir Naturgesch.,

1853, I, p. 35 (Central chain of the Alps).
1857. Vespemgo maurus Blasius, Saugethiere Deutschlands, p. 67.
1872. V[espertilid'\ agilis Fatio, Fauna 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 ocliromixtus Cabrera, Mem. Soc. Espafi. Hist. Nat., ii,

p. 267 (Sierra de Guadarrama, Madrid, Spain).
1910. Pipistrellus savii and P. savil ocliromixtus Trouessart, Faune ]\Iamm.

d'Europe, pp. 13-14.

Tijpe locality. — Pisa, Italy.

Geographical (listrihution. — 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.— LiSLVgest 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 hulilii
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 savii.
Nat. size.

conspicuous, a peculiar dull brownish ochraceous-bufi'; adult female :
light tips as conspicuous as in last, dull brownish cream-buff.

SJciill. — The skull is slightly larger than that of PtjnstreJhtyi
nathusii and P. kuhlii, 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. Tcnhlii, 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. hihlii,
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 pipnstrellus, 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»

Fia. 39.

Pipistrellus savii.
Anterior teetli. X 5.

222

CHIROPTERA

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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,
.-Sij 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 : —

SwiTZEELAND : 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).

BemarJcs. — This species is readily distinguishable among the
European members of the genus Pipistrellus by the pecuhar form
of the tragus, apart from its very pronounced cranial and dental
characters. Its colour gives it a superficial resemblance to
Eptesicus 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). 46. 11. 1. 3.

4 al. Genoa, Liguria, Italy. Genoa Museum (e). 86. 11. 3. 14-

17.
6. 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. France. Purchased (Lefebre). 46. 1. 2. 12.

6. 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. Cnepheeus Kaup, Entw.-Gesch. u. Natiirl Syst. Europ., Thierwelt, i,

p. 103 {serotinus).
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 312 (part).
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 313 (Sub-genus of Vesperitgo, 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, fasc. xxi, in account of

Vespertilio alcytlioe (serotinus).
1856. Catcorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,

neue Folge, ii, p. 131 [serotinus).

1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,

neue Folge, ii, p. 131 (part).

1857. Vespems Blasius, Siiugethiere Deutschlands, p. 51 (Sub-genus of

Vcspenigo), part.

1858. Ambhjotus Kolenati, Sitzungsber. kais. Akad. Wissensch. Wien,

Math.-Naturwissensch. Classe, xxix, p. 252 (atratus — nilssoni).
1868. Aristippe " Kolenati, Beitrage zur Kenntniss der Phthiriomyiarien,

Petersburg, 1863 " (part, included both discolor = murinus and

nilssoni).
1866. Pachyomits Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 90,

February, 1863 {pachijomus).
1870. Nyctiptenus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,

Math.-Naturwiss. Classe, lxii, p. 424 (smithii).
1878. Vesperus Dobson, Catal. Chiropt. Brit. Mus., p. 184 (Sub-genus of

Vesperugo), part.
1892. Adclonycteris 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 M6hely, Monogr. Chiropt. Hungarise, p. 219 (part).
1907. Eptesicus Miller, Families and Genera of Bats, p. 207, June 29, 1907.

Type species. — Eptesicus melanops Rafinesque = Verpertilio
fiiscus Beauvois.

Geographical distribution. — Europe, Asia (except Malay region),
Australia, Africa, Madagascar ; America from southern Canada
southward (except Lesser Antilles).

Characters. — Dental formula : i ;-?, c ^, pm ]f^^, m ?4! = 32.
Teeth strictly normal throughout, and showing no special pecu-
liarities. Both upper incisors well developed, the inner larger
than the outer and usually with distinct secondary cusp, the
outer separated from canine by a space equal to its greatest
diameter ; m^ variable in form, usually with well developed
metacone and three commissures in the smaller species, but with
metacone and third commissure obsolete in larger forms. Skull
without special peculiarities of form or structure, the rostrum
ilattish 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).

Hemarhs. — Among European bats the members of the genus
Epjtesicus may be distinguished by their dental formula combined
with a simple Pijjistrellus-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 CHIROPTERA

KEY TO THE EUEOPEAN SPECIES OP EPTESICUS.

Forearm 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. nilssonii, p. 234.

Forearm 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. 22G.

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). France,

based primarily on " La Serotine " 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 Fledermaus" of Schreber).
1827. Vespertilio ivieclii Brehm, Ornis, Heft iii, p. 24 (Renthendorf,

Thiiringen, Germany).
1827. Vespertilio okenii Brehm, Ornis, Heft iii, p. 25 (Renthendorf,

Thiiringen, Germany).
1844. Vesp[ertilio'] incisivus Crespon, Faune M6ridionale, i, p. 26 (Nimes,

Gard, France).
1857. Vesperugo serotinus Blasius, Saugethiere Deutschlands, p. 76.
1863. [Cateorus serotimis'] var. typus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 466 (Wiesbaden, Nassau,

Germany).
1863. \_Cateorus serotinus'] var. rufescens Koch, Jahrb. des Vereins fiir

Naturkdnde im Herzogthum Nassau, xviii, p. 466 (Freiburg,

Breisgau, Germany).
1878. Vesperugo serotinus Dobson, Catal. Chiropt. Brit. Mus., p. 191.

1885. Vesperus serotinus var. transylvanus Daday, Orvos-Terraeszettudo-

manyi ErtesetS, Kolozsvar, x, p. 275 (Als6-Sz6cs, Szolnok-Doboka,
Hungary).

1886. Vesperus serotinus var. transsylvayius Daday, Verhandl. u. Mittheil-

ungen des Siebenbiirgischen Vereins fiir Naturwissensch. in

Hermannstadt, xxxvi, p. 81.
1900. Eptesicus serotinus M6hely, Monogr. Chiropt. Hungariae, p. 209.
1904. Vespertilio serotinus insidaris Cabrera, Mem. Soc. Espaii. Hist. Nat.,

II, p. 263 (Minorca, Balearic Islands).
1904. Vespertilio isabellinus Cabrera, Mem. Soc. Espau. Hist. Nat., ii,

p. 264 (southern Spain). Not of Temminck.
1904. Vespertilio boscai Cabrera, Mem. Soc. Espan. Hist. Nat., ii, p. 265

(Muchamiel, Alicante, Spain).
1910. Eptesicus serotinus, E. serotimis transsylvanus, and E. boscai Troues-

sart, Faune Mamm. d'Europe, pp. 20-22.

Ty2^e 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.,

EPTESICUS 227

condylobasal length of skull more than 19 mm.) ; colour of upper
parts a yellowish brown without noticeably contrasted light tips
to the hairs ; under parts essentially similar, so that there is no
line of demarcation along sides of neck.

External characters. — General form robust, though less so
than in Nt/ctalus, the tail and legs rather short, the membranes
thick and opaque. Muzzle with moderately prominent glandular
swelHngs, its greatest width across this region Icvss than distance
from nostril to ear ; nostrils projecting very slightly, the concavity
between them not conspicuous, the orifice crescentic. Ear
moderately long, extending slightly more than half way from eye
to nostril when laid forward, its breadth when flattened about
equal to height above crown ; anterior border of conch abruptly
convex below, then nearly straight to narrowly rounded 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. — Fur soft and dense, the longest hairs on
back about 10 mm. in length, those of underparts shorter. It
is strictly confined to body, only extending as a thin pubescence
on extreme base of membranes and along a narrow line bordering
under surface of forearm ; free edge of uropatagium naked.
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

CHIROPTERA

Slcull. — 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 Nijctalus nociida, the straight, well developed
lambdoid crests which form its posterioi' 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 bullie small, the transverse diameter con-
siderably less than distance between bulla?. Interorbital region
moderately constricted, hour-glass shaped, the lachrymal region

decidedly less wide than brain-case,
with slight tubercular projection close
to anterior rim of orbit ; rostrum
flattened, with shallow but evident
lateral concavity on each side, dis-
tinctly narrower anteriorly than pos-
teriorly, the narrowly obovate narial
emargination extending about half
way back to level of lachrymal fora-
men ; rostral depth at front of orbit
less than distance from orbit to outer
incisor ; anteorbital foramen less i*e-
duced than in Ni/ctalus noctula, its
posterior border over region of con-
tact between large premolar and first
molar, lachrymal foramen directly
behind it, on inner side of orbital
rim. Palate long and narrow, slightly concave both laterally and
longitudinally, the anterior emargination small, squarish, extend-
ing back to level of middle of canine. Posterior extension of
palate nearly parallel sided, though narrowing a little posteriorly,
its width at level of 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-

FlG. 40.
Eptesieus serotinus. Nat. size.

EPTESICUS

229

quarter that of canine, the subterete shaft directed inward and
sUghtly 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, the two concavities some-
times separated by a low but evident ridge ; inner margin with
a low secondary cusp on well developed cingulum. A line
perpendicular to main axis of skull would pass through centre
of shafts of all four incisors ; the outer tooth is separated from
canine by a space about equal to breadth of its own crown. Lower
incisors large, much crowded, and very conspicuously imbricated,
e'l and i^ 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 12) ', cross section of crown somewhat triangular, the
posterior angle occupied by a low tubercle in i.^ and i^. 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 nodula ;
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 gi'oove ;
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^, 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
moi-e 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

CHIROPTERA

columnar thickening of posterior base of protocone ; paracone
lower and smaller than metacone, the contrast unusually notice-
able ; styles and commissures well developed, the VV-p^ttern
normal ; m? with crown area less than half that of m^, its longi-
tudinal diameter through metacone much less than half transverse
diameter, the mesostyle, metacone, and second and thii'd com-
missures greatly reduced, though not sufficiently to lose their
identity. Lower molars with no special peculiarities ; angles in
commissures between outer and inner cusps rather wide and
shallow, especially that between protoconid and metaconid ; area
of second V in '"3 scarcely half that of first.

Measurements.— AA\i\t 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, 16. Adult
male and female from Seville, Spain : head and body, 67 and 64 ;
tail, 46 and 51 ; tibia, 21*6 and 22-4 ; foot, 11 and 10-4 ; fore-
arm, 49 and 50 ; thumb, 9 • 2 and 8 • 4 ; third finger, 89 and 90 ;
fifth finger, 64 and 68; ear from meatus, 17*6 and 18. Adult
male and female from Rome, Italy : head and body, 72 and 73 ;
tail, 54 and 54 ; tibia, 21-4 and 21 ; foot, 11 and 11 '4 ; fore-
arm, 51-6 and 53 ; thumb, 9 • 6 and 9*8; third finger, 91 and 92 ;
fifth finger, 66 and 67 ; ear from meatus, 19 and 19 ; width of
ear, 15 '6 and 15*4. For cranial measurements see Table,
p. 232.

Specimens examined. — Seventy-eight, from the following localities: —

England: Kenley, Surrey, 1 ; Hawkhurst, Kent, 1; Whitstable, Kent, 2 ;
Wingham, Dover, Kent, 1 ; Yalding, Kent, 5 (B.M. and U.S.N.M.) ; Isle
of Wight, 5.

Prance : Barsac, Gironde, 1 (U.S.N.M.) ; near Nimes, Gard, 2 (Nimes ;
type of incisivus Crespon, and a specimen wrongly marked type of palustris
Crespon).

Germany: Ingelheim, Rheinhessen, 2; Magdeburg, Saxony, 2 (B.M.
and U.S.N.M.) ; Herrnhut, Saxony, 1 (U.S.N.M.) ; Strass, near Burgheim,
Bavaria, 3; Bavaria, no exact locality, 2 (U.S.N.M.) ; Tubingen, Silesia, 2.

Austria-Hungary : Moravia, 1 ; Csallokoz-Somorja, Pressburg, Hun-
gary, 1 ; Budapest, Hungary, 2 ; Transylvania, 5 ; Zara, Dalmatia, 2.

RouMANiA : Bustenari, Prahova, 3.

Greece: Patras, 14; near Athens, 1.

Italy : Siena, 3 (B.M. and U.S.N.M.) ; Florence, 2 (B.M. and U.S.N.M.) ;
Vallombrosa,! (U.S.N.M.); Volterra, 1 (U.S.N.M.); Rome, 3 ; SanMartino
al Cimino, Rome, 1 (Genoa) ; Ustica Island, Sicily, 1 (U.S.N.M.).

Sardinia : Cagliari, 2 (Genoa).

SrAiN : Pajares, Leon, 1 ; Seville, 3 ; Muchamiel, Alicante, 1 (Madrid ;
type of boscai Cabrera).

Bemarks. — Among the bats of Europe this species is recogniz-
able by its rather large size, noticeable free tip to the tail,
moderately long, narrow ear, and straight, erect tragus. With
the material now available for study it seems impossible to

EPTESICnS

231

distinguish any local geographical forms. Specimens from Seville
representing the isabelUnus of Cabrera I am unable to separate
from true serotinus ; the type of hoscai Cabrera is a young of the
same animal ; insularis I have not seen, but there is nothing in
the original description to indicate that it is distinct.*

2 9.
9al.
1st.

3 9.

9al.
2.
1.

1.

6, 9.

2 S, 9.

2al.

1 al.

2.

2al.
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6, 9.
6, 9.

19.

9.
9.

lal.

6.
9.

9al.

1 al.

1.

1910.

Kenley, Surrey, England.

Whitstable, Kent.
Wingham, Kent.
Yalding, Kent.
Yaldiug, 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.
[Kiirbitz.)

Tubingen, Silesia.

Moravia, Hungary.

Csallokoz-Somorja, Press-
burg.

Budapest.

Transylvania.

Zara, Dalmatia. {K. Bios.)
Bustenari, Prahova, 840 m.
Roumania. {W.Dodson.)
Patras, Greece. {C.Mottaz.)

W. R. Ogilvie-Grant

(c & p).
C. H. B. Grant (c & p).
G. Donker (c & p).
H. Reid (c & p).
W. R. Ogilvie-Grant

(c & p).
Rev. G. Bury (c & p).
F. Bond (c & p).
Tomes Collection.

Tomes Collection.
C. Hilgert (c).

Patras.
Athens.

(C. Mottaz.)

Florence, Italy.
Rome. (C. Coli.)
Pajdres, Leon,

(iV. Gonzalez.)
Seville.
Seville.
Europe.

Spain.

Lord Lilford (p).

Lord Lilford (p).

Dr. A. Giinther (p).
Purchased (Parreys).
Budapest Museum (e).

Budapest Museum (e).
C. G. Danford and J. A.

Brown (c & p).
Lord Lilford (p).
Lord Lilford (p).

Hon. N. C. Rothschild

C. 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-Hamiltoi

Vespertilio sodalis Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
8th ser., v, p. 291, March, 1910. Type in British Museum.
1910. Eptcsicus 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. Espan.
Hist. Nat., VI, p. 449, December, 1908).

232

CHIROPTERA

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Greece : Patras

EPTESICUS

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234

CHIROPTERA

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.

Roumania. {W. Dodson.) {Type of species.)

EPTESICUS NILSSONII Keyserling and Blasius.

1836. Vespertilio kulilii Nilsson, Ilium. Fig. Skand. Fauna, pt. 17, pi. 34
upper figure. Not of Kuhl, 1819.

1838. Vespert\ilio] borealis Nilsson, Blum. Fig. Skand. Fauna, pt. 19,

pi. 34 (renumbered 36) upper figure. Not of P. L. S. Miiller, 1776
(Scandinavia).

1839. V[espertilio'] nilssonii Keyserling and Blasius, Wiegmann's Archiv

fiir Naturgesch., 1839, p. 315 (Mountains of Scandinavia. Based
on the V. kuhlii of Nilsson, 1836).

1857. Vesperugo nilssonii Blasius, Saugethiere 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) Ehoads, Reprint 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.

Tyjje locality.^^weden.

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-
parts 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 mastoifl
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. serotinui<. Interorbital fig. 41.

region relatively less constricted than in Eptesicus wUsonii.
E. serotinus, but of essentially the same form ; ^at. 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

CHXROPTERA

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
Ej)tesicus serotinus, but disproportion between paracone and
metacone less evident ; m^ with crown area about two-thirds that
of to\ 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 ^^ '"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.

Austria-Hungary : Csallokoz-Somorja, Pressburg, 1.

Switzerland : Grotte de Vallorbe, Vaud, 1 (Mottaz).

RemarJcs. — This species is readily distinguishable from Eptesicus
serotinus by its smaller size and by the conspicuous pale tips to
the hairs of back. From Vespertilio 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-
ficiall}^. Pijnstrellus sarii, 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. R. Alston (p.) 81. 6. 9. 2.

4 9. Upsala, Sweden. {Kolthoff.) Lord Lilford (p). 11.1.1.39-42.

i, 9. Wernigerode, Saxony, Ger- Dr. W. Wolterstorff 0. 2. 8. 3-4.
many. (c & p).

1. Csallokoz-Somorja, Press- Budapest Museum (e.) 94.3.1.14.
burg, Hungary.

EPTESICUS

237

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238 CHIROPTERA

Genus VESPERTILIO Linnaeus.

1758. Vespertilio Linnaeus, Syst. Nat., i, 10th ed., p. 31 (murinus by

tautonymy).
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur-

gesch., 1839, i, p. 312 (part).
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fiir Naturgesch.,

1839, I, p. 313. Sub-genus of Vesperugo (part).

1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeituug, Dresden,

neue Folge, ir, p. 131 (part).

1857. Vesperus, misprinted Vesperugo Blasius, Saugethiere Deutschlands,

p. 69 (Sub-genus of Vesperugo).
1863. '■^ Aristippe Kolenati, Beitrage zur Kenntniss der Phthiriomyiarien,

Petersburg, 1863" (jDart).
1872. Marsipolsemus Peters, Monatsber. k. preuss. Akad. Wisseusch.,

Berlin, p. 260 (Sub-genus of Vesperugo for alhigularis — inurinus).
1878. Vesperus Dobson, Catal. Chiropt. Brit. Mus., p. 183. Sub-genus of

Vesperugo (part).
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,

October, 1897 (part).
1900. Vespertilio Mehely, Monogr. Chiropt. Hungarias, p. 219 (part).
1907. Vespertilio Miller, Families and Genera of Bats, p. 209, June 29, 1907.

Type species. — Vespertilio murinus Linnasus.

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.

Hemarks. — The genus Vespertilio as now restricted contains
only two species, V. inurinus 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 Linngeus.

1758. IVespertilio] rmirinus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden).

1819. Vespertilio discolor Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 187 (Vienna, Austria).

1857. Vesperugo discolor Blasius, Saugethiere Deutschlands, p. 73.

1872. Vesperus [Marsipolmmus) alhigularis Peters, Monatsber. k. preuss.
Akad. Wissensch., Berlin, p. 260 (Type supposed to have been
taken in Mexico ; for its reference to this species see M6hely,
Monogr. Chiropt. Hungariae, pp. 229, 341, 1900).

1878. Vesperugo discolor Dobson, Catal. Chiropt. Brit. Mus., p. 204.

1885. Vesperus siculus Daday, Orvos-Termeszettudomdnyi Erteseto, Kolozs-
var, X, p. 275 (Homorod-Almas cave, Hungary). See Mehely,
Monogr. Chiropt. Hungarise, pp. 229, 346, 1900.

VESPEETILIO 239

1886. Vespenis siculus Daday, Verhandl. u. Mittheilungen des Siebenbiirg-
ischen Vereins fiir Naturwissensch. in Hermannstadt, xxxvi, p. 82.

1897. Vespertilio murinus Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,
p. 382, October, 1897.

1910. Eptesicus siculus and Vespertilio viurinus Trouessart, Paune Mamm.
d'Europe, 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 li times height from crown; anterior basal lobe so much
reduced as to suggest the keel of the Molossidse, the anterior
border of conch nearly straight from forehead to broadly rounded
tip ; posterior border 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 EpAesicus nilssonii, but light tips to
hairs very pale, almost whitish cream-bufi", producing a decidedly
" frosted " appearance ; underparts cream-bufi", 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 nilssonii with the broad, flat rostrum of E. serotinus, but
diflfers conspicuously in aspect from both in the very deep,
Nyctaliis-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 bullse 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
xr , ,,v „\JJ,. .,-.,„ o to front of inner incisor ; anteorbital foramen

Vesperttlio murinus. \

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 sj^ine
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.

TeetJi. — 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 Epdesicus ; lower canine
with same peculiarity in outline of crown though to a less degree.
Upper premolar with longitudinal diameter of crown relatively

VESPERTILIO

241

242 CHIROPTERA

less than in the European species of Eptesiciis, 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^ and m"^ 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 ; forearmi, 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.

Denmaek: 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 alhigularis Peters (Berlin).

Remarks. — Superficially this bat somewhat resembles Eptesiciis
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.

9. 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 fiir Natur-

gesch., 1839, i, p. 312 (part).
1842. Nocttilinia Gray, Ann. and Mag. Nat. Hist., x, p. 258 (part ; contained

piroterus = 7ioctitla a,nd fnlviis = Scotophilus kulilii).

1856. Panugo Kolenati, AUgem. deutsche Naturhist. Zeitung, Dresden,.

neue Folge, ii, p. 131 {noctula and leisleri).

1857. Vesperugo Blasius, Saugethiere Deutschlands, p. 49 (part).

NYCTALUS 243

1878. Vesperiigo Dobson, Catal. Chiropt. Brit. Mus., p. 183 (part).

1893. Noctulinia H. Allen, Proc. U.S. National Lluseum, xvi, p. 30,

June 13, 1893.
1897. Pterygistes Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384,

October, 1897.
1899. Euvcsperugo Acloque, Fauna de France, Mammif^res, p. 32 (part,

included noctula, leisleri, maurus, kuhlii, 'pipistrellus, and

abrcmnus).

1907. Pterygistes Miller, Families and Genera of Bats, p. 207, June 27,

1907.

1908. Ni/ctalus Andersen, Ann. and Mag. Nat. Hist., 9th ser., i, p. 434,

'May, 1908.

Type sjjecies. — 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 b c d

Fig. 43.
A'l/ctalus maximus (a), N. noctula (b), N. leisleri (c), and X. azoreum (d). Jfat. size.

striction, and with large anterior palatal emargination ; fifth finger
much shortened, scarcely exceeding metacarpal of fourth or third.
Bemarhs. — This genus is well difierentiated from allied groups
by the peculiar narrowing of the wing due to the shortness of the
fifth finger. Three of the half dozen known species occur on the
continent of Europe, while a fourth inhabits the Azores. - The
most obvious character by which the European forms are dis-
tinguished is the size of the skull (see fig. 43).

KEY TO THE EUROPEAN SPECIES OP NYCTALUS.

Condylobasal length of skull more than 17 mm. ; fore-
arm more than 45 mm. ; hairs of back without
dark bases.
Condylobasal length of skull 22 to 23 mm. ; forearm

64 to 68 mm. (Southern Europe) N. maximus, ^p. 244.

Condylobasal length of skull 17 • 4 to 19 • 4 mm. ; fore-
arm 47 to 55 mm. (Distribution general) N. noctula, p. 245.

Condylobasal length of skull less than 17 mm. ; fore-
arm less than 45 mm. ; hairs of back with notice-
able dark bases.
Condylobasal length of skull 15 to 16 mm. ; forearm

39 to 43 mm. (Distribution general) N. leisleri, p. 252.

Condylobasal length of skull 13 to 14-2 mm. ; fore-
arm 35 to 42 mm. (Azores) N. azoreum, p. 254.

R 2

244

CHIROPTERA

NYCTALUS MAXIMUS Fatio.

1781. ? Vespertilio lasiopterus Schreber, Saugethiere, j)!. lvii b. See

Thomas, Ann. and Mag. Nat Hist., 8th ser., viii, pp. 379-380,

September, 1911.
1827. ?■? Vespertilio ferruginciis Brehm, Ornis, Heft iii, p. 26 (Renthendorf,

Thiiringen, Germany).
1869. [Vesperugo nochda] var. maxima Fatio, Faune Vert. Suisse, i, p. 57.

Co-type in Geneva Museum.
1900. Ptcrygistes maximns Miller, Proc. Biol. Soc. Washington, xiii,

p. 156, June 13, 1900.
1910. Nyctalus mMximus Trouessart, Faune I\Iamm. d'Europe, p. 19.

Tii/pe 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 iV^. noctula.

Far 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.

Shull 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 fingcir, 76 and 77 ; ear from meatus,
21 and 22 ; width of ear, 23 and 24. Adult male from Venice : *
head and body, 92 ; tail, 65 ; tibia, 26 ; foot, 14 ; forearm, 67.
Adult female from Amsteg, Uri, Switzerland : head and body, 90± ;
tibia, 21-6 ; foot, 13 ; forearm, 64; thumb, 9*4; third finger,
114; fifth finger, 72. For cranial measurements see Table, p. 250.

Specimens examined. — Five, from the following localities : —
Switzerland : Amsteg, Uri, 1 (Geneva ; co-type).
Italy: Pisa, 8 (B.M. and U.S.N.M.); Ravenna, 1.

* Ninni, Atti Soc. Ital. Sci. Nat., Milano, xxvi, p. 109, 1883.

NYCTALUS 245

Hemarks. — Nyctalus 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 (e). 94. 6. 3. 3.

6 al. Ravenna. Florence Museum (e). 85. 7. 6. 9.

6 al. Lidth de Jeude Coll. 67. 4. 12. 337.

NYCTALUS NOCTULA 8chreber.

1774. Vespertilio noctula Schreber, Saugthiere, i, pi. lii ; description, i,

p. 166, under name : Die Speckmaus (France ; based primarily

on "La Noctule " of Daubenton, Hist. Acad. Royale des Sci.,

Paris, 1759, p. 376. 1765).
1776. Vespertilio larclarius P. L. S. Miiller, Natursyst. Suppl. u. Regist.-

Band, p. 15 (France; based primarily on Schreber's Speckmaus).
1789. \^Vespertilio'] magnus Borkenhaut, Syn. Nat. Hist. Great Britain and

Ireland, i, p. 1 (Cambridge, England ; based on Pennant, Brit.

ZooL, No. 38).
1789. Tes2)ertilio altivolans White, Nat. Hist, and Antiquities of Selborne,

p. 93 (Selborne, Hampshire, England).
1816. ? Vespertilio major Leach, Syst. Catal. Spec. Indig. Mamm. and

Birds Brit. Mus. (Willoughby Society reprint, 1882), p. 5 Nomen

nudum : " Great Bat."
1818. Vespertilio proterus Kuhl, Ann. Wetterau, Gesellsch. Naturk, iv

(= Neue Ann., i), pt. 1, p. 41 (Substitute for noctula).
1829. Vespertilio 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[ertilio'] palustris Crespon, Fauna Meridionale, i, p. 22 (marshes

near Nimes, Gard, France).
1857. Vesperugo noctula Blasius, Saugethiere Deutschlands, p. 53.
1869. [Vesjjerugo noctula''] var. minima Fatio, Faune Vert. Suisse, i, p. 58

(Geneva, Switzerland).
1878. Vesperugo noctula Dobson, Catal. Chiropt. Brit. Mus., p. 212.
1897. Pterygistes 7ioctula Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,

p. 384, October, 1897.
1910. Nyctalus noctula Trouessart, Faune Mamm. d'Europe, p. 18.

Tiij)e 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, extendincf when laid forward about half

246

CHIROPTEKA

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 con^h finely papillose, without evident cross ridges ;
tragus very short, scarcely rising above level of anterior base of
conch, much wider above than below, its greatest width about
equal to height ; anterior and posterior borders concave, the
anterior more so than posterior, their length about equal to that
of expanded upper portion, which is more abruptly rounded
anteriorly than posteriorly. Wing long and slender, the fifth
finger exceeding foi'earm by only -f to ^ length of latter, the
membrane leathery and opaque, joining leg at ankle ; third and
fourth metacarpals sub-equal, the third slightly the longer and
about equal to forearm, fifth abruptly shorter by slightly more
than one-fifth forearm. Leg short and strong, the broad foot
about one-half as long as tibia ; calcar 1^ to twice as long as
tibia or as free border of interfemoral membrane, its distal
termination obscure but basal portion strong, well defined and
with keel about 2 • 5 mm. wide supported by a well defined
thickening and terminating in an abrupt convexity on side
nearest heel. Tail rather short, extending to between shoulders
when laid forward, the tip of last vertebra (about 2 mm.) free.

Ftir and colour. — Fur dense and velvety, the hairs at middle
of back only about o 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
Ridgvvay, the hairs showing a faint clouding of prouts-brown in
certain lights, their basal portion a light dull isabella-colour ;
underparts scarcely difterent 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.

247

Nyctalus noetula.
Nat. size.

the region on each side of it flat or sUghtly depressed ; lambdoid.
crest high, curved abruptly forward at point of junction with
sagittal crest ; floor of brain-case with small but evident lateral
pits, well defined anteriorly but communicating posteriorly with
vacuity between cochlea and basioccipital ; auditory bullae well
developed but of moderate size, the transverse diameter about
equal to distance between bulk^. 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 nai'ial 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
edge of which is on level with middle of
large premolar, rather evidently concave

laterally, less so longitudinally. Posterior extension of palate
nearly parallel sided, its width at level of posterior edge of third
molar slightly less than length ; hamulars barely turned inward ;
median sjjine large, acute. Mandible robust, noticeably deeper
at symphysis than immediately behind tooth-row ; posterior
jiortion 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-.'-ondary
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

CHIROPTERA

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 /j and i.^ compressed, tritid,
that of «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 reniform in
outline, its cusp low but distinct ; posterior upper premolar with
crown area somewhat more than half that of first molar, the
protocone well developed, nearly as high as in molars, the main
cusp rather high but much shorter than canine (measured along
cingulum), its two cutting edges well developed. Lower pre-
molars closely crowded, their crowns about equal in cross-section
and somewhat more than half that of canine, the shaft of fii-st
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 VV-pattern normal, m'^ with
crown area distinctly more than half that of m^, its longitudina.I
diameter through metacone about half ti'ansverse 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 ^^^fl '"2 ^^"^
approximate equality in m^ ; 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, IS; foot, 10;
forearm, 52 ; thumb, 9 ; third finger, 94 ; fifth finger, 58 ; ear
from meatus, 15 ; width of ear, 16. Adult female from Herrn-
hut, Saxony, Germany : head and body, 79 ; tail, 53 ; tibia, 19 ;
foot, 10-6; forearm, 53; thumb, 9*4; third finger, 97; fifth
finger, 60 ; ear from meatus, 17 ; width of ear, 17. Two adult
males from Pisa, Italy : head and body, 70 and 71 ; tail, 49 and
52-6 ; tibia, 18 and 19 ; foot, 10 and 11 ; forearm, 51 and 53 ;

NYCTALUS 249

thumb, 8 and 9 ; third finger, 92 and 97 ; fifth finger, 56 and 59 ;
ear from meatus, 14-6 and 16 ; width of ear, 14 and 15' 6. Two
adult females from the same locaHty : head and body, 74 and 75 ;
tail, 50 and 53 ; tibia, 18 '6 and 18 ; foot, 10 and 11 ; forearm,
51 -6 and 52 ; thumb, 9 and 10 ; third finger, 93 and 97 ; fifth
finger, 57 and 58; ear from meatus, 16 and 16; width of ear,
16 and 16. Two adult females from Corinth, Greece : head and
body, 69 and 76; tail, 51 and 54; tibia, 18 and 19; foot, 11
and 10 ; forearm, 52 and 54 ; thumb, 9 • 4 and 9 • 4 ; third finger,
96 and 102 ; fifth finger, 58 and 62 ; ear from meatus, 15*6 and
16 ; width of ear, 15 and 16. For cranial measurements see
Table, p. 250.

Specimens cxaviincd. — 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; Triug, 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.

AusTRiA-HuNGAEY : Csallokoz-Somorja, Pressburg, Hungary, 2.

Roumania: Gageni, 1; Bustenari, 1; no exact locality, 1 (U.S.N.M.).

Switzerland : Geneva, 4 (Geneva and Mottaz) ; St. Margarethen,
Appenzell, 8 (B.M. and U.S.N.M.).

Spain : Seville, 4 ; no exact locality, 2.

Italy : Pisa, 18 (U.S.N.M.) ; Florence, 4 (Mottaz) ; Vallombrosa, near
Florence, 1 (U.S.N.M.) ; Siena, 1 ; Rome, 2.

Greece : Corinth, 12 (U.S.N.M.).

BemarJcs. — 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.

6. Aberia, Merionethshire, Col. Yerbury (c & p). 11. 1. 3. 26.

Wales.
9. Bowdon, Cheshire, Eng- G. Barrett-Hamilton 11. 1. 2. 100.
land. (T. A. Cotuard.) (p).

(11. 1. 1. 126-
2 (5, 2 ?. Oundle, Northampton- Lord Lilford (p). I 127.

shire. 11. 1. 1. 129-

( 130.
9. Bedfordshire. J. S. Elliot (c & p). 11. 1. 3. 391.
1. Trumpington, Cambridge- G. Barrett-Hamilton 11. 1. 2. 101.
shire. (p).
9 al. Cambridgeshire. Rev. L. Jenyns (c & p).

250

CHIROPTERA

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252

CHIROPTERA

6. Henley-on-Thames, Ox-
fordshire.
6 st. Sandringham, Norfolk.

9. Chelmsford, Essex.

9. Wellyn, Hertfordshire.

2 c5, 9 st. Tring, Hertfordshire.

2 6. Shalford, Surrey.

3 d, 2 9. Earlsfield, Surrey.

6 st. Fulham Park, London.
6, 9. Eastwell, Kent.

9. St. Leonards, Sussex.

6. Farnborough, Hampshire.

9. Wareham, Dorset.

6. Oosterbeek, Guelderlaud,

Holland.
9. Niesky, Silesia, Germany.

{W. Baer.)
6. Schwartzburg, Thiiringen,

200 m.

3 (5. St r ass, Burgheim,

Bavaria. (Korbitz.)

1 al. Stuttgart, Wurtemburg.
2. Csallokoz-Somorja, Press-
burg, Hungary.

6. Gageni, Prahova, Rou-
mania. (IF. Dodson.)

9. Bustenari, Prahova. (T7.
Dodson.)
2(5,3 9. St. Margarethen,
Appenzell, Switzer-
land. (E.H.ZoUikofer.)

4 al. Seville, Spain. {A. Ruiz.)

2 al. Spain.
i juv. al. B.onie.

Heatley Noble (c & p).
H.M.King Edward VII.

M. Christy and L. E.

Thompson (c & p).
C. H. B. Grant (c & p).
Hon. N. C. Rothschild

(c & p).
W. R. Ogilvie - Grant

(c & p).
C. H. B. Grant (c & p).

J. Saunders (c & p).
C. H. Caton - Haigh

(c & p).
R. Butterfield (c & p).
H. S. Scott (c & p).
W. T. Blanford (c & p).
0. Thomas (c & p).

Lord Lilford (p).

Lord Lilford (p).

Lord Lilford (p).

Dr. A. Giinther (p).
Budapest Museum (e).

Lord Lilford (p).

Lord Lilford (p).

0. Thomas (p).

Seville Museum (e).
Lord Lilford (p).
Florence Museum (e).

11.

1.

3.

40.

96.

7.

27

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11.

1.

3.

41.

11.

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3.

33.

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22.

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55.

3.

34-

11.

1.

3.

28-

32.

87.

7.

2S

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11.

1.
37.

3.

36-

11.

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3.

392.

11.

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3.

39.

11.

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3.

38.

98.

2.

1.

5.

99.

1.

9.

7.

95.

4.

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11.

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30-

32.

66.

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6.

94.

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5.

8.

1-4.

72.

8.

21

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7.

6.

8.

NYCTALUS LEISLERI Kulll.

1818. Vespertilio leisleri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, ]). 46.
1818. Vespertilio dasycarpos Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 49 (Alternative name for leisleri).
1839. Vespertilio pachygyiathtis Michahelles in Wagner, Schreber's Saug-

thiere, SuppL, i, pi. lv b (Dalmatia. See Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii,

Abth. I, p. 222, 1870).
1857. Vesperugo leisleri Blasius, Saugethiere Deutschlands, p. 56.
1878. Vesperugo leisleri Dobson, Catal. Chiropt. Brit. Mus., p. 215.
1910. Nyctalus leisleri Trouessart, Faune 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.
nodula, 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. Eai-s
and membranes (dry) blackish. Two skins are a pale buffy wood-
brown throughout except for the usual dark bases of the hairs.

Skull and teeth. — Except for its smaller size and more delicate
structure the skull does not differ appreciably from that of
Nyctalus nodula. Teeth essentially as in the larger species, but
crown area of upper incisors nearly equal, and lower incisor row
forming a deeper, almost V-s^aped 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 linger, 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 : —
Ireland : Belfast, 1 ; Bray, Co. Wicklow, 2 ; Co. Armagh, 2.
England: Mexbro', Yorkshire, 1; Welford, Warwickshire, 2; Cleave
Priory, 1 ; no exact locality, 1.

RouMANiA : Bustenari, Prahova, 1.
Switzerland : Meiringen, Bern, 1.

Bemarks. — Though its smaller size is the most obvious
character of this species as compared with N. nodula, 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.
<5al.
9al.
9al.
2 9.

Belfast, Ireland.

Bray, Wicklow.

Armagh.

Armagh.

Welford, Warwickshire,
England.

Cleeve Priory, Warwick-
shire.

England.

Bustenari, Prahova, 840 m.
Roumania. (W.Dodson.)

Meiringen, Bern, Switzer-
land.

G. Barrett-Hamilton (p.)
J. E. Harting (c & p).
Dr. G. E. Dobson (c & p).
R. M. Barrington (c & p).
Tomes Collection.

Tomes Collection.

Dr. Leach (p).
Lord Lilford (p).

Tomes Collection,

1. 3. 15. 1.
90. 2. 14. 1.
89. 11.12.5.

74. 5. 28. 8.
7. 1. 1.386-

387.
7. 1. 1. 388.

63. A.
4. 4. 6. 6.

7. 1. 1. 889.

254

CHIROPTERA

NYCTALUS AZOREUM Thomas.

1901. Ptcrygistcs azoreuni Thomas, Ann. and Mag. Nat. Hist., 7th ser., viii,
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 ; underj^arts 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 • trans verse diameter of m^ relatively greater,
crown area of upper incisors nearly equal, and lower incisor row
forming a more abrupt convexity.

Measurements. — Type (adult male, St. Michael) : head and
body, 54 ; tail, 42 ; tibia, 17 ; foot, 7*7; forearm, 37 ; thumb, 6 ;
third finger, 62 ; fifth finger, 43 • 7 ; ear from meatus, 12. Adult
male from Terceira : head and body, 50 ; tail, 43 ; tibia, 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 : — Terceira, 4 (B.M. and U.S.N.M.) ; above Magdalena, Pico, 5 ; St.
Michael, 6 (B.M. and U.S.N.M.) ; St. George, 2.

Memarhs. — The Azorean Nyctalus is slightly more differentiated
from N. leisleri than the Continental forms are among themselves,
since it differs from the other members of the genus rather
noticeably in colour as well as in size. The specimens show
considerable variation in the extent of the light wash on upper
parts caused by the pale tips to the longer hairs, but this appears
to be independent of locality.

6 al. St. Michael, Azores. P. Du Cane Godman 65. 10. 2. 1.

(c & p). [Type of species.)
2 c5, 2 9. St. Michael. (TF. B. 0. Hon. W. Rothschild 3. 6. 5. 1-3, 7.
Grant.) (p).

255

Observations

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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Genus PLECOTUS Geoffroy.

1816. ? Macrotus Leach, Catal. Spec. Indig. Manim. and Birds Brit. Mus.
(Willoughby Society reprint, 1882), p. 1. Nomen nudum : "Euro-
pean Longear, Macrotus europxus.'"

1818. Plecotus Geoffroy, Description de I'figypte, ii, p. 112. For date see
Sherborn. Proc. Zool. Soc. Loudon, 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 : * ^, c Jfi, pm ||, m || = 36.
Skull with large, elongate and rounded brain-case, and slender
but normally formed rostrum ; auditory bullae 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.

Bemarks. — -The genus Plecotus is at once recognizable among
European bats by the very lai'ge 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 Linnpeus.

1758. Vespertilio auritus Linnaeus, Syst. Nat., i, 10th ed., p. 32 (Sweden).

1816. ? Macrotus europxus Leach, Catal. Spec. Indig. IMamm. and Birds
Brit. Mus. (Willoughby Society reprint, 1882), p. 1. Nomen
nudum: " European Longear, Macrotus europeeus.''''

1818. Plecotiis auritus Geoffrey, Description de I'Egypte, ii, p. 118.

1825. Vesperlilio otus Boie, Isis, p. 1206 (Copenhagen, Denmark).

1826. Vespertilio cornutus Faber, Isis, p. 515 (Jutland, Denmark).

1827. Plecotits communis Lesson, Man. de Mammal., p. 95 (France).

1828. P\lecotus'] brevimanus Jenyns, Trans. Linn. Soc, London, xvi, p. 55

(Grunty Fen, Isle of Ely, Cambridgeshire, England).

1829. Plecotus vulgaris Desmarest, Faune Franpaise, p. 18 (France).

PLECOTUS 257

1829. [VcsiJertilio anritus'] y austriacus Fischer, Synops. ]Mamm., p. 117.

Based on the "second variety" of Desmarest, Diet, des Sci. Nat.,

Lviii, p. 51, 1829 (Vieuna, Austria).
1832. Plecotus vclatus I. Gooffroy, Gu^rin's Mag. de Zool., ii, 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. Gesellsch.

Natur.- u. Heilkunde, Giessen, p. 40, May, 1860 (Dillonburg,

Oberhessen, Germany).
1863. {^Plecotus auritus'] var. typus Koch, Jabrb. des Vereins fiir 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 (Wcstorvvald, Nassau,

Germany).
1863. [Plecotus auritus] var. brcvipcs Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 407 (Substitute for kirscli-

baumii Koch).
1878. Plecotus auritus Dobson, Catal. Chiropt. Brit. Mus., p. 178.
1910. Plecotus auritus Trouessart, Faune Mamm. d'Europe, p. 12.

2'ypc locality. — Sweden.

Geographical distribution. — From Ireland eastward into Asia,
and from the Mediterranean north to the Scandinavian Peninsula.

Diagnosis. — Characters as in the genus ; auditory bull*
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 tijj 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

s

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 hi 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 w^hen laid forward, the extreme tip (about
1 • 5 mm.) free.

ShiU. — 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 bulla?. Dorsal profile rising gradually from front
of nares to middle of anterior portion of brain-case, with slight
concavity at interlachrymal region, then falling gradually to
low, strongly convex, slightly overhanging lambda ; ventral
profile scarcely elevated posteriorly. Brain-case large but rather
low, the depth at middle only a little more
than half mastoid breadth, the general out-
line when viewed from above squarish with
rounded occipital projection and abrupt
anterior constriction, the surface smoothly
rounded, with slightly indicated sagittal ridge
anteriorly and very low, short lambdoid crest,
the latter situated unusually close to upper
edge of foramen magnum ; an irregular area
in supramastoid region thickly sprinkled with
small vacuities ; base of brain-case smooth,
without special features, a narrow groove at
PI • t I tus each side of basioccipital ; . auditory bulla

Nat. size. relatively larger than in any other European

bat, its diameter about three times the
space between bullae. Interorbital constriction narrow, scarcely
hoiu'-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 off ; rostral
depth at front of orbit about equal to distance from orbit to

PLECOTUS 259

alveolus of ijiner incisor ; anteorbital foramen small, close to rim
of orbit, its oritice over parastyle of first molar ; lachrymal fora-
men slightly above and behind it, on inner edge of orbit. Palate
rather broad and short, sti'ongly 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^ 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 of canine
in basal area, its shaft with evident posterior cutting edge ; large
premolar with crown area about two-thirds that of first molar,
the inner side very narrow and without crushing surface, the
antero-internal cusp slender but well developed. Crown area of
anterior lower premolar 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 aci'oss 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'^

s 2

260

CHIROl'TKRA

does not extend outward to level of parastyle and meiasiyle ;
W-pattern normal ; m^^ with three cusps and tliree conmiissures,
its crown area about half that of m'^, its transverse diameter
through metacone about half length of anterior border. Lower
molars with no special peculiarities except that protoconids and
inner cusps are unusually high and slender ; ciugulum forming a
barely indicated postero-internal cusp behind entoconid ; second
triangle of 7/(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, 17 • 6. Adult female
from i'lorence, Italy : head and body, 49 ; tail, 48 ; tibia, 19'2 ;
foot, 9; forearm, 39-4; thumb, 7*8; third finger, 69; fifth
finger, 54 ; ear from meatus, 34 ; width of ear, 23*4 ; tragus, 17.
For cranial measurements see Table, p. 262.

Specimens examined. — Eighty, from the following localities : — ■

Scotland : Montrose, Forfar, 4.

Ireland : Antrim, 1 ; Co. Longford, 2.

England: Alderley, Cheshire, 1 ; Bowdon, Cheshire, 1; Diss, Norfolk, 1;
Winfarthing, Norfolk, 1 ; Woburn Sands, Bedfordshire, 1 ; Pembrokeshire,
no exact locality, 1 ; Chelmsford, Essex, 1 (U.S.N.M.) ; Tring, Plertford-
shire, 1 ; Boxmore, Hertfordshire, 1 ; Hoddesdon, Hertfordshire, 1 ;
London, 4 ; Godalming, Surrey, 1 ; Surrey, no exact locality, 1 ;
Bonchurch, Isle of Wight, 1; near Houiton, Devonshire, 2 ; Devonshire,
no exact locality, 8 ; no exact locality, 1.

Holland : Oosterbeek, Guelderlaud, 4.

France : Near Barcelonnette, Basses-Alpes, 1 (Mottaz).

Germany : Wernigerode, 3 ; Ummerstadt, Thiiringen, 2 ; Magdeburg, 2 ;
Niesky, Silesia, 4 ; Damsdorf, Silesia, 1.

Austria-Hungary : Csallokoz-Somorja, Pressburg, 7.

Switzerland : Geneva, 2 (Mottaz) ; Lausanne, Vaud, 1 (IMottaz) ;
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 ;
Rome, 1; no exact locality, 1; Sicily, 1.

Spain : Silos, Burgos, 2 (U.S.N.M.) ; La Granja, Segovia, 1.

Remarlcs. — Plecotus auritus is so readily distinguished from all
other European bats by the great size of the ears that no special
comparisons are required.

(?, 8 9. Montrose, Scotland. J. H. Coward (c & p). 11. 1. 3. 48-51
6 st. Antrim, Ireland. Hon. N. C. Rothschild 1. 9. 3. 5.

PLECOTUS 261

2 6 al. Longford. Dr. G. E. Dobson (c & p). 76. 2. 12. 2.

6. Pembroke, Wales. W. E. de Winton (c & p). 11. 1. 3. 393.

1 St. Winfarthing, Norfolk, E. Markham (c & p). 9G. 7. 27. 3.
England.
6. Diss, Norfolk. {W. E. C. H. B. Grant (p). 11.1.3.394.

Slierrin.)
1 al. Woburn Sands, Bed- A. Death (c & p). 7. 9. 2. 1.

fordsliire.
?. Boxmore, Hertford- H. Piffard (c & p). 11. 1. 3. 55.

shire.
1 St. Hoddesdon, Hertford- A. Chittenden (c & p). 86. 9. 27. 1.
shire.
6. Tring, Hertfordshire. Dr. E. Hartert (c & p). 11. 1. 3. 56.

9 al. London. No history.

6. London. Sergt. Brown (c & p). 11. 1. 3. 57.

6. S. Kensington, Middle- • 11.1.3.54.

sex.
6. Godalming, Surrey. W. T. Blanford (c & p). 11. 1. 3. 58.

1. Bonchurch, Isle of Rev. C. A.Bury (c & p). 4}. 3. 29. 6.

Wight.
6,9. Honiton, Devonshire, G. C. Shortridge (c &p). 11.1.3.52-53.
300 ft.
4 (5, 4 ? al. Devonshire. No history.

2 6,2 9. Oosterbeek, Guelder- 0. Thomas (c & p). 98. 2. 1. 1-4.

land, Holland.
?. Wernigerode, Harz, W. Wolterstorff (c & p). 0.2.8.1.

Germany.
6, 9. Wernigerode, Harz. Lord Lilford (p). 11. 1. 1. 45-46.

{Wolterstorff.)
6. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 47.

( Wolterstorff.)
6,9. Ummerstadt', Thiirin- Lord Lilford (p). 11.1.1.43-44.

gen. (Scliuchardt.)

6. Niesky, Silesia. {W. Lord Lilford (p). 99. 1. 9. 1.

Baer.)
6 al. Damsdorf, Silesia. Dr. A. Giinther (p). 06. 8. 1. 10.

7. Csall okoz-Somorja, Budapest Museum (e). 94.3.1.5-11.

Pressburg, Austria-
Hungary.
6. Thurgau, St. Gallen, 0. Thomas (p). 4. 4. 5. 14.

Switzerland. (Zolli-
kofer.)
1. Switzerland. Leon 0. Galliard (p). 75. 9. 20. 2.

? al. Rimini, Italy. Florence Museum (e). 85. 7. 6. 2.

6. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 15.

Skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 728.

(without parte.)
skull).

9 al. Sicily. Purchased (Parzudaki). 52. 2. 26. 18.

1 al. La Granja, Segovia, M. de la Escalera (c). 8. 7. 30. 5.
Spain.
1. Europe. ■ Leyden Museum (b). 37. 4. 28. 25.

262

CHIEOPTERA

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BARBASTELLA

263

Genus BARBASTELLA Gray.

1821. Barbastella Gray, London Med. Repos., xv, p. 300, April 1, 1821.

1825. Barbastdlus Gray, Zool. Journ., ii, p. 243, July, 1825.

1839. Synotus Keyserling and Blasius, Wiegmaun's Archiv fiir Naturgescb.,

1839, I, p. 305.
1857. Synotus Blasius, Siiugethiere Deutschlands, p. 42.
1878. Synotus Dobson, Catal. Cbiropt. Brit. Mus., p. 175.
1897. Barbastella Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 375,

October, 1897.
1907. Barbastella Miller, Families and Genera of Bats, p. 223, Juno 29,

1907.

Type species. — Vespertilio harhastellus Schreber.

Geographical distribution. — Northern Africa ; central and
southern Europe ; west-central Asia to the Himalayas.

Characters. — Dental formula : i |-^, c ^f^J, pm ||, m ^^ = 34.
Skull with rather long, rounded brain-case and weak rostrum, the
upper surface of which is occupied by a shallow, flattened-concave
area extending from nares to faintly developed supraorbital
ridges ; auditory bullae 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.

Bemarks. — The short broad ears joined together across fore-
head distinguish this genus from the other European members of
the family Vespertilionidse. Two species are known, one peculiar
to tlie Himalayan region, the other occurring in Europe.

BARBASTELLA BARBASTELLUS Schreber.

1774. Vespcrtilio barbastclhis Scbreber, Siiugtbiere, i, pi. LV (description, i,

p. 168, under name: Das Kurzmaul). Based primarily on "La

Barbastelle " of Daubenton, Hist. Acad. Royale des Sci., Paris,

1759, p. 377. 1765.
177G. Yespertilio barbastelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.-

Band, p. 17 (Burgundy).
1836. Barbastellit's daubentonii Bell, Hist. Brit. Quadr., pt. i, p. 63

(Burgundy; based primarily on Daubenton).
1838. Barbastelliis communis Gray, Mag. Zool. and Bot., ii, p. 495,

February, 1838 (Renanring of barbastellus).
1857. Synotus barbastellus Blasius, Siiugethiere 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, Fauue Mamm. d'Europe, p. 11.

Type locality. — Burgundy, France.

Geographical distribution. — Central and soiitliei-n Europe,

264

CHIROPTEUA

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. longT)y 1 mm. wide, its upper
border convex, its lower border concave) ; lower border of ear full
and rounded, but with no diflerentiated 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 excej^t at extreme tip and along posterior border
to a httle 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 lol)e 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 bulhie scarcely more in-
flated than in the small sj^ecies 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- „

orbital foramen relatively large, over anterior Barbastaia harha-
margin of m^. Rostrum short, its upper sur- steiius. Nat. size.
face flattened-concave, deeply emarginate in
front by the unusual backward extension of the uppei- margin of
anterior nares, the posterior border of which is at level of ante-
orbital foramina, exposing anterior extremity of vomer. Palate
short, noticeably concave both longitudinally and laterally ; its
anterior emargination small, extending slightly behind level of
middle of canine ; its posterior border slightly behind level of m^,
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 (J-'^liaped 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 Plecoius
auriius, 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 : bead
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, Gei'many : 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). Eor 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 ; Kugby, Warwick-
shire, 1; Welford, Rugby, 1; Pilton, Onndle, Northamptonshire, 1; Epping,
Essex, 1 ; Llanelwedd, Radnorshire, 1 ; Swindon, Wiltshire, 1 ; Milton,
Hampshire, 1.

Prance : 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).

Hemarks. — 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. Croicfoot.)

6. Arrow Lodge, Warwickshire. Tomes Collection. 7.1.1.348.

1. Alcester, Warwickshire. Tomes Collection. 7. 1. 1. 348.

6 St. Rugby, Warwickshire. G. Dalgleish (c & p). 4. 10. 13. 2.

6. Oundle, Northamptonshire. Lord Lilford (c & p). 94. 4. 16. 1.

9 al. Epping, Essex. H. Doubleday (c & p). 44. 5. 1. 1.

6. Milton, Hampshire. Rev. J. E. Kelsall (p). 11. 1. 3. 59.

BARBASTELLA

267

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268

CHIEOPTERA

i. Swindon, Wiltshire. Eev. E. A. Goddard 8. 5. 12. 1.

•(c &p).
6. Caterille, Haute-Garonne, O. Thomas (p). G. 4. 1. 7.

1000 m. France.

(A. Robert.)
i, ?. Niesky, Silesia, 181 m. Ger- Dr. E. Hamilton (p). 97. 12. 4. 2-3.

many. (W. Baer.)
2 9, Niesky, Silesia. {W. Bacr.) Lord Lilford (p). 99. 1. 9. 2, 4.

S. W e r n i ger od e, Saxony. Lord Lilford (p). 0.2.8.2.

{Walter storff.)

Sub-Family MINIOPTERINiE.

1878. Miniopteri Dobson, Catal. Chiropt. Brit. Mus., p. 170 (part).
1907. Minioptermx Miller, Families and Genera of Bats, p. 227, June 29,
1907.

Geographical distrihution. — Africa, southern Europe and
southern Asia, eastward to the Malay region, Japan and
AustraHa.

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.

HemarJcs. — The sub-family Miniopterinse , though widely dis-
tributed in the warmer portions of the Old World, is at present
known to contain the genus Mmiopterus only.

Genus MINIOPTERUS Bonaparte.

1837. Miniopterus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx, under

Vespertilio emarginatus (Sub-genus of Ves2:)crtilio).
1857. Miniopterus Blasius, Saugethiere Deutschlands, p. 45.
1866. Miniopteris Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 91,

February, 1866.
1878. Miniopteriis Dobson, Catal. Chiropt. Brit. ]Mus., p. 347.
1892. Minyopterus Winge, Jordfundne 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. schreihersii
Kuhl.

Geographical distrihution. — Same as that of the sub-family.

Characters. — Dental formula : i 'j0,, c ^^, pm ||, m [^j = 34.
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.

Bemarhs. — The peculiar shortening of the ears together with
the remarkable elongation of second phalanx of third finger leadily

WINIOPTERUS

269

disiin,i,'uis]i this genus aiiiong the members of the European fauna.
About a dozen forms have been described, one of which occurs
in southern Europe.

MINIOPTERUS SCHREIBERSII Kuhl.

1819. Vespc7-tilio schrcihersii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 185 (Hungary).

1837. Vespertilio wsinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Monte Corno, Ascoli, Italy). Type in British Museum.

1841. Vespertilio orsinii Temniinck, Monogr. de Mamm., ii, p. 179 (modi-
fication of ursinii).

1857. Minioptcrus schreibersii Blasius, Saugethiere Deutschlands, p. 46.

1878. Miniopte7-its schrcihersii Dobson, Catal. Chiropt. Brit. Mus., p. 348.

1910. Minioptcrus schrcibersi Trouessart, Paune Mamm. d'Europe, p. 34.

Type locniiti/. — Kulmbazer Cave, mountains of southern
Bannat, Hungary.

Geographical distribuiiun. — iiouthern 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
contiruious 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 CHIROPTERA

metacarpals sub-equal, about 3 mm. shorter than forearm ; iifth
metacarpal about 4 mm. shorter than third ; membrane joinint;'
leg at or a little above ankle. Leg rather slender, the foot
scai'cely half as long as tibia ; calcar about as long as tibia and
nearly equal to free border of uropatagium, its distal termination
very obscure, its posterior border without trace of keel. Tail
about as long as head and bod}', included in the membrane to
extreme tip.

Fur and colour. — The fur is of a 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 ;
(h) at middle (tips of shorter hairs, 2 mm.), ecru-drab ; and (c) at
tip (extremities of longer hairs, 2 mm.), drab. Muzzle, ears and
membranes brown, scarcely darker than body.

Skull. — The skull differs from that of all other European bats
in the great inflation of the anterior poi'tion 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
■L^!}-^^>^ 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 sufliciently to
conceal condyles when skull is viewed from
directly above ; ventral profile nearly flat,
Via. 47. slightly elevated posteriorly. Brain-case

Miniopterus schreibersii. in i- j.i i.T i i i. i

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 bi'eadth 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 bullte.
Interorbital region moderately constricted, slightly hour-glass
shaped, the lachrymal region scarcely more than half as wide
as brain-case, smoothly rounded at sides ; rostrum tapering
gradually, rounded off at sides, flattened concave along median
lihe, 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 tootli-i'ow, 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 postero-internal 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 ; «\ and i.^ with crown very low, the cutting edge
obscurely tritid, the crown of each tooth wider posteriorly (in line
of tooth-row) than anteriorly, but this more evident in second
than in first ; i\ 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 antei'o-internal cusp. Anterior upper premolar
with crown area about equal to that of canine, its general form

WU CITIHOI'TERA

essentially as in large premolar except that the antero-iuternal
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 oft', 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 v/ell 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,
W-p^t.tern 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^ 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.

MINIOPTERUS

273

Spechnens examined. — Three hundred and fifty-oue, 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.

Prance : Troubate, Hautes-Pyr6n6es, 2 ; Dions, Gard, 9 ; Marseilles, 1
(U.S.N.M.).

Switzerland: Geneva, 235 (Mottaz) ; Neuchatel, 2 (U.S.N.M.).

Austria-Hungary : Ofener Mountains, Hungarv, 1 ; Hungary, no exact
locality, 3 (U.S.N.M.).

Montenegro : Beri, 1 ; Velgi, Czolo, 1.

Italy : Western Liguria, 14 (Genoa) ; Pinalborgo, 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 Corno, Ascoli, 1
(type of ursinii Bonaparte) ; Rome, 10 ; Velletri, Rome, 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.

BemarJcs. — The lengthened second phalanx of the third finger,
the peculiar cropped a2:)pearance of the ears, and the short, dense,
velvety fur of the head are the most obvious external character-
istics of Miniojjterus scJireihersii as compared with other European
bats.

2.

6,9.

3 6.

6,<i.

8 6,19

1 al.

?.

9.

Silos, Burgos, Spain. Rev. S. Gonzalez (c).
Inca, Majorca, Balearic 0. Thomas and R. I.

Islands. Pocock (c & p).

San Cristobal, Minorca. O. Thomas and R. I.

Pocock (c & p).
Troubate, Hautes- 0. Thomas (p).

Pyr^n^es, 400 m.

France. {A. Robert.)
Dions, Gard, 70 m. 0. Thomas (p).

Franco. (C. Mottaz.)
Ofener Mountains, Budapest Museum (e).

Budapest.

Beri, Montenegro.
{L. Filhrer.)
Velgi, Czolo.

(L. Filhrer.)
1 6,2,9. Rome. (C. Goli.)
Skeleton Monte Corno, Ascoli,
(without Italy,
skull).
2 6. Marsala, Sicily.

(^1. Robert.)
6. Labyrinth, 200 ft.

Crete.

88. 1. 5. 46-47.
0. 7. 1. 4-5.

0. 7. 1. 32^34.

6. 4. 1. 8-9.

8. 8. 10. 1-9.
94. 7. 18. 10.
5. 8. 4. G.
5. 8. 4. 5.

11. 1. 2. 5-14.

7. 1. 1. 735.
{Type of Vespertilio ursmii Bonaparte.)

0. Thomas (p). G. 8. 4. 24-25.

Miss D. Bate (c). 5. 12. 2. 9.

O. Thomas (p).

0. Thomas (p).

Lord Lilford (p).
Tomes Collection.

274

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276 CHIROPTERA

Family MOL088ID^.

1865. Molossi Peters, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 258.
1872. Molossidx Gill, Arrangement of the Families of Mammals, p. 17.
1878. Emhallonuridm (part ; Molossime, part, Molossi) Dobson, Catal.

Chiropt. Brit. Mus., p. 402.
1907. Molossiclx Miller, Families and Genera of Bats, p. 241, June 29, 1907.

Geogrnphical distribution.' — Warmer parts of botli liemisplieres ;
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.

Remarks. — 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 Geofiroy, Descr. de I'Egypte, ii, p. 114 (asgyptiaciis) .
1821. Nyctinoma Bowdich, Anal. Nat. Class. IMamm., p. 28 (Modification
of Nyctinomus).

1821. Nyctinomes Gray, London Med. Repos., xv, p. 299, April 1, 1821

(Modification of Nyctinomus).

1822. Nyctinomia Fleming, Philos. of Zool., ii, p. 178 (Modification of

Nyctinomus).
1842. Mojjs Lesson, Nouv. Tabl. Regne Anim., p. 18 {Mo2}s indicus Lesson

= Dysopes mops F. Cuvier).
1878. Nyctinomus Dobson, Catal. Chiropt. Brit. Mus., p. 420 (part).
1902. Nyctmomops Miller, Proc. Acad. Nat. Sci. Philadelphia, p. 393,

September 12, 1902 [fenwrosaccus) .
1907. Nyctinomus Miller, Families and Genera of Bats, p. 251, June 29, 1907.

T?/j?e 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 ]^ or y,* c ^ , pm ?^, m i* | = 30
or 32 ; bi'ain-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.

Memarhs. — As thus defined the genus Nyctinomus contains

* In the species occurring in Europe.

NYCTINOMQS

277

about forty species, two-thirds of which are pecuHar to the Old
World, one of them occurring in the Mediterranean region of
Europe.

NYCTINOMUS TENIOTIS Ralinesque.

1814. Ccplialutcs tcniotis Rafinesque, Prec.desDecouv. SomioL, p. 12 (Sicily).
1825. Diyiops ccstoni Savi, N. Gioru. de' Letterati, Pisa, x, p. 235 (Pisa, Italy).
1840. Dysopcs savii Schinz, Europ. Fauna, i, p. 5 (Substitute for cestoni).
1871. [Dysopcs cestonii] var. nigrogriseus Schneider, Neue Denkschr.

Schweiz. Gesellsch. Naturwiss., xxiv, p. 5 (articles separately

paged). Basel, Switzerland.
1877. Nyctinonms cestonii Dobson, Catal. Chiropt. Brit. jMus., p. 423.
1891. Nyctinomus txnioiis 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. Nyctino7nzcs tseniotis Trouessart, Faune Tilamm. 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 segyptiacus 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 defined, wider
than high, the sub-circular nostrils opening forward and slightly
outward at its outer margin, its surface with very fine reticula-
tions and a few coarse wrinkles, glabrous except below, where it
is sprinkled with fine hairs, this hairy area continuous with the
brush-like fringe of recurved blunt hairs along middle of iipper
lip ; upper margin of pad broadly concave at middle, convex
latei'ally 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. AVings 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 stiflfened 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.

Fihr 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. Fr-nges on feet and hairy lines on ears, drab like
body.

ShuU. — The skull is large, but rather slender, about equal to
that of Myotis myotis iia 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-intei'orbital portion, and
the conspicuously emarginate inner side of auditory bulla?. Dorsal
profile essentially straight from nares to lambda, though with a

NYCTINOMUS

279

slight convexity over anterior two-thirds of brain-case, and a
more abrupt though not very conspicuous swelhng posteriorly,
the two convexities separated by a narrow concavity ; general
direction of dorsal pi'otile 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 iaterorbital 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 leather 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^ ; 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 sepai-ated 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 »i^ ; mesop-
terygoid space large, its width anteriorly equal to that of

FIG. 48.

Nyctinomus teiiiotis.
Nat. size.

280

CHIROPTERA

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^ and m"^. 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-externally ;
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 W'cll 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.

Xi/ctinomus tcnioiia. Teeth X 5.

NYCTINOMUS

281

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282

CHIROPTERA

of lower molcirs ; large upper premolar with crown area about
two-thirds that of m'^, 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 commissui-e 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, 11-4; forearm, 61; third finger, 117; fifth
finger, 61 ; ear from meatus, 31*4; width of ear, 31. Two adult
females from Cintra, Portugal : forearm, 60 and 61 ; third finger,
110 and 115 ; fifth finger, 59 and 63. For cranial measurements
see Table, p. 281.

Speciinens examined. — Six, from the following localities : —
Portugal : Cintra, 2.

Italy: No exact locality (probably from Pisa), 2. (B.M. aud U.S.N.M.) ;
Sicily, 1 (U.S.N.M.).

Greece : No exact locality, 1.

2 ?. Cintra, Portugal. 0. Thomas (c & p). 98. 2. 2. 6-7.

i. Italy (probably Pisa). Zoological Society (p). 10. 5. 24. 1.

{Savi.)

9. Greece. Parreys. 46. 6. 15. 121.

CARNIVORA

283

Order CARNIVORA.

1827. Carnivora Gra}', 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.

Hemarhs. — 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 CARNIVORA.

Larger cheek-teeth with crowns of a crushing type, the
cusps sub-equal, low, subterete, without noticeable
cutting edges ; upper carnassial 2-rooted, in front of
anteorbital foramen, its inner lobe posterior ; size
very large; form heavy; feet plantigrade (Bears) ... Ursidx, p. 281.
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 naolar
scarcely larger than outer incisor; claws com-
pletely retractile (Cats) Fclid.r, 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) Canidx, p. .303.

Represented in Australia by a species of Canis, probably introduced.

284

CARNIVORA

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) Vlvcrrichi', p. 440.

Auditory bulla simple ; upper molars 1-1, the

crown wider internally than externally Mustelidiv, 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) Mdinx, 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) Lutrinx, 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) MusteUn.T, p. 364.

Dentition not highly trenchant ; small pre-
molars opposite, not capable of shearing
action, the points of all but jjvi^ and pm^
widely separated when jaws are closed ;
upper carnassial with posterior cusp
broad, almost fiat-topped ; auditory bulla
broader than long ; external form heavy ;
feet sub-plantigrade (Glutton) Guloninse, p. 432.

Family URSID^.

1825. Ursidn- 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

URsus '285

upper premolar and first lower molar scarcely differentiated as
carnassials, the former 2-rooted, its inner lobe at posterior border
of crown, its position so far anterior to level of anteorbital
foramen as not to be at point of greatest mechanical efficiency ;
auditory bulla flattened, without septum ; form heavy ; size
large ; feet strictly plantigrade ; digits, 5-5.

Bemarhs. — The family Ursiclse, 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-teetli that
they require no special comparisons with other carnivora. Two
genera occur in Europe.

KEY TO THE EUROPEAN GENERA OF VR8IDJE.

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 Linnseus.

1758. Urs'iis Linnseus, Syst. Nat., i, 10th ed., p. 47 [ardos, by tautonymy).
1857. Ursus Blasius, Siiugethiere Deutschlands, p. 196.
1864. Euarctos Gray, Proc, Zool. Soc. London, p. 692 (americanus).
1864. Myrmarctos Gray, Proc. Zool. Soc. London, p. 694 (cversmanni - arctos) .
1898. Ursarctos Heude, M6m. Hist. Nat. Emp. Chinois, iv, pt. i, p. 18
{yesoe7isis).

Type species. — Ursus arctos Linnteus.

Geographical distribution. — Northern hemisphere from northern
limits of the great continental areas south to the Atlas Moun-
tains, the Himalayas and Mexico.

Characters. — Dental formula : i ^'■\ c ^"\ pm '^~, m — = 42 ;

3-3' l-l' i^ 4-4' •d-:i '

inner upper incisor well developed, permanent ; first, second and
third premolars in both jaws small, single-rooted, readily deciduous,
especially pm'^, pm^^ and pin^ ; molars large and robust, the length
of the two upper teeth together equal to width of palate.

BemcirJcs. — 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 definit(?ly
known to occur in Europe.

URSUS ARCTOS Linnajus.

1758. [Ursus'] arctos Linnaeus, Syst. Nat., i, 10th ed., p. 47 (Sweden).
1772. [Ursus] ?6rsMs Boddaert, Kortbegrip van het zamenstel der Natuur,

I, p. 40 (Renaming of arctos).
1788. [Ursiis arctos] a nigcr Gmelin, Syst. Nat., i, 13th ed., p. 100

(Northern Europe).

286 CAENIVORA

1788. [Ursics arctos] $ fusctos Gmelin, Syst. Nat., i, 13th ed., p. 100 (Alps).

1778. [Ursus arctos'] y albus Gmelin, Syst. Nat., i, 13th ed., p. 100
(Unknown ; based on the " ours blauc terrestro " of Buffou).

1792. U[rsus] arctos griscus Kerr, Anim. Kiugd., p. 184 (Germany; also in
northern North America).

1797. Ursus arctos rufiis Borkhausen, Deutsche Fauna, i, p. 4G (Swiss and

Tirolean Alps).

1798. Ursus badius Schrank, Fauna Boica, i, p. 55 (Forests on the

Bohemian boundary).
1808. Ursus fusciis 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 ynajor 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] & hrunncus Billberg, Synopsis Faunaa Scandiuaviffi,

p. 15 (Northern Scandinavia).
1827. [Ursus arctos] y annulatus Billberg, Synopsis Faunae Scandinaviaj,

p. 15 (Northern Scandinavia).
1827. [Ursus arctos] 5 arcjenteus Billberg, Synopsis Faunae Scandinavise,

p. 15 (Northern Scandinavia).

1827. [Ursus] myrmcpliagus Billberg, Synopsis Faunte Scandinavia, p. 16

(Northern Scandinavia).

1828. [Ursus] formicarlus Billberg, Synopsis Faunae Scandinavios, 2nd ed.,

p. 16 (Renaming of myrmepliagus).

1829. U[rsus] pyrcnaicus Fischer, Synopsis Mamm., p. 142. Latiuization

of "Ours des Pyrenees" F. Cuvier, Hist. Nat. des Mammif., v,

fasc. 44, 1824 (Asturias, Spain).
1829. U[rsus] norvcgicus Fischer, Synopsis Mamm., p. 142. Latiuization

of "Ours de Norwege " F. Cuvier, Hist. Nat. des Mammif., ii,

fasc. 7, 1819 (Norway).
1836. ? Ursus falcigcr~R.e\ch.enha,ch., Regn. Anim. Icon., i, p. 32 ("Pyrenees" ;

afterwards supposed to be an individual of " U. fcrox.'" See Natur-

gesch. des In- und Auslands, Raubstiugeth., p. 299, 1852).
1842. Ursus pyrenxus F. Cuvier, Hist. Nat. des Mamm., Tabl. gen., p. 3

(Described in fasc. 44, 1824) (Asturias, Spain).
1847. ? Ursus euryrldmis 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, Wisseusch.-pop. Naturgesch. der

Saugeth., i, p. 372 (Norway).
1864. [Urs^ls arctos] var. 1. normalis Gray, Proc. Zool. Soc, London, p. 682

(Renaming of arctos).
1864. [Ursus arctos] sub- var. a. scanclinavicus Gray, Proc. Zool. Soc,

London, p. 682 (Based on Nilsson, Ilium. Fig. Skand. Fauna,

pi. 23).
1864. ? [Urstcs aixtos] sub- var. c. rosslcus Gray, Proc. Zool. Soc, London,

p. 682 {nomcn nudum).
1864. [Uisus arctos] sub-var. f. poloniczis Gray, Proc. Zool. Soc, London,

p. 682 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, pi. xxii,

fig. 3).
1864. [Ursus arctos] var. 2. grandis Gray, Proc. Zool. Soc, London, p. 684

("North of Europe " ; based on " a male purchased at Hull, living

in the Zoological Gardens from 1852 to 1863").

URSUS

287

1864. [Ursiis arctos] var. 4. stenorostris Gray, Proc. Zool. Soc, London,

p. G85 (Poland ; based on Cuvier, Oss. Possiles, iv, p. 332, 2od var.,

pi. XXII, fig. 4).
18G4. Mijrmarctos evcrsvianni Gray, Proc. Zool. Soc, Loudon, p. G95

(Norway).
1910. Ursus arctos, U. arctos formicarius, U. arctos alpinus, and TJ. arctos

prjrenaicus Trouessart, Panne Mamm. d'Europe, pp. G7-68.

Tyi)e 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 fi-om about 260 to 350 mm. ; interor'bital 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 butty, 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 moi-e 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

CARNIVORA

quite to heel ; region between pad and balls of toes, and at outer
side of backward extension densely furred ; fringe along edge of
foot conspicuous. Tail very short, concealed in the fur.

Colour. — The colour of body is usually a light brown or dull
buff, the head not essentially different, but feet and outer surface
of legs darker. Many individual differences in colour have been
described, some of which are probably characteristic of geographical
races.

Skull. — General form of skull rather robust, the rostrum

Fig. 50.
Ursus arctos.

moderately long (distance from orbit to front of premaxillary
contained about 2^ times in condylobasal length), the brain-case

uRsus 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

I

Fig. 51.
ITisus arctos. x J.

evenly concave throughout. Brain-case broadly ovate in outline, its
greatest breadth about equal to distance from bregma to lambda,""
its depth at middle, exclusive of sagittal crest, slightly less than
greatest breadth ; lambdoid and sagittal crests well developed in
adults, 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.

C

290

CARNIVORA

runs to extremity of each postorbital process, the hinder margin
of which it forms ; occiput wlien viewed from behind moderately
broad, the depth from himbda to lower lip of foramen magnum
contained If to 1 1 times in mastoid breadth ; mastoid processes,
paroccipital processes, and condyles extending to about the same

Fig. 52.
Ursns arctos. x h

level. Floor of brain-case nearly flat, the basioccipital with
raised edges applied to inner surface of bulla?, 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 mai'gins slightly everted ; nasal bones elongate wedge-
shaped, si^uarely truncate anteriorly, their posterior extremity on
level with or extending slightly behind nasal branch of maxillary ;
anteorbital foramen over metacone of m^ or paracone of m^ ;
palate narrow, its width between posterior molars contained about
3h times in median length ; extension behind molars nearly
parallel-sided, its length equal to about three-quarters breadth ;
mesopterygoid space 1 h times to twice as long as wide, its anterior
boi'der 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 ;
coronoid process broad, its width at level of alveolus slightly
greater than height, the anterior border at first straight then
evenly convex to overhanging tip, the posterior border concave ;
angular process short, extending slightly if at all behind level of
articular process, its inner border nearly straight, its outer border
convex.

Teeth. — The teeth are moderately large relatively to size of
skull. Upper incisors forming a continuous row, separated at
each side from canine by a diastema about as wide as inner
incisor ; i^ and P sub-equal, the former slightly the smaller, the
anterior face smoothly rounded, a little more than half as wide
as high, the posterior face abruptly concave, with shelf-like
posterior extension, the cingulum slightly developed and forming
a rudimentary nodule on inner and outer side of P and on outer
side of i^ near- level of middle of anterior surface ; P with crown
area nearly double that of P, 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 /3 ; posterior border slightly

u 2

292

CAKNIVORA

Fig. 53.
Ursus arctos. 'i'eetli iiat. size.

uRsus 293

concave, with inner, middle and outer ridges, the inner and middle
low and confluent in i^. Canines large and strong, oval or
slightly ovate in cross section, the longest diameter at level of
alveolus equal to or slightly greater than distance from alveolus
to median line of palate ; lower canine shorter and a little more
curved than upper ; a slightly 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 postero-internal 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
sube([ual 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 I'ather 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 appi'oximately 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

CARNIVOKA

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oblique to it ; postero- external cusp slightly smaller than antero-
external, and contrast betAveen outer and inner cusps less marked
than in m^ ; posterior third of crown occupied by a flattened heel,
variable in form and size, but usually narrowing off, chiefly by
slanting inward of outer boi'der, 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 ; pai'aconid forming naiTow
anterior extremity of crow^n 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-ofl^ 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.^, its surface
entirely flat except for a slightly raised rim which forms a
small antero-internal cusp.

3Ieasurements. — Adult male from Sweden (mounted) : head
and body, 1900; tail, 80; hind foot, 195 ; ear, 90. For cranial
measurements see Table, p. 294.

Specimens examined. — Eight, from the following localities : —
Sweden : Kvickjock, Norbotteu, 1 ; uo exact locality, 3 (B.IM. and
U.S.N.M.).

Switzerland : Engadine, 1.
Austria-Hungary : Near Hatszeg, Hunyad, 3.

BcmarJcs. — 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 arcfos 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. horrihilis perhaps averaging somewhat the larger. The teeth
of the two animals are also much alike.

9. Kvickjock, Norbottcn, Stockholm Museum 90. 8. 1. 3.

Sweden. (e).

2 6. Sweden. {Lloyd.) Purchased (Stevens). 62.3.29.7-8.

9 St. Engadine, Switzerland. H. Justen (p). &G. 1. 23. 1.

3 skeletons. Hatszeg, Transylvania, C. G. Danford (c). 78. 1. 16. 1-3.
Hungary.

Note on the Ursus formicarius of Bieler.

A peculiar small bear supposed to have been taken in the
Canton of Grisons, Switzerland, has been described by Professor
Bieler of the Lausanne Agricultural College as Ursus formicarius
Eversmann.* Through the author's kindness I have had the
opportunity to examine this skull. It is that of a rather young
individual, apparently a female, with basal suture open, but with
teeth showing slight indications of wear. In size it is smaller
than in a female of the same age or slightly younger from
Sweden (see Table of cranial measurements, p. 294), the inter-
orbital region is much depressed, so that the frontal profile is
nearly flat, and the mesopterygoid fossa is unusually broad and
short, the width between pterygoids equal to a little more than
half distance from hamular to level of last molar. The teeth on
the other hand are slightly larger than usual in females (see
Table, p. 295) ; 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 Myrmarctos eversmaymi. 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. Thalarctos Gray, Ann. of Philosophy, N.S., x, p. 62, July, 1825.
1825. Thalassarctos Gray, Ann. of Philosophy, N.S., x, p. 339, November,

1825.
1896. ThalassiarcJms Kohelt, Bericht Senckenbcrg. naturforsch. GescUsch.

Frankfurt am Main, p. 93 (Substitute for Thalarctos).

Type species. — Tlialarctos polaris Gray = Ursus maritimus
Phipps.

* Compte-Kendu des Stances du Sixieme Congres Internationale de
Zoologie, Berne, 1904, p. 248. 1905.

298

CARNIVOKA

Geographical disfrihution. — 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 TJialarctos, though not veiy 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, Reise durch verschiedene Provinzen des

russischen Raichs, iii, p. 691 (Arctic Ocean).
1792. Ursus polaris Shaw, Museum Leverianum, i, p. 7 (Renaming of

marinus).
1862. Thalarctos maritimus Gray, Catal. Bones Mamm. Brit. Mus., p. 105.
1908. ? Tlialassarctos jenacnsis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Jena Island,

Spitzbergen).
1908. ? Tlialassarctos sjntsberg crisis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Seven Island,

Spitzbergen).
1910. Ursus {Tlialassarctos) maritimus Trouessart, Fauue Mamm. d'Europc,

p. 66.

Tiipe locality. — Spitzbergen.

Geofjraphical 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. — Porm 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 bi'ain-case essentially as in
U. arctos, but portion at base of condyles more narrowed and

THALARCTOS

299

elongate. Palate noticeably broader than in Ursiis arctos, a
character made more conspicuous by the I'clative weakness of the
teeth. Mandible with no special peculiarities except that the

Fig. 54.
Thalarctos maritiinus. x

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

CARNIVORA

notably from that of Ursus ardos in tho reduction of tlie molars
and increased size and prehensiveness of the canines and incisors,
the details of the individual teeth present little that is specially

Via. 55.
Thalaretos mariUmua. x

noteworthy. General form of upper incisors as in Ursus arctos,
but points of i'^ and i'^ narrower and more hooked backward ; P
with cusp more slender and ridges nearly obsolete ; lower incisors

THALARCTOS

301

with lobes more sharply defined. Canines both above and below
essentially similar to those of Ursus arctos, except for their greater
size. Owing to the greater width of palate the proportion of

Fig. 56.
Thalarctos maritimus. X J.

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

CAKNIVORA

^M

Fig. 57.
Thalarctos maritinms. Teeth nat. size.

THALAECTOS 303

anterior cusp than in U. arctos, its inner side more flattened,
giving the tooth a more carnassial appearance ; jjostero-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 in^ not peculiar, though outer cusps are higher and narrower
and inner cusps relatively lower ; m'^ 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^.
1821. Canidn: Gray, London Medical Repository, xv, p. 301, April 1, 1821.

Geograpliical distribution. — Essentially cosmopolitan ; in E urope
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 etliciency ; auditory bulla moderately or considerably
inflated, without septum ; form rather light, the legs long ; size
moderate ; feet digitigrade ; toes, 5-4 or 4-4.

RemarTcs. — 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 CARNIVOEA

KEY TO THE EUROPEAN GENERA OF CANIDu^.

Interorbital regiou distinctly elevated ; postorbital processes

convex above ; pupil of eye round Cflwis, 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 Aloj^cx, p. 318.

Postorbital processes distinctly concave above ; forehead

rising gradually above level of rostrum ; ear pointed Yulpcs, j). 325.

Genus CANIS Linnaeus.

1758. Canis Linnaeus, Syst. Nat., i, 10th ed., p. 38 (type by tautonymy

C. familiaris).
181G. Lupus Oken, Lenrb. d. Naturgesch., iii, pt. 2, p. 1039 [Canis liqnis,

by tautonymy).
1837. Viilpicanis Blainville, Ann. Sci. Nat., Paris, 2nd ser., Zool., viii,

p. 279, November, 1837 (Canis aureus Linnaeus).
1839. Lyciscus H. Smith, Jardine's Naturalists' Library, Mammals, ix,

J). 160 {Canis latrans Say).
1839. Thous H. Smith, Jardine's Naturalists' Library, Mammals, ix,

p. 193 [Canis anthus F. Cuvier).
1839. Sacalius H. Smith, Jardine's Naturalists' Library, IMammals, ix,

p. 213 [Canis aureus Linnaeus).
1841. Oxygons Hodgson, Calcutta Journ. Nat. Hist., ii, p. 213 [Canis

aureus Linnaeus).
1857. Canis Blasius, Saugethiere Deutschlands, p. 177.

1868. Ncocyan Gray, Proc. Zool. Soc, London, p. 506 [Canis latrans Say).

1869. Dicha Gray, Catal. Carn. Pachyd. and Edentate Mamm. Brit. Mus.,

p. 180 [Canis antlius F. Cuvier).

Type s])ecies. — Canis familiaris Linnaaus.

Geograjjliical 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 postoi-bital
processes thick, convex above, their edges rounded off; dorsal
profile of forehead rising rather abruptly and noticeably above
level of rostrum ; dental formula : i ||, c 1^, j:>Ht t^, m || = 42 ;
teeth heavy and large, the length of carnassial and upper molars
together contained about 2| 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).

Bemarhs. — 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, tind the Old World jackals. Two species are known
to occnr in Europe, one of which is represented by several
geographical races.

KEY TO THE EUROPEAN FORMS OF 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.) ; cingulum on
outer imargiii of ?»' 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 ?«' narrow, tending to bo
incomplete at middle (Distribution general; true

Wolves) C. lupus, p. 805.

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. iui>m, p. 313.
White of throat extending uninterruptedly on to

cheek (Spain except extreme south) C. l. siijnatu^, p. 314.

CANIS LUPUS Linnaeus.
(Synonymy under subspecies.)

Geograpliical distribution. — Originally throughout Eui'ope 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 balks narrowly edged with short hair.

Colour. — General colour of upper parts, tail, and outer surface
of legs yellowish brown or bufT, darker along median region of

X

306

CARNIVORA

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.

Fig. 58.
Canis lupus, x

especially behind middle and at base of tail ; pencil narrowly
clear black, rest of tail essentially like back. Underparts and
inner surface of legs pale 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

Fig. bd.
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 2i

FIG. 60.
Canis lupus, x i.

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
1| 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
featui'es, the auditory bullae 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 postei'ior 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 ; miixillary 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 robvist, the depth at posterior edge of carnassial
contained about six times in length ; symphysis rather long ;
coronoid process high, the depth of mandible through its middle
noticeably greater than distance from last molar to back of
condyle ; angular process heavy, nearly horizontal, distinctly
raised above level of under margin of ramus.

Teeth. — The teeth are relatively larger than in any of the
races of domestic dogs, though in form they show no tangible
features by which they may be distinguished. Upper incisors form-
ing a continuous, slightly convex row, the outer tooth separated
from canine by a distinct space ; size, when viewed from in front,
increasing regularly from first to third, but third abruptly much
larger than the others in cross-section and nearly half the size of
canine ; anterior surface of i^ and i- slightly more than half as
wide as high, smoothly I'ounded oft*, 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

CARNIVORA

lobes, its general form intermediate between that of canine and
of inner incisors. Lower incisors forming continuous row between
canines, the three teeth essentially alike in form, but inci'easing

Fig. 01.
Canis lujniK. 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 l)ilobed, 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? 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 pmj subterete, that of jmi^ neai'ly 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 pn^ and pm")
slightly concave, the long axis nearly parallel with sagittal plane
except in pm^, which is obliquely set ; main cusp a little in front
of middle of crown, its height distinctly more than half length of
crown, its anterior and posterior cutting ridge well developed, the
posterior bearing a distinct secondary cusp situated over middle
of posterior root and relatively larger in lower than in upper
teeth ; a slight shelf-like projection behind secondary cusp ; pm^
similar to pm.^ 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 comjDlete 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 ofiset, 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 ecpially 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 behiiul 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 cori-esponding 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 lowei- molar single-i'ooted, 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 commissui'es, each of which joins ciugulum 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
Ih-st, 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 tlie 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 rii^ it is usually
obsolete (compare figs. 61 and 62).

Jxemarlcs. — 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 I'eason to doubt their validity.

CANIS

313

The only known characters by which the skull of Canls hqjus
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 caruassials is respectively 21*6 and 25 '0 mm.
In ten skulls with condylobasal length of more than 200 mm.
the average and extremes for these teeth are : upper, 20 • 5
(19-22) ; lower, 24 • 0 (22 • 8-26 • 0).* In all the dog skulls which
I have examined, representing such different breeds as the pug,
fox-terrier, bloodhound, mastiff, ancient Egyptian, ancient
Peruvian, Eskimo (Greenland and Alaska) and American Indian,
the teeth are strictly of the wolf type, never showing any
approach to that of the jackal (fig. 62).

Canis lupus lupus Linnaeus.

1758. [Cams'] liqms Linnaeus, Syst. Nat., i, 10th ed., p. 39 (Sweden).
1792. C[anis] Uipiis flavus Kerr, Auim. Kingd., p. 137 (France and

Gerinauy).
1804. Canis lupus nicjer Hermann, Observ. Zool., p. c!2. Not of Kerr,

1792 (Forest of Hagenau, Alsace, Germany).
1839. ? {Canis hqms] var. canus de Selys-Longchamps, Etudes de Micro-

mamm., p. 144 (nomen nudum).
1839. ? [Canis lupus] var. fulvus de Selys-Longchamps, Etudes de Micro-

mamm., p. 144 (nomen nudum).
1841. Lupus oricntalis Wagner, Schreber's Saugthiere, Suppl., ii, p. 367

(Europe).
1857. Ca7iis lupus Blasius, Saugethiere Deutschlands, p. 180.
18G3. [Canis lupus] var. major Og6rien, Hist. Nat. du Jura, iii, p. 59

(Lower slopes of the Jura).
1803. [Canis lupus] var. minor Ogerien, Hist. Nat. du Jura, iii, p. 09

(Higher portions of the Jura).
1897. Canis lupus minor Mojsisovics von Mojsvar, Thierleben der osterr.-

bung. Tiefebeuen, p. 241 (Southern Hungary). Based on the

"Rohrwolf," an animal supposed to be smaller and greyer tban

true luptts.
1910. Canis lupus and C. lupus lycaon-f Trouessart, Faune jMamm.

d'Europe, p. 90.

Tyjye 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, wbile 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 nxxxix,
the basis of the name, is a copy of Buffon's plate xli, representing an
animal brought alive to Paris from Canada.

314

CARNIVORA

not markedly tawny ; white of tliroat not extending to cheeks.
The few skulls examined agree with Asiatic specimens in having
the outer cusps of 7u^ moderately large, the iiaracone with trans-
verse diameter of base about equal to width of large flattened
portion of crown.

Measurements. — For cranial measurements see Table, p. 316.

Speciincns examined. — Four 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 signatii.s Cabrera, Bol. Real Soc. Espafi. Hist. Nat.,

Madrid, vii, p. 195.
1910. Canis lupus signahis Trouessart, Faune Mamm. d'Europe, p. 91.

Ttjpe locality. — Escorial, Madrid, Spain.

Geographical distrihution. — 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 atid female from Province of Burgos, Spain : head and
body, 1130 and 1180 ; tail, 350 and 380 ; hindfoot, 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.

Itemarhs. — 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)- conspiciiously 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
bulhe.

i. Seville, Spain. {A. Ruiz.) Lord Lilford (p). 95. .3. 3. G.

i, 9. Riocabado, Burgos. Hon. N. C. Roths- 11. 10. 5. 1-2.

[Ecv. S. Gonzalez.) 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 consi^icuous 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. Eeal Soc. Espafi. Hist. Nat.,

Madrid, vii, p. 197.
1910. Canis lupus deitanus Trouessart, Faune Mamm. d' Europe, p. 91.

Tijpe locality. — Moratalla, Murcia, Spain.

Geographicnl distribution. — Now known from the type locality

only- _ .

DicK/nosis. — Smaller than Canis lupus lupus and brighter in
colour, the general ajipearance much as in C. aureus.

Measurements. — Unknown.

Specimens examined. — I have seen the two living examples in Madrid
on which the form was based.

Remarlcs. — Tn general appearance the two Moratalla wolves
are strikingly different from Canis lupus. Unfortunately their
true characters are not yet known.

CANIS AUREUS Linnaeus.

1758. [Cams'] aureus Linnaeus, Syst. Nat., i, 10th ed., p. 40.

1835. Canis aureus var. moreotica I. Geoffroy, Exp6d. Sci. de Mor6e. Zool.,

pi. I (Morea, Greece).
1841. Clams'] dalmatinus Wagner, Schrober's Siiugthiere, Suppl., ii, p. 383

(Dalmatia).
1841. C[a.nis] cjnecus Wagner, Schreber's Sangthiere, Suppl., ii, p. 383

(Peloponesus).
1892. Canis aureus balcanicus Brusina, Glasnik Hrvatskoga Naravoslovnoga

Drustva, Zagreb, vii, p. 317 (Drava River, Croatia).

Ti/2>e locality. — Province of Lar, Persia.

Gcoe/raplilcal dlstributton. — 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 buff'y cinnamon, clouded Ijy black hair
tips along dorsal region, nearly clear on sides, and becoming clear
Ijright cinnamon on outer surface of legs and ear and on area
around and behind base of ear ; muzzle more heavily washed
with black ; from middle of back to base of tail the cinnamon is
replaced by whitish, causing a rather noticeable contrast between
this region and the surrounding parts when fur is disarranged ;
tail essentially like back, the basal half above greyish, the
terminal half and underside bufiy much o^•erlaid with black ;
pencil blackish ; underparts and inner side of legs dull bufiy
grey ; chin blackish ; feet dull buffy.

316

CARNIVORA

Observations.

Basal suture closed.
„ closed.

„ open.
,, closed.

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C. lupus lupus.

Sweden .

Northern Sweden .
Russia .

Italy : near Sassello, )
Liguria . /

C. lupus signatus.

Spain : near Seville
Prov. Burgos

C. aureus.

Greece : Pirteus
Asia Minor : Khotz

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318

CARNIVORA

Skull. — The skull is much smaller than that of Can!.s lupus.
In form it differs slightly in the less elevated frontal region and
somewhat more inflated auditory bullae.

Teeth. — While agreeing with those of Canis lupus in general

form and in the position of the cusps, the teeth

are on the whole more trenchant in character, as

shown by the general tendency toward narrowness

of crown and prominence of ridges. This is

particularly noticeable in the upper molars, in

which the large cusps are relatively higher, more

slender, and less terete than in Canis lupus, their

cutting ridges much more developed ; transverse

diameter of metacone in m^ noticeably less than

width of inner portion of crown ; cingulum on

outer border of both upper molars wide and

conspicuous in contrast with narrow cusps,

Fig. 62. showing no tendency to become obsolete in

Canis aureus, region between paracone and metacone. Lower

^*chee ™teeuf'^ carnassial with metaconid actually as well as

Nat. size. relatively larger than in Canis lupus, and posterior

heel with area equal to nearly half that of anterior

portion of tooth, its cusps strongly developed.

Measurements. — For cranial measurements see Table, p. 316.

Specimens examined^ — One from Greece (Piraeus) ; numerous others
frora Asia Minor and India.

Remarls. — The single specimen from Greece agrees sufficiently
with a series of five from Khotz, near Ti'ebizond, Asia JNIinor,
to make it appear vmwise, in the absence of more satisfactory
material, to use one of the Balkan names.

1. Pirccus, Greece. (C. Mottaz.) Hon. N. C. Eoths- 8. 10. 2. 49-50.

child (p). _

Genus ALOPEX Kaup.

1829. Alopex Kaup, Entw.-Gesch. uud Natiirl. Syst. Europ. Thierw., i,

p. 83.
1857. Leiicocyon Gray, Proc. Zool. Soc, London, p. 512.

Type species. — Canis lagopus Linna?us.

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 Cants ; 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 Ganis 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
Cams 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 OF ALOPEX.

Condylobasal length of skull about 130 in males,

121 in females (Scandinavia and Finland)... A. Icujopus, p. 319.
Condylobasal length of skull about 120 in males,

114 in females (Spitzbergeu) A. spitzbcrgcncnsis, p. 324.

ALOPEX LAGOPUS Linna3US.

1758. [Canis] lagopus Linnaeus, Syst. Nat., i, 10th cd., p. 40 (Lapland).
1816. V[ul2}es] arctica Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1033

(Renaming of Canis lagopus).
1820. C[_anis] vulpes cxrulca Nilsson, Skand. Fauna, i, p. 88 (Lajjland).
1827. [Canis lagopus] & argentcus Billberg, Synopsis Faunaj Scaudinavise,

p. 14 (Lapland).
1910. Vulpes lagopiis Trouessart, Fauue IMamm. 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 cliaracters. — 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 uuderfur 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

CARNIVORA

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.
Alopez latjopus. x i.

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 ; underfur and posterior surface of ear
light drabby grey. In the " blue " phase the entire animal is at
all seasons a bluish drab, usually washed with sepia on head and
feet, and sprinkled with pure white hairs on face, chin and throat.

Skull.- — The skull is shorter and narrower than that of VuJpes
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
F. 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
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foramen over space between pm^
and pm'^ ; auditory bulla relatively
as large as in the larger animal ;
depth of brain-case through bulla
equal to greatest breadth above
roots of zygomata ; postorbital pro-
cesses slightly less concave on
underside and somewhat more
flattened above ; mandible with
ramus relatively a little deeper
and more compressed than in the
common fox.

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closely resemble those of Vulpjes
vulpes except for their slightly
smaller size. Incisors both above
and below with relatively wider crowns than in the common fox,
the cingulum of i^ and «"- much more developed and forming a

Fig. 64.
Alopcx lagopus. Teeth.

322

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noticeable inner and outer secondary cusp (the former soon
wearing away in i^) much as in Canis ; along posterior border of
crown the cingukim shows a strong tendency to develop irregular
tubercles, a condition not observed in Vulpes vuljws. Canines
and premolars with no special peculiarities, the form of the upper
carnassial quite as in V. vuljpes. 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 vidpes.

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 Bechstein, Thomas Pennant's allgem. Uebersicht
d. vierfiiss. Thiere, i, p. 270 (Iceland).

1799. ? Canis grocnlandictis Bechstein, Thomas Pennant's allgem. Ueber-
sicht d. vierfiiss. Thiere, i, p. 270 (Greenland).

1898. Canis lagopus spitzhcrgenensis 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,
Faune Mamm. d'Europe, p. 97.

Type locality. — Spitzbergen.

Geographical distribution. — Spitzbergen ; also Iceland and
Greenland 1

Diagnosis. — Like Alopex lagopus but smaller, condylobasal
length of skull about 120 in males, about 114 in females.

Colour. — Type (summer pelage) : colour pattern well defined
and exactly as in A. lagopus, but dark areas wood-brown against
which the cream-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 aud dental measurements see
Tables, pp. 322, 323.

vuLPES 325

SpecUnens examined. — Six, all from Spitzbergen(B.M.and U.S.N.M.): —

6 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.)
6, 9 skulls. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 4-5.

Genus VULPES Oken.

1816. V[ulp)es] Oken, Lebrb. d. Naturgesch., in, pt. 2, p. 1033, in full on
p. 1034 and in index, p. 1268 (Vulpes communis Oken = Canis
vulpes LinnEEUs).

1821. Vulpes Bowdich, Anal. Nat. Classif. Mamm., p. 40 {Canis vulpes).

1857. Vulpes Blasius, Siiugethiere Deutschlands, p. 178 (Sub-genus of
Canis).

Type species. — Canis vulpes Linnteus.

Georjrapliical distribution. — Northern portion of the northern
hemisphere from about the hmit of tree growth south to Morocco,
India and Mexico ; in Europe west to Ireland.

Characters. — Skull slender and low (depth of brain-case less
than one-third condylobasal length) ; interorbital region nearly
flat, the frontal sinuses scarcely inflated, the postorbital 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 (tig. 65).

Bemarhs. — 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 OF 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. iclmusx, 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 T'. vulp>cs, p. 326.

Teeth larger and more robust, the premolars tend-
ing to be slightly spaced or in contact, their
secondary cusgs well developed (Scandinavia)... V. v. vulpes, p. 330.
Teeth smaller and less robust, the premolars tend-
ing to be widely spaced, their secondary cusps
usually obsolete or absent.
Yellowish and reddish tints bright ; posterior
half of back seldom much frosted with whitish ;

tail never clear grey (Central Europe) T. v. crucigera, 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. silaeea, p. 333.

326

CARNIVORA

Fig. 65.
Incisors and canines from front, of Caniji (n), Alo2)cx (h), and Vttlpes (c).

VULPES VULPES LinnsBus.

(Synouyniy under subspecies.)

Type locality. — TJpsala, Sweden.

GeograjjJncal clistrihution. — 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,

VULPES 327

thick and bushy, with abundant underfur ; longer hairs of back
normally concealing the underfur ; feet as in Cauls, 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 nai-es 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 bvdlai 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 Canis. Rostrum moderately long (the distance from
orbit to gnathion about equal to that from postorbital jarocess 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^ or over space between pjx^
and pviK Palate relatively narrower tlian in the European

Fig. 66.
Vulpes vulpes. X i.

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
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^ 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. pjg gy

Upper carnassial with inner VtUpes vuipes. Teeth. Nat. size.

lobe better developed than in

Canis, and bearing a distinct terete cusp, its position a little
more forward and outward than in the dogs, so that it appears
to lie in or nearly in the main axis of the tooth. Lower carnassial
with 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
I'elatively 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.

330

CAKNIVORA

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 VULPES Linnseus.

1758. ICanis'] vtilpes Linnaeus, 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{^idpes\ vulgaris Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1034

(Renaming of vulpes).
1820. Canis nigro-argenteus Nilsson, Skand. Fauna, i, p. 91 (Lofoten

Islands, Norway).
1827. ICariis vitlpus'] y nlgrocaudatus Billbarg, Synopsis Faunae Scandi-

navise, p. 12 (Uppland, Sweden).
1827. [Canis vulpus'] ^ variegatus Billberg, Synopsis Faunae Scandinaviae,

p. 13 (Uppland, Sweden).
1827. [Canis vulpus] t) lineatus Billberg, Synopsis Faunae Scandinaviae,

p. 13 (Skane, Sweden).
1830. ? [Vulpes] comimmis Burnett, Quart. Journ. Sci. Lit. Art, xxviii,

1829, p. 349 (Substitute for vulpes). Nomen nudum.
1898. Vulpes vulpes Thomas, The Zoologist, 4th ser., ii, 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.

Bemarks. — 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). G4. 3. 8. 2.

11 skulls. Egersund, Stavanger, K. H. Schaanning (c). 11. 6. 3. 1-11.
Norway.

VULPES 331

VuLPRS VULPES CHUCIGERA Bechstein.

1789. [Canis] cmcigera Bechstein, Gemeinn. Naturgesch. Deutschlands,
I, p. 250 (Thuriugen, Germany).

1792. C[anis] Vulpes alopex europaeus Kerr, Anini. Kingd., p. 142 (Bur-
ginidy, France).

1797. Canis vulpes alba Borkhausen, Deutsche Fauna, i, p. 33 (Vogelsberg,
near Rudigshain, Hessen, Germany).

1797. Canis vulpes nigra Borkhausen, Deutsche Fauna, i, p. 33 (Hessen
and Thiiringen, Germany).

1801. C[anis'] vliidpes] lutea Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 628 (Thiiringen, Germany).

1801. Clams'] ■u[«Zp<;s] cinerea Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 628 (Thiiringen, Germany).

1832. Canis mclanogaster Bonaparte, Iconogr. Fauna Ital., i, fasc. 1
(Neighbourhood of Rome, Italy).

1841. Vulpes hyponielas Wagner, Schreber's Siiugthiere, Suppl., ii, p. 405
(Oberbayern, Germany).

1857. Canis vulpes Blasius, Saugethiere Deutschlands, p. 191.

1861. ? V\ulpes'] vulgaris meridionalis Fitzinger, Wissensch.-pop. Natur-
gesch. der Siiugeth., i, p. 194 (Dalmatia).

1910. Vulpes vulpes (part) and V. vulpes melanogaster Trouessart, Faune
Mamm. d'Europe, pp. 93-94.

Type locality. — -Thuringen, 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 yellowLsh
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

CAENIVORA

size as in true Vulpes imlpes, and the small premolars are seldom
in contact, the spaces between them visually 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, 144db ; 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).

Feance : Ax-les-Thermes, Ari^ge, 3 ; Oaterille, 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) ; Riesengiberge, 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, 4 (Ghidini) ; Garbagna, Piedmont, 1 (Genoa) ;
Borghetto S. Spirito, 1 (Genoa) ; Vargo, Liguria, 1 (Genoa) ; Torriglia,
Liguria, 1 (Genoa) ; Cornigliano, Liguria, 2 (Genoa) ; near Genoa, 1 ;
Molasana, 1 (Genoa) ; Tuscany, 1 ; Pisa, 1.

Sardinia: Zinnigas, Siliqua, 1 (Genoa).

Greece : Cephalonia, 1 ; Tatoi, near Athens, 2.

Memarhs. — 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, Sutherlaudshire, E. R. Alston (p). 79. 9. 25. 80.

Scotland. {H. Broivn.)

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. 4.

9. Cappagh House, Waterford. R. J. Ussher (c & p). 96. 12. 28. 1.

1. Northumberland, England. Rev. H. H. Slater 0. 2. 24. 5.

(0 & p).

333

2 6. Thame, Oxfordshire.

2 6. Ditchling, Sussex.

3. Tunbridge Wells, Sussex.

2 6,9. Ax- les- Thermes, Ariege,

Prance.
?. Caterille, Haute-Garoime,

900 m. {A. Robert.)
9. PicSessire,Haute-Garonne.

{A. Robert.)
6. St. Aventin, Haute-Ga-

ronne, 900 m. {A. Robert.)
6. Ingelheim, Rheinhessen,

Germany.
3 skulls. South Germany.

6.
skull.

lyg-

9.
6,9.

Genoa, Liguria, Italy.

Pisa.

S. Italy. (Prof. Savi.)

Tuscany.

Tatoi, Athens,
(G. Mottaz.)

Greece.

Hon. N. C. Roths-
child (p).

Guy Dollman
(c & p).

C. H. B. Grant (c).

V. BuiUes (c & p).

O. Thomas (p).
0. Thomas (p).
0. Thomas (p).
C. Hilgert (c).
Dr. A. Giinther (c).

Marquis G. Doria (p),

Zool. Soc. Mus.

Zool. Soc. Mus.

Purchased (Dr.
Riippell).

Hon. N. G. Roths-
child (p).

0. 10. 31. 1-2.

(9. 11. 8. 1.
(10. 9. 13. 1.

1. 2. 15. 1-3.
8. 3. 27. 6-7.
8. 3. 27. 14.
8. 7. 15. 2.

8. 7. 15. 3.

8. 7. 15. 4.

8. 11. 2. 15.

59.9.6.83,88.

175. k.

88. 12. 1. 3.

58. 5. 4. 126.

55.12.24.240.

45. 7. 22. 15.

8. 10. 2. 22-23.

VULPES VULPES SILACEA Miller.

1907. Vulpes vidpes silaceus Miller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 393, November, 1907. Type in British Museum.
1910. Vulpes vulpes silaceus Trouessart, Faune Mamm. d'Europe, p. 45.

Type locality. — Near Silos, Province of 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 buli" without bright
yellowish or reddish tints, the posterior half of back conspicuously
frosted with whitish (except in abraded pelage) the tail often a
clear greyish.

Measurements. — Type (adult male) : head and body, 750 ;
tail, 370; hind foot, 150. Adult female from near the type
locality: head and body, 670; tail, 330; hind foot, 125. Old
male from Elche, Alicante : head and body, 770 ; tail, 480 ;
hind foot, 160 ; ear from meatus, 113.

Specimens examined. — Eighteen, from the following localities in
Spain : Olot, Gerona, 1 (probably not typical) ; vicinity of Silos, Burgos, 5 ;
near Burgos, 1 ; Arrechavaleta, Vitoria, 1 ; Torres del Alio, Coruna, 1 ;
Madrid, 1 ; near Seville, 3 ; Goto Donana, Huelva, 3 (B.M. and U.S.N. M.) ;
Elche, Alicante, 1 ; Barracas, Castellon, 1.

Itemarlcs. — 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.
(ZV. Gonzalez.)

0. Thomas (p).

8. 2. 9. 49.

i.

Torres del Alio, Coruna.

Dr. v. L. Seoane
(c & P).

94. 5. 29. 1.

6.

Madrid.

M. de la Escialera (c).

8. 9. 24. 7.

6, 9.

Seville. {A. Ruiz.)

Lord Lilford (p).

95. 9. 4. 7-i

2 6.

Goto Dofiana, Huelva.

A. Chapman (c & p).

8. 3. 8. 4-5.

6.

Elche, Alicante.

G. S. Miller (c).

8. 8. 4. 48.

6 juv.

Barracas, Castellon.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 50.

VULPES ICHNUS.S: Miller.

1907. Vulpes ichnusx Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 391,

November, 1907. Type in British Museum.
1910. Vul]yes vulpes ichnusie Trouessart, Faune Mamm. d'Europe, p. 94.

Type locality. — Sarrabus, Sardinia.

Geographical distributi on. ^Sebrdinia, and Corsica.

Diagnosis. — Smaller than any of the races of Vtiljyes 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-
rufous 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 ; ear from
meatus, 70. Adult male and female from the type locality :
head and body, 640 and 590 ; tail, 280 and 290 ; hind foot,
127 and 125 ; ear from meatus, 74 and 71. Adult female from
Siliqua : head and body, 600 ; tail, 350 ; hind foot, 127 : ear, 74.
Adult from near St. Florent, Corsica ; hind foot, 123 ; ear from
meatus, 73.

Specimens examined. — Eight, from the following localities : —
Corsica : Grotto Campu Consule, near St. Florent, N.W. Corsica, 1

(Major).

Sardinia : No exact locality, 1 (Turin) ; Lanusei, 1 ; Sarrabus, 3 (B.M.

and Genoa) ; Zinnigas Siliqua, 2.

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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 Sai'dinia ma}^ l)e due to recent
introduction. The Corsican specimen, though a mummy and
without fur, appears to be a typical example of Vuljjes 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 MUSTELID^.

835. Miistelidfn Swainson, Nat. Hist, and Classif. of Quadrupeds, p. vii, 361.

Geographical (Jistrtbutioii. — 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.

BemorJcs. — The family Mitstelidse is, next to the Ganidee, 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 MUSTELID^.

Grown of upper carnassial triangular or rhombic in
outline, its length and width sub-equal ; lower car-
nassial with anterior triangle distinct, the meta-
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

Meliniv) Melcs, p. 341.

Upper molar about equal to carnassial ; skull broad
and fiat, the rostrum broader than long ; tail long,
densely furred, very muscular ; habits aquatic

(Otters, sub-family Lutriniv) Lutra, p. 354.

Crown of upper carnassial not triangular or rhombic in
outline, its length much greater than its width ;
lov/er carnassial with anterior triangle obsolete or
absent (represented by the two outer cusps only),
the metaconid when present much smaller than the
other cusps.

MELES 341

Premolars — .

Lower caniassial without metaconid ; skull robust
(in European species considerably more than
100 mm. in coudylobasal length) ; form robust ;
tail short (Wolverenes, sub-family Guloniiuv)... Giilo, p. 433.

Lower carnassial with evident though small meta-
conid ; skull slender (in European species con-
siderably less than 100 mm. in coudylobasal
length) ; form slender ; tail long (Martens, sub-
family Mustdinn) 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 i^f^/s^eZ^lw^i) Vormela, ]). 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., snh-isiuiilj Musteliritr) Mustela, p. 381.

Sub-Family MELIN.^.
1857. Melinx Baird, Mamm. North Amer., p. 148.

Geographical distribution. — Temperate an<l 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.

i^ewirtr/cs-.— 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. Melcs Brisson, Regn. Anim. in Classis ix, distrib., 2nd ed., p. 13

(Meles Brisson = Ursns mcles Linnseus).
1780. Mcles Storr, Prodr. Meth. Mamm., p. 34. First use of name by an

author following the Linnsean system (Ursns meles).
1795. Taxus Geott'roy and Cuvier, Magasin Encyclopedique, ii, p. 184

(Urstis meles Linnaeus).
1815. Melesium Ralinesque, Analyse de la Nature, p. 59 (Modification of

Meles).
1857. Meles Blasius, Siiugethiere Deutschlands, p. 202.

Type species. — Meles * Brisson = XJrsus meles Linnaeus.

* Not ^^ Meles 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 Linntean system.

342

CAENIVORA

Geograpliical distribution. — Northern temperate portions of
Old World ; in Europe from central Scandinavia to the
Mediteri'anean, and from Ireland eastward.

Characters. — Skull narrow and high (depth of brain-case
much more than half mastoid breadth), the zygomatic arches not
widely expanded, and postorbital region not specially narrowed
(distance from point of greatest narrowing to zygoma less than
breadth of postorbital constriction) ; rostrum elongate, the
distance from orbit to gnathion about three-fourths length of
brain-case ; auditory bulla elevated along inner margin, but else-
where somewhat flattened, the meatal tube distinct ; paroccipital
process robust, standing out conspicuously behind bulla ; dental
formula : i ~, c ^, ])m ~* in ^ = 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 lai'ge 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 cusj^s 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.

Remarks. — The genus Meles contains about half a dozen
named forms, the status of several of which is at jaresent not
clearly understood. Two species occur in Europe.

KEY TO THE EUROPEAN FORMS OF MELES.

Maxillary tooth-row, exclusive of incisors, about
35 mm. ; auditory bullae rather strongly in-
flated, the inner border not ridge-like (Crete) M. arcalus, p. 352.
Maxillary tooth-row, exclusive of incisors, about
40 mm. ; auditory bullte 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. marianemis , p. 352.

* In adults usually ^~ , owing to the early disappearance of the small

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 I'ange 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-deiined 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 buflfy
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.

SJcull. — Except for the greatly developed sagittal crest, the

344

CARNIVOKA

height of which in old individuals is equal to nearly half inter-
orbital breadth, the skull of Melea meles is rather smooth and
without marked angularity. Depth of brain-case (without crest)
and depth through posterior portion of orbit nearly equal, and
about two-thirds mastoid breadth ; depth at front of nasal about
two-thirds that at orbit. Dorsal profile (without crest) nearly
flat from middle of brain-case to interorbital region, convex over
posterior half of brain-case ; rostrum sloping downward at angle
of about 30° with surface of brain-case. When crest is fully

Fig. 68.
Meles meles. x i.

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 througli-
out. Palate moderately wide, the distance between molars
contained about 2h times in distance from gnathion to level of
posterior edge of molar. Incisive foramina small, at level of
space between canine and /^, 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

Fig. 69.
Meles tneles. x i.

space shoi't 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

CARNIVORA

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.
Mcles meles. X '}.

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 Musfelidse. Upper
incisors robust, the crowns somewhat higher than wide when
viewed from in front, the teeth closely crowded in a slightly
convex row, the longitudinal diameter of crown a little greater
than transverse diameter ; i^ and 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 Ijut distinct
heel ; P with crown area about double that of f- and nearly half

347

that of canine, its crown higher than in the other incisors,
owing to lower insertion of root, but extremity of cutting edge
not extending beyond level of smaller teeth ; laterally the cutting
edge is continued down outer-posterior margin of shaft where it
functions against lower canine. Lower incisors not so large as
upper, forming a continuous, nearly straight row between
canines, the shafts straight, sloping obliquely forward, the root
of ^2 implanted behind the others ; cutting edge irregularly
2-lobed, that of i^ longer than the others. Canines rather short
and weak, the shaft simple, that of upper tooth nearly straight,
that of lower tooth strongly curved backward, the enamel surface
of both essentially smooth, though that of lower canine shows
indications of a rudimentary cingulum in front. Anterior
premolar both above and below a minute or spicular tooth closely
crowded between canine and
second premolar, pm"^ early
deciduous, pm^ usually more
persistent. Second premolars
similar to each other, the crown
area about equal to that of P,
the outline irregularly oval when
viewed from above, the cusp
about as high as long, its apex
slightly in front of middle of
crown ; root of each tooth single,
that of pni.^ showing a tendency
to become divided longitudi-
nally. Other small premolars
(pm-^, pmi^ and p)n^ alike in form,
the crowns laterally compressed,
triangular when viewed from
the side, with apex slightly in
front of middle, prii^ and pm^
nearly alike in size, pm^ some-
what longer ; cingulum very slightly developed, not forming
secondary cusps. Upper carnassial triangular in outline, the
pustero-internal border longest ; outer, trenchant portion of teeth
consisting of a main antei'ior cusp resembling pwt^ 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 nietaconid 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

Fig. 71.
Meles meles. Teeth. Nat. size.

348

CARNIVORA

which the anterior is the larger), a large cusp similar to postero-
external cusp at middle of inner margin ; posterior margin
occupied by three or four minute cusps soon disappearing with
wear, and a similar minute cusp sometimes in angle behind
protoconid and metaconid. Second lower molar flat, terete, its
area about half that of crushing portion of carnassial ; middle of
crov/n with basin-like concavity ; margin with small but evident
antero-external and postero-external cusps, and a smaller
elevation at middle of inner edge. Upper molar rhomboidal
in outline, the inner and outer margins parallel, the former
nearly li times as long as latter; outer border occupied by two
rather large, subterete, conical cusps, the anterior (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 tive 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 hyjjocone.

Bemarks. — 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 Linnaeus.

1758. [Ursus] meles Linnseus, Syst. Nat., i, 10th ed., p. 48 (Sweden).
1785. [Meles] taxus Boddaert, Elenchus Animalium, i, p. 80 (Europe).
1789. [TJrsus meles'] 0 alba Gmeliu, 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. 876 (Reuaming of Urstis
meles) .

1816. Meles europRus Desmarest, Nouv. Diet. d'Hist. Nat., in, p. 465

(Renaming of Ursus meles).
1822. Taxus meles F. Cuvier, Hist. Nat. des ]\Iamm., iii, fasc. 36, January,

1822.

1827. [Meles] communis Billberg, Synopsis Faunae Scandiuavise, p. 16
(Renaming of Ursus meles).

1827. [Meles communis] )3 caninus Billberg, Synopsis Faunae Scandinavise,
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[cles] typicus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., iv, p. 384, November, 1899.

MELES 349

1906. Melcs meles britannicus Satunin, ]\Iitteilungen des Kaukasischen
]\Iuseums, ii, 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. Melcs melcs and M. meles britannicus Trouessart, Paune Mamm.
d'Europe, p. G9.

Type localily. — 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 modei-ately
light, the sides not often conspicuously whitish.

Measurements. — Adult male and female from Woodpark,
Gal way, 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 Bouconne, 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.

tSpecimcns examined. — Thirty-five, from the following localities : —

Ireland : 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; Cornwall, 1; no exact locality, 4.

Norway : Holme, ilandal, 7 ; Egersund, Stavanger, 2.

Sweden : ^Middle Sweden, no exact locality, 1 (Stockholm) ; Takern,
Ostergotland, 1 (Stockholm) ; Smaland, 1 (Stockholm) ; Skime, 2 (Stock-
holm) ; no exact locality, 1.

France : Foret de Bouconne, Gers, 1.

Germany : Ingelheim, Rheinhessen, 1 ; Strass, near Burgheim,
Bavaria, 1.

Italy: Liguria, 1 (Genoa).

6, 9. Woodpark, Galwav, Ire- R. F. Hibbert (p). 95. 6. 2. 1.
land. " 95. 12. 3. 1.

{Co-types ot M. m. britannicus, Satunin.)
Burnley, Lancashire, Hon. N. C. Roths- 0. 10. 31. 4.

England.

child (p).

• 6.

Ross, Herefordshire.

Hon. N. C. Roths-
child (p).

0. 10. 31. 3.

', 3 jxiv. st.

Kentchurch, Hereford-

W. E. de Winton

shire.

(c & P).

6.

Tetworth, Oxfordshire.

Hon. H. Parker
(c & P).

6.

Ibstone, Buckingham-
shire.

A. H. Cocks (c & p).

4. 1. 25. 1.

6.

Horsham, Sussex.

E.G. 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-
wright).

64. 3. 8. 1.

4 <5, 2 ?.

Holme, Mandal, Norway.

R. J. Cuninghame
(c & p).

S. S. 9. 7-12.

2 skulls.

Egersund, Stavanger.

K. H. Schaanning

(11. 6. 3. 13.

(c).

111. 10. 23. 2.

350

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CARNIVORA

?. Foi'et de Bouconne, Gers, O. Thomas (p). 8. 7. 15. 5.

250 m. France. (A.

Bobert.)
?. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 16.

Germany.

Meles meles marianensis Graells.

1897. \_Meles taxus] var. marianensis Graells, Mem. Beal Acad. Gien.,

Madrid, xvii, p. 170 (Central Spain).
1899. Meles meles mediterraneiis 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, Fauna Mamm. d'Europe, p. 70.

TyjJe locality. — Central Spain.

Geograjjh ical disfrihufion. — Iberian Peninsula.

Diagnosis. — Skull and teeth large, the teeth frequently attain-
ing maximum size for species ; colour light, the sides often
conspicuously whitish, especially in region bordering dark ventral
area.

Measurements. — For cranial and dental measurements see
Table, p. 351.

Specimens examined. — Seven, from the following localities in Spain :
Arrechavaleta, Vitoria, 1 ; Quintanar de la Sierra, Burgos, 2 ; near
Seville, 2 ; Goto Doilana, Huelva, 2.

Remarhs. — 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.

(J, ?. Quintanar, Burgos. Rev. Saturio Gou- 8. 7. 7. 19-20.

zalez (c).

6, ?. Seville. {A. Pmiz.) Lord Lilford (p). 95. 3. 3. 7-8..

(95. 3. 3. 7. Type ol mediterraneus Barrett-Hamilton.)

6, ?. Goto DoBana, Huelva. A. Chapman (p). 8. 3. 8. 6-7.

MELES ARCALUS Miller.

1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and ]\Iag. Nat.
Hist., 7th ser., iv, p. 131, November, 1899 (part).

1906. Meles meles mediterrane7is Bate, Proc. Zool. Soc, London, 1905, ii.

p. 318, April 5, 1906.

1907. Meles arcalus Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 394,

November, 1907. Type in British Museum.
1910. Meles arcalus Trouessart, Faune ]\Iamm. d'Europe, p. 70.

Type locality. — Lassethe Plain, Crete.
Geographical distribution. — Islaaid of Crete.
Characters. — Size small (upper length of skull in adult less than
110 mm. ; maxillary tooth-row, exclusive of incisors, about 35 mm. ;

MELES 353

DENTAL MEASUREMENTS OP MELES MELES AND M. ARC ALUS.

■ Lower molar

Locality.

Number.

Sex. '

i

Lpper.uolar. , ^° (^^st).

M. meles meles.

1

Sweden : no exact locality

64. .3. 8. 1

9

13-0 X 11-4

15-4 X 7-0

Wegeliu ....

Stockholm

6?

15-0 X 13-0

16-2 X 7-8

Skane ....

,,

J?

15-2 X 13-0

18-0 X 8-2

Ulriksdal ....

,,

6 juv.

13-4 X 11-6

15-8 X 7-8

SmSland ....

,,

9?

14-2 X 11-8

16-2 X 7-4

Orebro ....

J,

(? juv.

14-2 X 11-8

15-8 X 7-0

T&kern ....

,,

6?

13-6 X 11-2

15-6 X 7-2

Norway : Holme, ]\Iandal

152622

6

11-6 X 10-4

14-0 X 6-4

.. ,,

8. 8. 9. 9

S

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

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 : Ross, Hereford

0. 10. 31. 3

6

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

6?

13-4 X 12-0

16-4 X 8-0

„ . .

211c

9?

14-0 X 12-4
14-2 X 12-2

16-2 X 8-2

—

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 : Woodpark, Co. Galway

95. 12. 3. 1

15-6 X 12-8

17-0 X 7

8

95. 6. 2. 1

14-8 X 12-2

16-0 X 7

8

Denmark: Zealand.

Andersen

6

14-0 X 13-2

17-0 X 7

8

,,....

Copenhagen*

6

15-0 X 12-6

18-5 X 8

2

^,

6

15-0 X 13-7

18-3 X 8

3

^,

6

15-5 X 13-6

18-0 X 9

0

" '. '. '.

J,

9

14-6 X 12-5

17-5 X 7-7

various localities, (Maxi-1
13 specimens : \ mum J

16-3 X 13-6

/Mini-)

13-0 X 12-0

" (mum J

"

various localities, jMaxi-
17 specimens : \ mum

18-7 X 8-5

"

j Mini-)
" \mum!

15-0 X 7-0

France: Bouconne, Gers.

8. 7. 15. 5

9

14-0 X 12-0

16-2 X 7-2

Germany: Ingelheim, Rheinhessen .

8. 11. 2. 16

9

14-8 X 12-8

l'^-2 X 7-8

Italy: Borghetto S. Spirito, Liguria

33 Genoa

9

13-6 X 11-0

15-6 X 7-2

M. meles marianensis.

Spain : Quintanar, Burgos

8. 7. 7. 19

: 6

15-2 X 12-6

17-4 X 8-6

Seville ....

i 95. 3. 3. 7t

6

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 Donana, Huelva .

8. 3. 8. 0

6

16-4 X 13-2

17-8 X 8-0

M . .

8. 3. 8. 7

9

14-8 X 12-0

17-6 X 8-2

M. arcalus.

Crete

5. 12. 2. 17t

9 juv

1 13-6 X 11-2

15-4 X 7-2

5. 12. 2. 38

ad.

13-8 X 11-6

Measured liy H. AVinge.

t Type of m«Utermvei(x Baiiett-Hamilton.

t Type.

2 A

354

CARNIVORA

hind foot about 25 mm.) ; auditory bulla strongly inflated, the
highest region near middle of bulla proper (exclusive of meatal
tube) and so broadly rovxnded as not to form a longitudinal
ridge, the region between highest portion and meatus not notice-
ably concave ; teeth smaller than in Meles meles (lower molars
together about 20 mm. in length), with smaller cusps tending to
be more strongly developed ; metacone of m^ relatively largei-
than 31. meles, so that breadth of crown through this cusp is as
great as or greater than that through paracone ; colour as in
Meles meles marianensls or slightly paler.

Measurements. — For cranial and dental measurements see
Tables, pp. 351, 353.

Specimens examined. — Four, all from Crete.

6,9. 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.

?. Crete. H. 0. Jones, R.N. (c). 99.6.13.1.

Sub-Family LUTEINS.
1857. Lutriniv Baird, Mamm. North Amer., p. 148.

Geographical distrihtdion. — 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.

Bemarhs.- — 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 ix distrib., 2nd ed., p. 13

(Lutra Brisson = Mnstela lutra Linnteus).
1780. Lutra Brilnnich, Zoologiae Fundamenta, p. 34. First use of name

by an author following the Linnsean system [Mustcla Intra).
1806. Lutris Dum6ril, 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

= Ltitra lataxina F. Cuvier).
1857. Lutra Blasius, Saugethiere Deutschlands, p. 236.
1865, Barangia Gray, Proc. Zool. Soc. London, p. 123 {B. sumatrana Gray

= L'utra harang F. Cuvier).

LUTRA 355

1865. Lutrogale Gray, Proc. Zool. Soc. London, p. 121 [Lit tr a monticola

Hodgson).
1865. Nutria Gray, Proc. Zool. Soc. London, p. 128 (Luira felina Molina).
1867. Liiironectes Gray, Proc. Zool. Soc. London, p. 180 {L. wliitcleyi

Gray = L. Intra Linnseus?).

Type species. — Lutra * Brisson = Mustela Intra 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 J^J, j;«i ^^^, 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 LinnaiUS.

1758. [Mustela'] Intra Linnseus, Syst. Nat., i, 10th ed., p. 45 (Sweden).
1777. [Lutra] vulgaris Erxleben, Syst. Regni Anim., i, p. 448 (Renaming of

lutra).
1792. M[2istela'] Lutra piscatoria Kerr, Anim. Kingd., p. 172 (Renaming of

lutra).

* Not " Lutra lutra" (see footnote under Meles, p. 341).

2 A 2

356

CARNIVORA

1816. ■? LtUra fluviatilis Leach, Syst. Catal. Spec. ludig. Matnm. and

Birds Brit. Mus., p. G (uomen nudum : " lliver Otter").
1827. {Lutra vulgaris] /3 vmrinits Billberg, Synopsis Faunaj Scandinaviae,

p. 28 (Coasts of Scandinavia).
1830. ? [_Lutra\ fluviatilis Burnett, Quart. Journ. Sci. Lit. Art. sxviii,

1829, p. 349 (Substitute for Intra), nomen nudum.
1834. [Lutra] nudipcs Melchior, Den Danske Stats og Norges Pattedyr,

p. 50 (Coast of nortliern Norway).
1884. Lutra roensis Ogilby, Proc. 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, xxxviii, p. 34.

1885. [Luti-a] 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 Intra Trouessart, Faune Mamm. d'Europe, p. 86.

Tj/pe 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
thi'ee-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 stifi" 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-i-etractile, those on fore-feet best developed (about 8 mm. in

LDTRA 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 j^ad 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-

FlG. 72.
Lutra Intra. Nat. size.

what darker), with a drabby cast more evident in some lights
than in others, the hairs with a conspicuous metallic gloss ;
underfur light grey, the extreme tips of its hairs changing
abruptly to prout-brown ; on uiiderparts 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, cii, y. 1043, December lU, 1908.

358

CARNIVORA

Sometimes the brown is darker and richer than usual, or the
drab may be especially pronounced. Occasionally the long hairs
of tlie 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

Fig. 73.
Lutra hitra. 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 con.siderable portion of border
of rather large, somewhat upturned orbit, the greatest diameter

LUTRA 359

of which nearly ecjuals width of flattened interorbital region.
Rostrum short, broad and deep, the distance from front of
zygoma to gnathion barelj^ equal to greatest breadth across
canines, about equal to depth at front of orbit, and contained
about i^ 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

Fig. 74.
Lutra lutra. Nat. size.

angular projection marking posterior border of orbit. Palate
rather narrow, the distance between molars contained about
three times in that from gnathion to level of posterior border of
molar : incisive foramina moderate, about half as wide as long,
lying entirely between canines ; posterior extension of palate
about 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

CARNIVORA

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 jDrocess ; angular process
very short ; coronoid process narrowly rovinded 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. [Tpper
incisors forming a nearly straight
transverse row, the anterior faces
of the smaller teeth exactly in line,
those of the two larger teeth slightly
more posterior ; i'^ and P small,
sub-equal, the shafts compressed,
the cutting edges rounded, simple ;
P higher than the others, somewhat
resembling the lower canine in form,
its apex curved outward and back-
ward beyond level of smaller teeth,
its postero-external surface with two
concavities separated by a trenchant
ridge ; lower incisors subterete, the
Nat. size. crown of i^ obscurely and unsym-
metrically bilobed, that of the others
simple ; cross section of «\ about one-third that of i^. 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
ovitward. 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 ; pwi^ small, single
rooted, its crown area about equal to that P, its form essentially
like that of succeeding tooth, though with cusp relatively less
developed ; j>nt^ and pm^ successively lai'ger, two- rooted, the
crown area of pj/i-' 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 p«i- elliptical, that of
pm^ with inner margin bulging inward behind middle, though
without developing a definite inner lobe ; pnij and pm^ much like

Fig. 75.
Luti-a Intra. Teeth.

LUTRA 361

pm- in both size and form, the three teeth cutting against each
other when jaws are closed ; pm^ larger than the other small
jDremolars, 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 jjosterior 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 I'aised 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,
subteretft, 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; eai", 22. For cranial and dental measurements
see Tables, pp. 362, 363.

Specimens examined. — -Tliirty-three, from the following localities : —

Ireland : Londonderry, 1 (type of rocnsis) ; Ahascragh, Co. Galway, 1.

England : Pembrokeshire, 1 ; River Stoiir, Dorsetshire, 1 ; Rugby,
Warwickshire, 1; Norfolk, 1; no exact locality, 1.

Norway : Egersund, Stavanger, 5.

Sweden : Jockmock, Lappmark, 1 (Stockholm) ; Gnesta, Soderman-
land, 1 (Stockholm) ; Skaue, 1 (Stockholm) ; no exact locality, 2.

France: St. Gilles, Gard, 1 (Lataste) ; Etupes, Doubs, 1 (Mottaz).

Germany : Southern Germany, 2.

362

CARNIVORA

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364

CARNIVOEA

Austria-Hdngaey : Karlsbad, Bohemia, 1 ; Csehtelek, Bihar, Hun-
gary, 1.

Italy: Torriglia, Liguria, 1 (Lataste).

Spain : Castrillo de la Reina, Burgos, 3 (B.M. and U.S.N.M.) ; Galicia, 1 ;
Seville, 5.

Bemai-ks. — 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,

Zoological Society's

57. 12. 14. 4.

Londonderry, Ireland.

Museum. (Typ

le of L. roensis

{Aliss A. Moody.)

0!

?iiby).

6 St.

Ahascragh, Galway.

Purchased (Row-
land Ward).

5. 7. 10. 1.

9, 3 juv. St.

Pembrokeshire, Wales.

Purchased (Row-
land Ward).

5. 5. 21. 1-4.

S.

Rugby, Warwickshire,

Dr.T.S. Townsend

9. 12. 16. 1.

England.

(c & P).

$st.

R. Stour, Dorset.

J. C. Mansel Pley-
deU (c & p).

98. 5. 13. 1.

1.

England.

99. A.

4 c5, 1 9,

Egersund, Stavanger,

K. H. Schaanning

11.6.3.14-18.

skulls.

Norway.

(c).

ad., juv.

South Germany.

Dr. A. Gunther (c).

59. 9. 6. 62-63.

6.

Cs6htelek, Bihar, Hun-

Hon. Mrs. N. C.

10. 9. 14. 1.

gary.

Rothschild (p).

1.

Karlsbad, Bohemia.

Lord 0. Russell

(c & p).
Rev. Saturio Gon-

57. 2. 14. 1.

6,9.

Castrillo de la Reina,

8. 7. 7. 16-17.

Burgos, Spain.

zalez (c).

1 juv.

Galicia.

Dr. V. L. Seoane
(c & P).

94. 6. 18. 1.

skull.

Seville.

Lord Lilford (p).

74. 10. 7. 2.
76. 3. 4. 1.

9, juv. ■

Seville. (Dr. A. Ruiz.)

Lord Lilford (v).

95.9.4.10-11.

skull.

Seville.

Col. L. H. Irby
(c & P).

2. 11. 16. 1.

Sub-Family MUSTELINE.
1835. Mustdinfe Swainson, Nat. Hist, and Classif. Quadr., p. vii.

Geofi'rapliical distribution. — Northern hemisphere, south into
northern Africa, the Malay Archipelago, and northern and
western South America ; in Europe west to Ireland.

Character.^. — Dentition highly trenchant, upper carnassial
without crushing surface other than a small concave 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

MARTES 365

form but never greatly flattened and never very robust, the
rostrum always at least as long as broad ; external form slender,
the legs usually rather short ; feet digitigrade ; toes partly
webbed ; tail varying in length, slender or bushy, never con-
spicuously muscular.

jReviarJcs. — As here understood the sub-family Mustelinse
contains the three genera, Martes, Mustela and Vormela, all of
which occur in Europe.

Genus MARTES Pinel.

1792. Martes Pinel, Actes Soc. d'Hist. Nat., Paris, i, p. 55 {M. domestica

Pinel = M. foiiia Erxleben).
1820. Martes Nilsson, Skaucl. 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, Siiugethiere Deutschlands, p. 211.
1911. Martes Thomas, Proc. Zool. Soc, London, p. 139, March, 1911.

Type species. — Mn-stela martes Linnieus.

Geographical distrihutioyi. — Noi'thern hemisphere from the
limits of tree growth south to the Mediterranean, the Malay
Archipelago, and the central United States ; in Europe west to
Ireland.

Characters. — Skull narrow, moderately high (depth of brain-
case much more than half mastoid breadth), the dorsal profile
moderately curved, the zygomatic arches not specially wide-
spreading, and po.storbital 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
bullae 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^, c t], pm i* m ^~- = 38 ; cutting edges of five small

.i-y 1-1 -*■ 4-4' 2-2 ^ o <::>

premolars (2 upjjer 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

CARNIVOEA

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
cat-like 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 Maries 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 MARTES.

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. latinorum^ p. 373.

Third upper premolar with crown evenly bicon-
vex ; width of inner lobe of upper car-
nassial barely half that of trenchant
portion of crown ; greatest diameter of
upper molar noticeably less than length
of outer border of carnassial ; fur usually
coarser and less soft, and throat-patch more
whitish (Beech Martens).
Condylobasal length of skull in adult male,
76 to 79 mm. ; pale throat patch always
much encroached on by brown of sur-
rounding parts, occasionally obliterated

(Crete) M. bunites, p. 380.

Condylobasal length of skull in adult male,
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) ill. foina, p. 374.

General hue of upper parts drab M.f. foina, p. 375.

General hue of upper parts wood-brown .. M.f. mediterranean p. 380.

MARTES MARTES LinnaBUS.

(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 gi'owth.

Diagnosis. — Third upper premolar with outline of crown.

MARTES 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 ; upi^er molar large, its greatest diameter about
equal to outer length of carnassial, the metacone more than half
as large as paracone, the outer border of the crown usually
notched ; general colour yellowish brovvn, the throat patch

Fig. 76.
Maries 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

CARNIVORA

well defined, completely naked, communicating with upper lip
by narrow median line. Feet densely furred throughout in
winter, the pads bare in summer ; palm with deeply divided,
trilobed, heart-shaped tubercular mass at bases of median digits,
a small round pad at base of thumb, and a lai-ger lound 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
ajDOut 25 mm. long at middle of back in winter, shorter in
summer, the longer hairs reaching about 40 mm. Mammse : 4.

Colour. — The colour varies considerably with season and
climate, but is usually characterized by rich yellowish brown and
blackish tints, the legs, tail and underparts darker than the
back, the head noticeably lighter than the body, the throat and
fore part of chest yellowish, and interramia, chin and muzzle
dusky. Claws light horn-colour.

Skull. — As compared with that of the European species of
Mustela the skull of Mnrtes martes is characterized by a general

Fig. 77.
Martes inartes. 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

MARTKS 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 bullae
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 bullae
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

Fia. 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 lA 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 hannilars short,
slightly everted.

Teeth. — In proportion to size of skull the teeth are larger
than in the members of the other genera of Mustclimx. occurring
in Europe ; transverse diameter of upper molar much more than
half width of palate between molars. Upper incisors forming a
straight row separated at each side from canine by a space about
equal to transverse diameter of i^ and i"^ together ; crowns strongly
compressed, the antero-poscerior diameter fully twice transverse
diameter, the anterior face convex, the posterior ccmcave with

faintly indicated cingulum ; f- slightly
larger than i^, P 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^ implanted in jaw behind
level of the other two ; crowns ob-
scurely bifid, the outer lobe smaller
than the inner ; crov/n area of z\
j.j(j 79 about one-third that of the com-

Martes martes. Teeth. Nat. size, pressed i^ Or the subterete i^. 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^ and i^
together), subterete, the height of the single ill-developed cusp
scarcely equal to diameter of crown in pm^, much less in pm^.
Other premolars, except upper cai^nassial, two-rooted, compressed,
the outline of crown triangular when viewed from the side, with
apex slightly in front of middle of tooth. Cx'own area of p)m'^
about three times that oiprn^ ; outline of ci'own 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 jmi^ somewhat longer than that oi pm"^,
but its area much greater owing to widening of strongly convex
inner side ; cusp essentially as in pwi^ though somewhat higher.
Second lower premolar about equal in size to pwi'^, but axis of
shaft more anterior in position ; pm.j essentially like jira?2, but
cusp with faintly indicated anterior longitudinal ridge, and
posterior ridge with a slight nodule or rudimentary basal cusp
(sometimes absent) ; piii^ noticeably larger than jnii.^ and with a
well developed secondary cusp at middle of posterior border of
main cusp. Upper carnassial ( pm*) long and narrow, the width
of crown just behind internal lobe less than half length along-
outer border, the internal lobe robust, its diameter in line of
tooth-row nearly or quite equal to width of trenchant portion of
crown, its axis nearly perpendicular to main axis of tooth, its cusp
well developed, about as high as that of pra^, and separated from
main cusp by deep concavity ; main cusp robust, its height
contained aVjout If times in length of crown, its axis slanting
a little backward, its anterior border v/ith 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 A times as long as broad, the sectorial portion of the
tooth consisting of a much distorted triangle, the paraconid
forming anterior extremity of tooth, its anterior border nearly
perpendicular, its posterior commissure meeting anterior commis-
sure of the higher protoconid at an abrupt angle, the two together
acting in opposition to the angled commissure of upper carnassial ;
metaconid reduced to a subterete postero-internal process on
base of protoconid, its area scarcely one-sixth that of larger
cusp ; crushing portion of crown slightly broader than trenchant
portion, its outline sub-circular with slightly raised edge, this
edge forming a noticeable though low 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 oiiter and
inner portions ; cingulum low but evident, especially around
inner margin of crown.

Bemarks. — The well-known pine marten is so strongly
characterized as to require no special comparisons with any other
European species except Maries foina (see account of latter).
Two geographical races are currently recognized, though their
status is by no means clear.

Martes martes martes Linmeus.

1758. \^Mustcla'\ martes Linnseus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1816. M\tistela] syliesiris Oken. Lehrb. d. Naturgescli., iii, pt. 2, p. 1020

(Renaming of martes).
1827. M[artes] vulgaris Griffith, Cuvier's Anim. Kingd., v, p. 123

(Reuamiug of martes).
1847. Martes sylvatica Nilssou, Skand. Fauna, i, 2nd ed., p. 41 (Renaming

of martes).
1857. Micstela martes Blasius, Saugethiere Deutschlands, p. 213.

1910. Mustela martes Trouessart, Faune Mamm. d'Europe, p. 72.

1911. [Martcsl martes Thomas, Proc. Zool. Soc. London, p. 139, March,

1911.

Tjipe locality. — Vicinity of Upsala, Sweden.

Geographical distribution. — Europe north of the Mediterranean
region, from Ireland eajjtward into Asia.

Diagnosis. — Throat patch cream-buff or slightly more yellow.

Colour. — Upj)er 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 buffy 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, Stavauger, 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

?.

Kenmaro, Kerry, Ire-
land.

E. Dodson (c).

94. 3. 27. 1.

1.

Kerry.

J. H. Gurney (p).

72. 1. 10. 1.

1.

Scotland. {McLcmj.)

Hargitt Collection.

8G. 9. 9. 1.

St.

Keswick, Cumberland,
England.

W. n. Wilson (c).

9G. 11. 6. 1.

6.

Cockermouth, Cum-
berland.

W. R. Wilson (c).

97. 1. 15. 1.

S.

Pontrilas, Hereford-

Mrs. St. John A.

85. G. 10. 1.

shire.

Matthews (p).

S.

Ludford Park, Here-
fordshire.

H. J. Bailey (c & p).

0. 2. 23. 1.

2.

Wales (1S28).

Earl Cawdor (p).

61. C. D.

2 (?, 1 9,

Egorsund, Stavanger,

K. H. Schaanning

11.6.3.19-21.

skulls.

Norway.

(c).

2 skulls.

Southern Germany.

Dr. A. Giinther (c).

188i, 1299f.

6.

Bohemia.

Lord 0. Russell
(c & P).

50. 12. 23. 1.

Martes MARTES LATiNORUM Barrett-Hamiltoii.

1904. Mustcla martes latinorum Barrett-Hamilton, Anu. and Mag. Nat.

Hist., 7th ser., xiii, p. 389, May, 1904 (Nurri Mountains, Sardinia).
1910. Mustela martes latinorum Trouessart, Faune Mamm. d'Europe,

p. 72.

Type localiti/. — Nurri Mountains, Sardinia.

Geograpliical distrihution. — Mediterranean region (Italy, Sar-
dinia, Balearic Islands).

Diagnosis. — Throat patch bufi-yellow ; general colour lighter
than in true martes.

Colour. — Upper parts a rather light brown, the longer hairs
between the raw-umber and mai's-brown of Ridgway, 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 butf ; 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
irom 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

(CARNIVOEA

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.)

G. Barrett-Hamilton 11. 1. 2. 29.

E. Cavendish Taylor 5. 5. G. 8.

(!■)■
E. N. Buxton (p). 95. 4. IG. 1.

{Type of subspecies.)
0. Thomas (p). 1. 3. 6. 1.

0. Thomas & R. I. 0. 7. 1. 43.
Pocock (c & p).

1.

I\Iilau.

1.

9.
9.

Nurri Mountains, Sar-
dinia.
jMancor, Majorca, Bale-
aric Islands. (Riutort.)
San Cristobal, Minorca.

MARTES FOINA Erxleben.
(Synonymy under subspecies.

Central and southern Continental
coast eastward, and from the

KiG. 80.

Larger cheek-teeth of Marten
mai-tes (a) and M.foina (0).
Nat. size.

Geographical distribution.-
Europe, from the Atlantic
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
as in M. martes, its gx-eatest diameter less
than outer length of carnassial, the meta-
cone less than half as large as paracone ;
external form as in M. martes, but fur
of less fine quality ; colour usually more
greyish or drab than in 31. martes and seldom with the rich
brown tints of the related species, the throat-patch never strongly
tinged with yellow. Mammae : 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 3Tustela erminea and distinctly
less elevated than in Martes martes. Interorbital region and
rostrum wider than in Magics 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

Bemnrhs. — 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.

Martes foina foina Erxleben.

1777. [Mustela] foina Erxleben, Syst. Regni Anim., i, p. 458 (Germany).
1792. Martes domestica Pinel, Actes Soc. d'Hist. Nat., Paris, i, p. 55 (France).
1801. Mltistela] f[oina] alba Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 759 (Thiiringen, Germany).
1857. Mustela foina Blasius, Saugetbiere Deutscblands, p. 217.
1869. Mustela martes var. fagormn Fatio, Fauna Vert. Suisse, i, p. 318

(Sweden) : name wrongly attributed to Linnseus, Syst. Nat., i,

12th ed., p. 67.
1910. Mustela foina Trouessart, Faune Mamm. d'Europe, p. 72.

Type locality. — Germany.

Geographical distribution. — Range oi" the species except
southern Spain.

Diagnosis. — Longer hairs of back tipped with sepia, the
genei'al 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 insulting 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, IMeurthe-et-Moselle, 2.

Germany : Ingelheim, Rheinhessen, 2 ; Nuremberg, Bavaria, 1
(U.S.N.M.) ; southern Germany, 7 (skulls).

Switzerland : Geneva, 1 (Mottaz) ; Vall6e-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.IM.).

Italy : Near Rome, 1.

376

CARNIVORA

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380

CARNIVORA

Geeecr: Mt. Vuno, Cepbalonia, 1 (colour normal; size apparently
rather small).

?, juv. Manonville, Meurthe-et- Lord Lilford (p). 95. 9. 5. 1-2.

Moselle, France.
6. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 17.

Germany.
6. Ingelheim, Rheinhessen. G. Barrett-Hamilton 11.1.2.102.

7 skulls. South Germany. Dr. A. Giinthcr (c). 1299, a-g.

2 6 juv. Schaffhausen, Switzerland. 0. Thomas (p). 2. 8. 4. 22-23.

{E. H. Zollikofcr.)

6. Oberrich, St. Gallen. 0. Thomas (p). 2. 8. 4. 21.

(E. H. ZoUikofer.)

6. Silos, Burgos, Spain. Rev. Saturio Gonza- 8. 7. 7. 21.

lez (c).

1. Rome, Italy. E, Cavendish Taylor 5. 5. 6. 7.

1. Mt. Vuno, Cephalonia, J. I. S. Whitaker (p). 8.10.1.9.
Greece. (C. Mottaz.)

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 w^ith a light
yellowish brown, the general hue of upper parts lighter, yellower
and less drab than in trnQ 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.

Bemarks. — 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 (c & p). 98. 3. 18. 1.

(Type of subspecies.)

MARTES BUNITES Bate.

1899. Mustela foina Iciicolachnfia Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., iv, p. 383 (Not of Blanford).
1906. Mustela foina hunites Bate, Proc. Zool. Soc. London, 1905, ii, 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

MUSTELA 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 folna 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 gi'eatly 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.

Sliidl and teeth. — The skull and teeth resemble those of
Martoi 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.

6, 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, R.N. (c). 99. 6. 13. 2-3.

Genus MUSTELA Linnaeus.

(Synonymy under sub-genera.)

Type species. — Mustela erminea Linn<ieus.

Geographical distrihution. — 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 — , pm ^~

* 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 CARNIVORA

m l^l = 34 ; upper carnassial as in Maries, 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 jjosterior heel crossed by a longitudinal
trenchant ridge ; other teeth essentially as in Maries ; external
form slender, the muzzle obtuse, the ears low and rounded, the
legs short, the tail variable in length and in quality of hair, but
never so bushy as in Maries.

Bemarhs. — The genus Mustela, tlie most widely distributed of
the strictly terrestrial Mmielinse, 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 x-epresented in Europe.

KEY TO THE EUROPEAN FORMS OF MUSTELA.

Brain-case broad, the mastoid width decidedly
greater than distance between basiou and
palation ; auditory bullae triangular in outline

(Sub-genus Putornis, Polecats) M. piitorius, p. 419.

Underfur usually buffy grey (Central Europe) M- P- putorius, p. 423.

Underfur usually yellowish (Southern Spain) M. ^j. aureolus, p. 425.

Brain-case narrow, the mastoid width less than
or about equal to distance between basion and
palation ; auditory bullse not triangular in
outline.
Rostrum flattened above ; inner margins of
auditory buUss strongly divergent pos-
teriorly ; tail bushy ; habits semi-aquatic

(Sub-genus Lutreola, Minks) M. lutreola, p. 415.

Rostrum convex above; inner margins of auditory
bullae 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. hibernka, -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).

383

Brown of sides not encroaching on yellowish
or whitish of underparts, the lino of
demarcation straight ; upper lip wholly

or partly white M. erminca, p. 385.

Region 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. e. crmi>u'a,Tp. 387.

Eegiou 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) ^t- e- neinx, 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 coat normally assumed ;
condylobasal length of skull in
adult male seldom over 50 mm. ;
in adult female seldom over
45 mm. ; upper carnassial not
enlarged and thickened (Conti-
nental Europe from southern
Sweden to the Alps and Pyrenees) M. v. xstiM, 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. e. staUUg, 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
ill. afncana) 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 CARNIVOEA

Brown of sides noticeably encroaching on
yellowish or whitish of underparts,
the line of demarcation irregular

(Malta, Azores) M. africana, 112.

Brown of sides not encroaching on yellowish
or whitish of underparts, the line of

demarcation straight (Crete) M. gaUntJiias, 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) ill. nivalis, p 401.

Winter pelage often white ; size small, hind
foot of males, 29 to 34 mm. (Northern

and central Europe) ^f. n. nivalit', 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. boccamela, 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. iherica^ p. 407.

Sub-genus MUSTELA Linnaeus.

1758. Mustela Linnseus, Syst. Nat., i, 10th ed., p. 45 (Type by tautonymy

M. erminca Linnseus).
1829. Ardogalc Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,

p. 30 [erminea).
1829. Ictis Kaup, Entw.-Cresch. u. Natiirl. Syst. Europ. Thierw., i, p. 85

(vulgaris = nivalis). Not of Schinz, 1824-1828.
1841. Gale Wagner, Schreber's Siiugthiere, SuppL, ii, p. 234 {vulgaris =

nivalis).
1857. Fcatorius Blasius, Saugethiere Deutschlands, p. 219 (part).
18G5. Neogale Gray, Proc. Zool. Soc, London, p. 114 {brasilicnsis, aiireo-

ventris and xanthogenys).
1871. Mustelina, Bogdanow, Ti)V4. Ouiq. EciecTB. KasaiU'K. JHiiBepc. I, Mem. i,

p. 167.
1899. Emnustela Acloque, Fauna de France, Mammiferes, p. 62 [vulgaris

and erminea).
1911. Mustela Thomas, Proc. Zool. Soc. London, p. 138, April, 1911,

Type species. — Mustela erminea Linnseus.

Geocjrapliical distribution. — ISTorthern 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 bullae not triangular, the inner and
outer borders nearly parallel ; posterior border of pm^ shearing

MUSTELA 385

against anterior border of pjjig ; other small premolars not
capable of trenchant action.

BemarJcs. — The sub-genus Musfela 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 LinnfGUS.
(Synonymy under subspecies.)

Geographical distribution. — ■ Europe from the Arctic coast
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 bullee 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 wliitish rim.

External characters. — General form long and slender, the
legs short, the body cylindrical, the neck long and nearly as
thick as body, tail considerably longer than outstretched hind
leg. Ear short but appearing distinctly above fur of head, its
outline evenly rounded, both outer and inner surface densely
clothed with short hair ; muzzle rather broad and short, the
nostril pad sharply defined, entirely naked, separated from upper
lip by a narrow hairy area. Palm and sole completely furred in
winter, the tubercles bare in summer ; on both palm and sole
there is a trilobed, heart-shaped tubercular mass at base of
median digits and a small round pad at base of inner digit ; on
palm there is an additional posterior rounded pad near wrist.
Fur dense and soft, the longest hairs on back about 10 mm.
in winter, less in summer ; tail rather closely haired, the pencil
full, usually wider than basal portion of tail, its longest hairs
about half as long as vertebras ; fur turning white in winter
except in warmer parts of range. Mammie : 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 underf ur and bases of the longer hairs much paler ; uuder-
parts, inner surface of legs and upper surface of feet whitish or
yellowish in strong contrast, the line of demarcation between

2 c

386

CARNIVORA

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

Fig. si.
M'ustela erminea. 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
Mastela 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. puforius, 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.pHtorius. 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 fl;it, 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
M. putorius.

Teeth. — Except for their smaller size the teeth are essentially
like those of Mustela putorius. Enamel of lower canine not
rugose. Second upper premolar and third lower premolar more
compressed than in the larger animal, and with base of cusp
shorter, so that there is a slight but evident concavity or
flattening at both anterior and posterior base. Upper carnassial
with inner lobe projecting more forward than in M. putorius, so
that its anterior border extends nearly to level of that of outer
margin of tooth. Upper molar and posterior lower molar as in
Mustela putorius, but m^ with metacone more reduced, and m.2
with crown area distinctly smaller than that of heel of carnassial.

Mustela erminea erminea Linnaeus.

1758. [Mustela] erminea Linnaeus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1792. Mlzistela] erminea hybcrna Kerr, Anim. Kingd., p. 181 (name applied

to the northern true ermine).
1816. M[;ustcla] herminea Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1026

(Renaming of erminea).
1827. [Mustela erminea] /3 maculata Billberg, Synopsis Faunae Scandinavise,

p. 8 (Scandinavia).
1857. Fcetorius erminea Blasius, Saugethiere Deutschlands, p. 228 (part).

2 c 2

388

CARNIVORA

1877. Patorius erminea Coues, Pur-bearing Animals, p. 109 (part).
1910. Patorius [Ictis) ermineus Trouessart, Faune Mamm. d'Europe, p. 78
(part).

Type locality. — Upsala, Sweden.

Geograpldcal distribution. — Scandinavian Peninsula, except
extreme south of Sweden ; eastern limits of range unknown.

Characters. — Region extending from front of rostrum to
and including postorbital processes relatively broad and short;
size below maximum for the European races : condylobasal length
of skull in adult male, 43 "4 to 48 "6 mm.; mandible in adult
male, 23 to 26*4 mm. ; basal portion of tail (in summer pelage)
usually lighter below than above.

Measurements. — Hind foot in each of two males from the
vicinity of Christiania, Norway, 43 mm. ; in a female from the
same region, 35 mm. Male and female from Nordre Fron, Lower
Gudbrandsdal : hind foot, 43 and 34 mm. Two males from
Lappmark, Sweden : hind foot, 44 and 46 mm. Adult female
from Upsala, Sweden : hind foot, 36 mm. Adult male from
Stockholm, Sweden : hind foot, 44*6. For cranial measurements
see Table, p. 392.-"-

Specimens examined. — Twenty-five, from, the following localities : —
Sweden: Karesuando, Lappmark, 1 (U.S.N.M.); Wilhelmina, Lapp-
mark, 1 (U.S.N.M.) ; Jemtland, 1 (U.S.N.M.) ; Upsala, 1 (U.S.N.M.) ;
Stockholm, 2 (U.S.N.M. and Stockholm).

Norway : Aker, near Christiania, 2 ; Brekkebygden, Trondhjem, 1 ;
Nordre Fron, lower Gudbrandsdal, 2; Marestuen, Fillefjeld, Bergen, 1,
Egersund, Stavanger, 13.

ItemarJcs. — The Scandinavian form of Mustela erminea is
distinguishable from the other European races by its slightly
smaller size and by the broader rostro-froiital 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 noi'mally assumed throughout the range of this form.

6, 6 juv. Aker, Christiania, Nor- Christiania Museum 93. 3. 1. 2, 4.
way. (e).

?. Brekkebygden, Trond- Christiania Museum 93. 3. 1. 3.

hjem. (e).

6, 9. Nordre Fron, Gudbrands- Christiania Museum 93. 3. 1. 5-6.
dal. (e).

9. Marestuen, Fillefjeld, E. R. 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
tj^pe, 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 jESTIVA KeiT,

1792. M[ustela] erminea sestiva Kerr, Anim. Kingd., p. 181 (Germany;

based on Schre bar's pi. 137a).
1820. Mustela erminea major Nilsson, Skaud. Fauna, i, p. 34 (Carlskrone,

Blekinge, Sweden).
1857. FcBtorius 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 AIjds and Pyrenees.

Characters. — Region extending from front of rostrum to and
including postorbital processes relatively narrow and long ; size
larger than in 31. erminea erminea ; condylobasal length of skull
in adult male, 47 to 51 mm. ; mandible in adult male, 25 to
29 mm.

Measurements. — Two young adult males from Pic du Midi,
Hautes-Pyrenees, France : head and body, 235 and 240 ; tail,
82 and 85 ; hind foot, 44 and 44 ; ear from meatus, 17 and 17.
Three males from Strass, near Burgheim, Bavaria : head and
body, 265, 270 and 280 ; tail, 90, 100 and 102 ; hind foot, 44,
47 and 48. Two females from the same locality : head and body,
240 and 240 ; tail, 80 and 90 ; hind foot, 38 and 40. Average
and extremes of eight adult males from the vicinity of
St. Gallen, Switzerland: head and body, 271 (251-292); tail,
104 (94-111); hind foot, 47 (4.5-50). Three adult females
from the same region : head and body, 226 (218-242) ; tail, 89
(85-98) ; hind foot, 38 • 8 (37 ■ 6-41 ).

Specimens examined. — Seventy-nine, from the following localities : —

Denmark : No exact locality, 2 (skulls, Copenhagen) ; Amager, 1
(skull, Copenhagen) ; Faaborg, Fyn, 4 (skulls, Copenhagen) ; Vestervig,
Jutland, 1 (skull, Copenhagen).

Holland : Texel Island, 1.

Prance: Pic du IMidi, Hautes-Pyr6n6es, 2; Manouville, Meurthe-et-
Moselle, 1.

Germany: Neustadt, Wied, 1 ; Ingelheim,Rheinhesseu,8 ; near Magde-
burg, 2 ; Bleiche, Saxony, 5 ; Moritzburg, Saxony, 1 (U.S.N.M.) ; Rudol-
stadt, Thiiringen, 3 ; Marxheim, Bavaria, 4 ; Strass, near Burgheim,
Bavaria, 5; no exact locality, south Germany, 6 (skulls).

Austria-Hungary: Csallokoz-Somorja, Pressburg, Hungary, 1.

Switzerland : Frete-de-Sailles, Vaud, 1 (U.S.N.M.) ; Frutigeu, Bern, 1
(U.S.N.M.); Rorbas, Bern, 1 (U.S.N.M.); Ziirich, 1 (U.S.N.M.); St.
Gallen, 4 (U.S.N.M.) ; Degersheim, St. Gallen, 5 (U.S.N.M.) ; Rorschach,
St. Gallen, 1 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 ; Untervatz, Grisons, 8
(B.M. and U.S.N.M.) ; Priittigau, Grisons, 1 (U.S.N.M.) : Vals, Grisons, 1
(U.S.N.M.); Obersaxen, Grisons, 1 (U.S.N.M.); Pontresina, Grisons, 1
(U.S.N.M.) ; Scanfs, Grisons, 1 (U.S.N.M.) ; Poschiavo, Grisons, 1 (U.S.N.M.);
Ems, Grisons, 2; Osogna, Ticino, 1 (U.S.N.M.).

Remarks. — The ermine of central Europe differs from that of
Scandinavia in slightly greater size and in the narrower rostro-
frontal region of skull. The white winter pelage is normally
assumed throughout the ranore of the form.

390 CARNIVORA

As shown by both external and cranial measurements the two
specunens from the Pic du Midi are unusually small. It is
possible that they represent a peculiar Pyrenean race.

6. Texel Id., HoUaud. J. L. Bonhote (c & 8. 10. 20.' 1.

P).
2 6. Pic du Midi, Hautes Pyr6- O. Thomas (p). 8. 9. 1. 52-53.

n6es, France. (A. Bobert.)
i. Manonville, Meurthe-et- Lord Lilford (p). 11.1.1.139

Moselle. {Lomont.)
9. Untervatz,Grisons,Switzer- 0. Thomas (p). 4. 4. 5. 33.

land. {E.H.Zollihofer.)
J, 9. Ems, Grisons. (£. H. ^o«i- 0. Thomas (p). 2.8.4.25-26.

hofer.)

5. Neustadt, Wied, Germany. Lord Lilford (p). 11. 1. 1. 138.

{Schneider.)
5 (?, 3 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 22-28.

2. Magdeburg, Prussia. Dr. W. Wolterstorff 0. 2. 8. 6-7.

4 6. Bleiche, Saxony. [H. Tor- Lord Lilford (p). 11. 1. 1. 140-
nitz.) 143.

6. Marxheim, Bavaria. Lord Lilford (p). 11.1.1.145.

{Wolterstorff.)

5. Strass, Burgheim. {Kur- Lord Lilford (p). 11. 1. 1. 144.

hitz.)

6. South Germany. Dr. A. Giinther (c). 59.9.6.64-69.
1. Csallokoz - Somorja, Press- Budapest Museum 94.3.1.1.

burg, Austria-Hungary. (e).

MusTELA ERMINEA STABiLis Barrett-Hamilton.

1904. Putorius ermineus stabilis Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xiii, p. 394, May, 1904.

1910. Putorius [Ictis) ermineus stabilis Trouessart, Faune Mamm. d'Europe,
p. 79.

Ti/pe locality. — Blandford, Dorset, England.

Geographical distrihution. — Mainland of Great Britain.

Diagnosis. — Size slightly greater than in M. erminea sestiva :
condylobasal length of skull in adult male, 49 to 52 • 4 mm. ;
mandible in adult male, 27 to 30 mm. ; teeth usually larger than
in the continental races, a peculiarity especially noticeable in the
upper carnassial ; colour averaging slightly darker above and
less strongly yellow below than in the continental forms (though
in extreme instances the underparts are between buff and straw-
yellow, quite as in the brightest continental specimens) ; tail
usually without distinct lighter area below.

Measureraents. — External measurements of type (adult
female) : head and body, 244 ; tail, 105 ; hind foot, 43 ; ear from
meatus, 22. Another adult female from the type locality : head
and body, 245 ; tail, 102 ; hind foot, 43; ear from meatu.s, 21.
Two adult males from Bamstead, Surrey : head aud body, 270
and 278 ; tail, 119 and 119; hind foot, 47 and 49. Two adult
males from Wales : head and body, 274 and 280; tail, 120 and

391

120; hind foot, 48 and 48*6. Adult male from Farr, Daviot,
luverness : head and body, 254 ; tail, 110 ; hiud foot, 48 "5.

Specimens examined. — Seventy-four, from the following localities : —

Scotland : Thurso, Caithness, 1 ; Sutherlaudshire, 1 ; Cromarty, 4 ;
Annadale, Skye, 2 ; Farr, Daviot, Inverness, 1 (Wilson) ; Stockbriggs,
Lanark, 9 ; Wyseley, Dumfries, 2 ; Ecclefechan, Dumfries, 1 (U.S.N.M.).

Wades : North Wales, no exact locality, 1 ; Nannerch, Flintshire, 2 ;
Vaynol, Carnarvonshire, 2 ; Usk, Monmouthshire, 2 ; St. Brides, Pembroke-
shire, 1 ; St. Fagan's, Cardiff.

England : Riding-ilill-on-Tyne, Northumberland, 1 ; Westmoreland, 1 ;
Doncaster, Yorkshire, 2 ; Leeds, Yorkshire, 1 (U.S.N.M.) ; Gainsborough,
Lincolnshire, 1 ; Derbyshire, 1 ; Sandringham, Norfolk, 3 ; Lowestoft,
Suffolk, 1 ; Bury St. Edmunds, Suffolk, 1 ; Priswell, Cambridgeshire, 8 ;
Rugby, Warwickshire, 1 ; Graftonbury, Hereford, 3 ; Tring, Hertford, 2 ;
Felden, Hertfordshire, 1 ; Banstead, Surrey, 2 ; Laseley Park, Guildford,
Surrey, 1 ; Buckland, Somerset, 2 ; Blandford, Dorset, 3 ; Eversley,
Hampshire, 1; Whitechurch, Hampshire, 1; Selborne, Hampshire, 1;
Staplehurst, Kent, 1 ; Horsham, Sussex, 2 ; Mayfield, Sussex, 8.

Bemarhs. — In the British stoat the change to the white
winter coat does not take place so regularly and completely as in
the continental forms. This is particularly true in central and
southern England. Apart from this character, of problematic^]
value, the animal is distinguishable by its heavy teeth. A few
small though apparently well developed skulls from the north
of Scotland indicate the possible existence there of a local form
somewhat resemblin" true erminea.

9.

3 9.

6, 9.

6.

3 6, St.

Thurso, Caithness, Scot-
land (ill?s. J. Edis).
Cromarty.

Annadale, Skye.
Sutherlaudshire.
Stockbriggs, Lanarkshire.

W. R. Sherrin (p). 9. 1. 9. 1.

W.R.Ogilvie-Grant

(c & p).
Dr. Hastings (c&p).
W. Paterson (c&p).
E.R.Alston (c&p).

3 S, 9. Stockbriggs, Lanarkshire. E. R. Alston (c & p).

9. Stockbriggs, Lanarkshire.

i, 9. Nannerch, Flintshire,

Wales.
2 9. Vaynol, Carnarvonshire.
2 i. Usk, Monmouthshire.

St. Brides, Pembroke-
shire.
1. St. Fagans, Cardiff,
st. Derbyshire, England,
st. Sandringham, Norfolk.

2 st. Sandringham, Norfolk.

9 st. Lowestoft, SuSolk.

st. Bury St. Edmunds, Suf-
folk.
3 (J, 4 9. Friswell, Cambridgeshire.
S. Rugby, Warwickshire.

E.R.Alston (c&p).
A. Richardson

(c & p).
J. E. Harting (p).
J. S.Phillips (c&p).
Hon. C. Edwardes

(c & p).
J. Cording (c).
Index Museum.
H.M. King Edward

VII. (p).
H.M. King Edward

VII. (p).
F. S. Worthiugton

(c & p).
T. Harcourt-Powell

(c & p).
W. Farren (c).
E.E.Austen (c&p).

11.1.3.175-177.

11. 1. 3. 395-396
8. 11. 26. 1.
79. 9. 25. 16, 18,

20.
79. 9. 25. 14, 15,

17, 19.
79. 9. 25. 84.
11. 1. 3. 192-193.

11. 1. 3.194-195.
87. 8. 6. 1-2.
90. 12. 5. 1.

90. 6. 19. 1.
94. 4. 8. 1.
96. 4. 13. 1.

96. 11. 24. 1-2.

98. 2. 15. 1.

87. 2. 28. 1.

4. 5. 3. 1--S.
11. 1. 8. 181.

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MUSTELA 397

6, 2 ?. Graftonbury, Hereford- W. E. de Winton (p). 11. 1. 3. 178-180.

shire.
2 ?. Tring, Hertfordshire. Hon. N. C. Roths- 11. 1. 3. 182-183.

child (p).

6. Felden, Hertfordshire. 11. 1. 3. 184.

1. Nr. Hounslow, Middle- W. Phillips (p). 60. 1. 23. 2.

sex.
?. Guildford, Surrey. G. Dalgleish (c &p). 4.4.3.1.

S st. Horsham, Sussex. Sir E. G. Loder, 6. 7. 14. 1.

Bart, (c & p).
(5,9. Mayfield, Sussex. C.H.B.Grant(c&p). 11. 1.3. 186-187.

6. Selborne, Hampshire. Dr. E,. Bowdler 2. 11. 29. 1.

Sharpe (c & p).
9. Whitchurch, Hampshire. W. E. de Winton 11. 1. 3. 185.

(c & p).
S, 9. Buckland, Somerset. R. Hooper (c & p). 11. 1. 3. 190-191.

9. Blandford, Dorset. J. C. Mansell Pley- 98. 5. 13. 2.

dell (c & p). [Type of subspecies.)
2 9. Blandford, Dorset. W. T. Blanford' 11.1.3.188-189.

(c & p).

MuSTELA ERMINEA RIOIN^ Miller.

1904. Putorius erminea Barrett-Hamilton, Ann. Scottish Nat. Hist., v,

p. 203, October, 1904.
1907. Putorius erminea ricinx Miller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 395, November, 1907.
1910. Putorius ermineus ricinie Trouessart, Faune j\Iamm. d'Europe, p. 80.

Type locality. — Islay House, Islay Island, Scotland.

Geographical distribution. — Islands of Islay and Jura, off coast
of south-west Scotland.

Characters. — Size a little less than in M. erminea stahilis and
about equal to that of M. e. sestiva : condylobasal length of skull
in adult male, 47 to 50 mm. ; mandible in adult male, 27 to
29 mm. ; zygomatic arches unusually wide-spreading ; posterior
extension of palate relatively wider than in any of the other
known forms.

Measurements. — External measurements of type (adult male) :
head and body, 254 ; tail, 105 ; hind foot, 43 ; ear from meatus,
22. Two other males from the type locality : head and body,
270 and 270; tail. 111 and 114; hind foot, 47 and 46. Two
adult females from the type locality : head and body, 231 and
234 ; tail, 95 and 105 ; hind foot, 38 and 39. Two adult males
from Jura : head and body, 220 and 225; tail, 120 and 125;
hind foot, 45 and 46*5.

Specimens examined. — Thirteen, six from Jura and seven from Islay
(B.M. and Cambridge).

Remarks. — ^While readily distinguishable from the large true
stahilis of England, the Islay and Jura stoat will probably prove
to be more nearly related to the smaller form occurring on the

398

CARNIVORA

maiiilaacl of Scotland, when the status of the Scotch animal is
more clearly understood.

1. Islay, Scotland. H. Morrison (c & p). 7. 10. 9. 1.

{Type of subspecies.)
6, 2 9. Islay, Scotland. H. Morrison (c & p). 11. 1. 3. 196-198.

MUSTELA HIBERNICA Thoma,s and Barrett-Hamilton.

1895. Putorius liihcrnicus Thomas and Barrett-Hamilton, The Zoologist,

3rd ser., xix, p. 125, April, 1895 (Ireland).
1895. Putorius hihernicus Thomas, The Zoologist, 8rd ser., xix, p. 226,

June, 1895 (Isle of Man).
1900. '\_M%istcla] hibernica Schulze, Zeitschr. fiir Naturwissensch., lxxiii,

p. 221.
1910. Putorius {Ictis) hihernicus Trouessart, Faune Mamm. d'Europe, p. 80.

Type locality. — Enniskillen, Co. Fermanagh, Ireland.

Geographical distribution. — Ireland and the Isle of Man.

Diagnosis. — Like Miistela erminea but slightly smaller (hind
foot in adult male rarely attaining 50 mm. ; condylobasal length
of skull in adult male, 44 to 50 mm.) ; oar and upper lip without
whitish edging ; brown of sides of body usually encroaching on
yellowish or whitish of underparts, frequently so much so as
to extend completely across throat and middle of belly, the
line of demarcation, except in rare instances, very irregular.

External characters. — The general external charactei^s, includ-
ing the proportion of tail to head and body, and the long,
full pencil, are as in Mustela erminea ; mammfe usually more
numerous : a 2 — 2, i 3 — 3 or 4 — 4 = 1 0 or 1 2.

Colour. — Except in the rare instances in which a partial or
complete white coat is assumed in winter, the actvial colour is
very constant throvighout the year, and such variation as there
is appears to be due chiefly to the greater or less abundance of
the long hairs on back. These hairs are a dark glossy brown
very nearly the bistre of Ridgway. The hairs of the dense
underfur are smoke-grey at base, changing to a yellowish broccoli-
brown distally. From the varying combinations of these elements
different shades of brown result which range from a yellowish
bistre toward isabella-colour, the bistre usually more pure along
median dorsal region and on head, the isabella-colour on sides,
legs and tail. Upper lip and entire ear concolor with dark
area. No dark spot behind angle of mouth. Feet usually dark
but frequently varied with whitish, this colour appearing at tips
of toes and spreading upwards. Tail without evident lighter
area on under side, Underparts varying from a creamy white
to a pale straw-yellow. The line of demarcation between the
dark and light areas is very variable in both form and position.
In the extreme development of the light area the line of demarca-
tion extends along sides in the same position as in Mustela
erminea, except that it passes somewhat further below ear and
eye and ends at or just above angle of mouth. In the extreme

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extension of the dark area the brown forms a band 20 mm. wide
across throat just in front of fore legs, and spreads over whole
median region of belly, so that the white is confined to three
patches, one extending from chin to throat-band, another occupy-
ing chest between front legs and spreading on inner side of
forearm, and the third in region between hind legs and spreading
on inner surface of thighs. Between these extremes eveiy
gradation may be found, including the most irregular arrange-
ments of isolated dark spots in the regions where the dark cross
bands occur.

Skull and teeth. — Except for their slightly smaller size the
skull and teeth are indistinguishable from those of Mustela
erminea. The difference is, however, particularly well marked as
compared with the large British M. erminea stabilis, the nearest
geographical ally.

Measurements. — External measurements of type (adult male) :
head and body, 228 ; tail, 88 ; hind foot, 42 ; ear from meatus,
21. Average and exti^emes of four males from Cappagh, Water-
ford : head and body, 271 (256-283); tail, 111 (103-117); hind
foot, 48*2 (46-51). Average and extremes of three females
from Powerscourt, Co. Wicklow : head and body, 209 (205-221) ;
tail, 78 (77-81) ; hind foot, 37-6 (36-40). For cranial measure-
ments see Table, p. 399.

Specimens examined. — Forty-four, from the following localities : —
Ireland : Carrick, Donegal, 1 ; Colebrooke, Fermanagh, 8 ; Enniskillen,
Fermanagh, 2 ; Clandeboye, Down, 1 ; Glaslough, Monaghan, 1 ; Board-
mills, Down, 2 ; Castle Hamilton, Cavau, 1 ; Mountainstown, Meath, 3 ;
Clonbrock, Galway, 1 ; Woodfair, Galway, 1 ; Carna, Galway, 1 ; Temple-
more, Tipperary, 1 ; Geashill, Kings, 1 ; Powerscourt, Wicklow, 4 ;
Bagenalstown, Carlow, 1 ; New Ross, Wexford, 4 ; Arthurstown, Wex-
ford, 2 ; Cappagh, Waterford, 4 ; Lismore, Waterford, 1.

Isle op Man : Lewaig, Ramsey, 1 ; Santon, 2 ; Tholt-y-Will, Snaefell, 1.

Remarks. — The Irish stoat is strikingly distinct from the
other European species of the sub-genus Mustela, though super-
ficially resembling certain North American members of the
group. It is at once recognizable by tlie combination of black-
tipped, heavily pencilled tail with entirely dark ear and upper
lip. It is also the only European stoat in which the line of
demarcation on sides of body is irregular and in which the dark
colour of sides tends to invade the light ventral area. In
exceptional instances''^ the colour pattern of body is like that of
Mustela erminea. While the colouring of the body and head
suggests that of Mustela nivalis, the animal is evidently a true
stoat, as shown by the heavily-pencilled, black-tipped tail and
the form of the skull.

6 St. Carrick, Donegal, Ireland. Hon. N. C. Roths- 0.5.17.1.

child (p).
3 6. Colebrooke, Fermanagh. Sir D. Brooke (c & p). 11. 1. 3. 199-201.

* In four of the forty-one skins examined.

MUSTELA 401

6. Eniiiskillen, Fermanagh. J. E. Harting (c&p). 95.4.5.1.

(Type of species.)
6 st. Clandeboye, Dowu. Hou. N. C. Kotlis- 0. 5. 17. 2.

child (p).
6. Carna, Galway. Col. J. W. Yerbury 93. 1. 6. 1.

(c & p).
st. Templemore, Tipperary. E. Lynan (c & p). 99. 7. 3. 1.
2 S. Arthurstown, Wexford. G. Barrett-Hamilton 11. 1. 3. 202-203.

(c & P).
?. Ramsay, Isle of Man. P. M. C. Kermode 95. 5. 80. 1.

(c & p).
?. Snaefell, Isle of Man. P. M. C. Kermode 11. 1. 3. 204.

(c & p).
6, 1 al. Santon, Isle of Man. J. C. Bacon (c & p). 1. 5 12. 1-2.

MUSTELA NIVALIS Linnoeus.

(Synonymy under subspecies.)

Geograpliical distribution. — Europe from the Arctic coast to
the Mediterranean (including the Balearic Islands, Corsica,
Sardinia, Sicily, and Malta) and from Great Britain .eastward
into Asia.

Diagnosis. — Size usually less than in Mtisiela erminea and
M. hibernica, and tail usually shorter in proportion to length of
head and body, the pencil small, its longest hairs not half as long
as vertebrae ; skull like that of Mustela erminea, but with rostral
breadth over canines relatively greater, frequently exceeding
interorbital breadth, especially in the larger races in which the
skull approaches that of M. erminea in size ; colour much as in
M. erminea, but brown of upper parts usually (except in
Mediterranean races) encroaching on whitish of belly, the line
of demarcation irregular ; tail with no black except occasionally
a tuft in pencil ; a white winter pelage in colder portions of
range.

External characters. — In general like Mustela erminea, but size
usually less, and tail relatively shorter (scarcely more than one
quarter head and body, except in the forms inhabiting the
Mediterranean region), the pencil thin, tapering, never as wide
as median portion of tail, its longest hairs only one-quarter to
one-third as long as tail vertebne ; palms and soles hairy, the
pads bare in summer in northern forms, always in southern
forms ; ear short but appearing distinctly above fur, its outline
evenly rounded, both inner and outer surfaces densely clothed
with short hairs ; fur as in M. erminea. Mammje : i 4 — 4 = 8.

Colour. — Upper parts a yellowish brown similar to that of
Mustela erminea, but seldom darkening along median dorsal
region or on head, the brown usually extending to or covering
dorsal surface of feet ; underfur throughout the dark regions
paler than distal portion of long hairs ; underparts and inner
surface of legs whitish or yellowish in strong contrast, but line
demarcation low on sides of body and usually irregular, the

2 D

402

CARNIVOEA

brown tending to encroach on white to such an extent as
occasionally to pass completely across chest, and frequently to
form spots and blotches in same region ; upper lip usually with
some white ; a brown spot varying much in size, and sometimes
joined with brown of checjk, usually present behind angle of
mouth ; ear brown like surrounding j^arts, the rim not
whitened ; tail concolor with back, the pencil faintly darker or
occasionally with black hairs, the median ventral region some-
times faintly paler than rest of tail, but never with definite
light stripe. In white winter pelage the entire animal is white
or whitish, the tail sometimes with a few black hairs in pencil.
The change does not take place regularly except in the colder
portions of the animal's range, as in the Alps and central and
northern Scandinavia.

Skull and teeth. — The skull resembles that of Mustela erminea
in all respects except that the interorbital
region is often narrower than the rostrum,
a character most pronounced in the larger
races that approach M. erminea in size.
Postorbital process frequently more promi-
nent than in M. erminea, and zygoma often
less evenly bowed up-ward, its upper border
flattened or sometimes with a distinct median
concavity and posterior widening as in
M. puioriiis. Teeth not distinguishable from
those of Mustela erminea except by their
smaller size.

BemarJcs. — Although the most vai'iable of
the European Mustelidae this species is always
readily distinguishable externally fi'om the
other members of the genus Mustela with
which it is associated by the short thin pencil.
The discrepancy in width between the rostrum and interoi'bital
region, while generally characteristic of the species as compared
with Mustela erminea, is not perfectly constant. It is invariably
well marked, however, in adults of the larger southern animals.
Three races are represented in Europe, a smaller northern and
central form, and two larger subspecies in the Mediterranean
region.

Fig. 82.

Mustela nivalis (male).
Nat. size.

Mustela nivalis nivalis Linnaeus.

1766. [Mustela] nivalis Linnaeus, Syst. Nat., i, 12th ed., p. 69.
1777. [Mustela'] vulgaris Erxleben, Syst. Regni Anim., i, p. 471
(Temperate Europe).
Mustela gale Pallas, Zoogr. Rosso-Asiat., p. 94 (Eenaming of

vulgaris).
Mustela minor Nilsson, Skand. Fauua, i,p. 35 (Renaming of nivalis).
Phitorius] minutus Pomel, Catal. M6th. et Descr. Vert. Foss. Loire,
p. 51 (Neighbourhood of Paris, France).

1811.

1820.
1853.

MUSTELA 403

1857. Fcetorius vulgaris Blasius, Siiugethiere Deutscblands, p. 231.

1869. Fcetorius picsillus Fatio, Faune Vert. Suisse, i, p. 332 (Not of Audubon

and Bachman ; substitute for vulgaris).
1895. Putorius nivalis Tbomas, The Zoologist, Brd ser., xix, p. 177,

May, 1895.
1900. Putorius nivalis typicus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v,' p. 42, January, 1900.
1900. Putorius nivalis vulgaris Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 42, January, 1900.
1908. P[utorius] nivalis var. monticola Cavazza, Ricerche sui" Putorius

nivalis " e sui "Putorius ermineus" d' Italia, Bologna, p. 37 (High

valleys of the Alps).
1910. Putorius (Ictis) 7iivalis, P. (I.) nivalis vulgaris and P. (I.) vulgaris

minutus Trouessart, Faune Mamm. d'Europe, pp. 81-82.

Type localitij. — Province of Vesterbotten, Sweden.

Geographical distributioit. — Europe from the Arctic coast to
the Alps and Pyrenees, and from Great Britain eastward.

Characters. — Size small : hind foot of adult males 29 to 34 mm. ;
condylobasal length of skull in adult males usually less than
41 mm. (36 to 42 mm.).

Measurements. — ^Two adult males from Cromarty, Scotland :
head and body, 211 and 215; tail, 65 and 59; hind foot, 34
and 34 ; ear from meatus, 16 and 15. Two adult females from
the same region : head and body, 161 and 174 ; tail, 39 and 49 ;
hind foot, 24 and 25 ; ear from meatus, 12 and 12. Two adult
males from Guines, Pas-de-Calais, France : head and body, 209
and 212; tail, 60 and 58; hind foot, 31 and 30. Adult male
and female from Porte, Pyrenees-Orientales, France : head and
body, 213 and 187 ; tail, 60 and 54 ; hind foot, 33 and 26. Adult
male and female from Hatszeg, Hunyad, Hungary : head and
body, 196 and 150; tail, 70 and 41; hind foot, 31-4 and 21.
For cranial measurements see Table, p. 408.

Specimens examined. — Ninety-nine, from the following localities : —

Scotland : Thurso, Caithness, 1 ; Cromarty, 2 ; South Sutor, Cro-
marty, 1 ; Black Island, Cromarty, 1 ; Elgin, 1 ; Dava, Elgin, 1 (Wilson) ;
Cortacby, Forfar, 1 (Wilson) ; Stockbriggs, Lanark, 3 ; Lamancha, Peebles,
1 ; Ecclefecban, Dumfries, 2, Wyseby, Dumfries, 1 ; Jardine Hall, Dumfries,
2 (U.S.N.M.).

England : Leeds, Yorkshire, 1 (U.S.N.M.) ; Grimsby, Lincolnshire, 1 ;
Aberia, Merionethshire, 1 ; Pembrokeshire, 4 ; Methwold, Norfolk, 1 ;
Sandringham, Norfolk, 4 ; Suffolk, 2 (skulls) ; Histon, Cambridgeshire, 2 ;
Tring, Hertfordshire, 3 ; Soham Fen, Cambridgeshire, 1 ; Saffron Waldeu,
Essex, 4 ; Southall, Middlesex, 1 ; Eashing, Surrey, 1 (U.S.N.M.) ; Cole-
ford, Gloucestershire, 1 ; Froyle, Hampshire, 1 ; Blandford, Dorset, 1 ;
Poughill, Shalton, Cornwall, 1 ; Stratton, Cornwall, 1.

Norway : Asker, near Christiauia, 1.

Sweden: Stockholm, 1 (U.S.N.M.).

Denmaek : Helsingor, Zealand, 1 (Copenhagen) ; Amager, 1 (Copen-
hagen).

France : Guines, Pas-de-Calais, 2 ; Leguevin, near Toulouse, Haute-
Garonne, 1; Luchon, Haute-Garonne, 1; Porte, Pyr6n6s-Orientales, 4;
Ax-les-Thermes, Ariege, 1 ; Barcelonnette, Basses-Alpes, 1.

Germany; Brunswick, 1 (U.S.N.M.); Konigsberg, 3 (U.S.N.M.);

2 D 2

404

CARNIVORA

Nuremberg, Bavaria, 1 (U.S.N.M.) ; Strass, near Burgheim, Bavaria, 1 ;
Ingelheim, Rheinhessen, 3; South Germany, 4 (skulls).

AusTBiA-HuNGARY : Halda, Arva, Bohemia, 1 ; Hainspach, Bohemia, 4
(U.S.N.M.) ; Csallokoz-Somorja, Pressburg, Hungary, 3 ; Hatszeg, Hunyad,
Hungary, 4.

Switzerland; Thayngen, Schaffhausen, 3 (U.S.N.M.); Andermatt,
Uri, 1 (U.S.N.M.); St. Gallen, 1; Mels, Sfc. Gallen, 1 (U.S.N.M.); Watt-
wil, St. Gallen, 1 (U.S.N.M.) ; Weiern, St. Gallen, 1 (U.S.N.M.) ; Ziiber-
v^angen, St. Gallen, 2 (U.S.N.M.) ; Oberengadin, 1 ; Untervatz, Grisons, 1
(U.S.N.M.).

Remarks. — The weasel of central and northern Europe never
attains the large size often found in the Mediterranean races.
The material at present available is not sufficient to show
whether or not the northern animals are all referable to a
sincjle form.

S.

8 i.
i.

2 i, 9,

Thurso, Caithness, Scot-
land.
Cromarty.

Lamancha, Peebles.
Stockbriggs, Lanark-
shire.

Mrs. J. Edis (p).

11. 1. 3. 300.

G. St. Quentin 11. 1. 8. 205,

(c & p). 207, 432.

J. L. Bonhote (c & p). 11. 1. 3. 433.
E. R. Alston (c & p). 79. 9. 25. 23,

85-86.

Aberia, Merionethshire, G. H. Caton Haigh 11. 1. 3. 443.

6.
(5,9.

Wales.
Pembrokeshire.

Grimsby, Lincolnshire,

England.
Norfolk.
Sandringham, Norfolk.

2 st. Sandringham, Norfolk.

(c & p).
G. H. Mills (c & p). 11. 1. 3. 444-

447.
G. H. Caton Haigh 11. 1. 3. 434.

(c & p).
Dr. C. Hose (c & p). 11. 1. 3. 208.
H.M. King Edward 96. 4. 13. 2. 3.

VII. (p).
H.M. King Edward 96.4.28.14-15.

VII. (p).
Col. E. S. Butler (p).

2. Suffolk.

6. Histon, Cambridgeshire. Dr. S. F. Harmer

(c & p).
9 (albino) Soham Fen, Cambridge- P. Bond (p)
shire.
Saffron Walden, Essex

2 iJ, 2 9.
i.
6.

A. Wris

Southall, Middlesex.
Froyle, Hampshire.

t (c & p).
Wroughton

94. 1. 6. 6-7.
11. 1. 3. 206.

90. 6. 21. 1.

11. 1. 3. 436-

439.
11. 1. 3. 440.

R. C

(c & p).
W. R. Ogilvie-Grant 11. 1. 3. 441.

(c & p).
W. L. S. Loat(G&p). 11.1.3.442.
Christiania Museum 95. 5. 29. 1.

(e)
O. Thomas (c & p)

9. Stratton, Cornwall,

juv. Asker, Christiania, Nor-
way.
2 6. Guines, Pas-de-Calais, O. Thomas (c & p). 94. 6. 6. 10-11.
France.
6. L u c h o n. Haute- O. Thomas (p). 6. 4. 1. 36.

Garonne. (A. Robert.)
6. Leguevin, Haute- 0. Thomas (p). 6.4.1.37.

Garonne. (A. Robert.)
6. Ax-les-Thermes, Ariege. V. Builles (p). 8. 3. 27. 8.

2 <5, 1 9. Port6, Pyr6n6es-Orien- G. S. Miller (c). 8.8.4.166-168.

tales.
9. Barcelonnette, Basses- O. Thomas (p). 8. 8. 10. 44.

Alpes. (C. Mottaz.)

MUSTELA 405

6 juv. Strass, Burgheim, Ba- Lord Lilford (p). 11. 1. 1. 96.

varia. (Korbitz.)

3. lugelheim, Rheinhes- C. Hilgert (c). 8. 11, 2. 29-31.

sen, Germany.

4. South Germany. Dr. A. Giinther (c). 59. 9. 6. 71-78,

122.
8. Csallokoz - Somorja, Budapest Museum 94.3.1.2-4,

Pressburg, 400 ft. (e).
Hungary.
3 6,9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3.2.2.17-19.
sylvania. 3. 11. 8. 19.

MusTELA NIVALIS BoccAMELA Bechstein.

1801. Mustela boccamela Bechstein, Gemeinn. Naturgesch, Deutschlands,

I, 2nd ed., p. 819 (Sardinia).
1869. Putorius vulgaris var. meridionalis Costa, Annuario del Mus. Zool.

della R. Univ. di Napoli, 1865, p. 40 (Southern Italy).
1900. Putorius 7iivalis italicus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 45, January, 1900 (Grezzana, highlands of

Verona, Italy). Type in British Museum.
1900. Putorius nivalis boccamela Barrett-Hamilton, Ann. and jMag. Hist.,

7th ser., v, p. 46, January, 1900.

1900. Putorius nivalis siculus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 46, January, 1900 (Marsala, Sicily). Type in
British Museum.

1901. Mustela (Ictis) dombrowskii Matschie, Sitz.-Ber. GeseUsch. Natur-

forsch. Freunde, Berlin, p. 231 (Siulnita, Roumania).

1905. [Fcetorius piisillus'] major Fatio, Arch. Sci. Phys. et Nat. Geneve,
4th ser., xix, p. 512, May 15, 1905 (Poschiavo, Grisons, Switzer-
land ; see Mottaz, Bull. Soc. Zool., Geneve, i, p. 169, November 15,
1908). Type in Mottaz collection.

1910. Putorius (Ictis) boccamela, P. [I.) nivalis meridionalis, P. (J.) nivalis
dombrowskii, and P. (J.) nivalis siculus, Trouessart, Fauue Mamm.
d'Europe, pp. 81, 83.

Type locality. — Sardinia.

Geofjraphical distrihution. — Italy and coa.st of France as far
as the Department of Var ; island.s of Sicily, Malta and Sardinia :
Corsica ?

Diagnosis. — Similar to Mustela nivalis nivalis, but decidedly
larger, and tail relatively longer ; hind foot of adult males 34, to
41 mm. ; condylobasal length of skull in adult males usually
more than 41 mm. (40 to 46-6 mm.).

Measurements. — Two adult males from Valescure, Var, France :
head and body, 252 and 250 ; tail, 88 and 80 ; hind foot, 37 • 6
and 38 ; ear from meatus, 21 and 19. Adult male from Agay,
Var, France : head and body, 265 ; tail, 86 ; hind foot, 41 ; ear
from meatus, 20. Adult male from vicinity of Genoa, Italy :
head and body, 240; tail, 80; hind foot, 37*6. Young adult
male from near Verona, Italy (type of italicus) : hind foot (dry),
32 • 4. Adult male from Marsala, Sicily : head and body, 250 ;
tail, 90 ; hind foot, 39. A second male from the same locality
(type of siculus) : hind foot, 34 • 2. Adult male from Malta ;

406

CARNIVORA

head and body, 173 ; tail, 62 ; hind foot, 34.
measurements see Table, p. 410.

For cranial

Specimens examined. — Forty-six, from the following localities : —

France. — Valescure, Var, 2 ; Agay, Var, 1.

Switzerland : Somvico, Ticino, 1 (U.S.N.M.) ; Boguo, Ticino, 1
(U.S.N.M.); Buggiolo, Ticino, 1 (U.S.N.M.); San Lucio, Ticino, 1 ;
Poschiavo, Grisons, 4 (U.S.N.M. and Mottaz, including type of major).
These Swiss specimens are intermediate between ioccamela and nivalis,
though nearer the former.

Italy : Padola, Cadore, 1 (Turin), not typical ; Turin, 1 ; near Genoa, 1 ;
near Verona, 3 ; near Florence, 2 (U.S.B.M.) ; near Rome, 2.

Sicily : Balestrate, 2 ; Marsala, 3 ; Palermo, 5 (B. M. and U.S.N.M.).

Sardinia : No exact locality, 4 ; Cagliari, 3.

Malta; No exact locality, 3 ; Ghallis, 4; Bingemma Fort, 1.

Roumania ; Malcoci, Dobrudscha, 1 (U.S.N.M.).

Bemarhs. — Although showing a perplexing amount of indi-
vidual variation'"' the weasel of the central Mediterranean region
must be regarded as distinct from that of central and northern
Europe. A small percentage of southern specimens cannot be
identified with certainty ; but typical fully adult males of
boccamela are strikingly different from the animal occurring
north of the Alps.

6. Valescure, Var, France.
6. Agay, Var.
9. St. Lucio, Ticino, Switzer-
land. {E. H. Zollikofer.)

5. Turin, Italy.

6. Genoa, Liguria.

2 c5, 1 9. Verona (Conte clegli Odcli).

(99.
6. Balestrate, Sicily.
4 6. Palermo.

2 c5, 1 9. Marsala.

G. S. Miller (c).
G. S. Miller (c).
0. Thomas (p).

8. 8. 4. 170.
8. 8. 4. 169.
2. 8. 4. 27.

6, 2 al.
3al.

]UV.

9.

3 (5, 1 9.

9.

Cagliari, Sardinia.

Sardinia.

Sardinia.

IMalta.

IMalta. (Micallef.)

Ghallis.

Bingemma Fort.

E. C a V e n d i s h 5. 5. 6. 11.

Taylor (p).
Lord Lilford (p). 95. 11. 4. 1.

0. Thomas (p). 99. 11. 11. 1-8.

11. 11. 1. Type of italiciis Barr.-Ham.)
J. I. S. Whi'taker (p). 95. 3. 4. 4.
J. I. S. Whitaker (p). 98. 10. 6. 8.

95. 3. 4. 1-3.
J. I. S. Whitaker (p). 95. 3. 4. 5-7.
(95. 3. 4. 5. Tiipe of siculus B.-Ham.)
E. N. Buxton V)- 95. 4. 16. 1-3.
Genoa Museum (p). 86. 11. 8. 1-3.
Purchased (Linnsea, 86. 7. 10. 2.

Berlin).

C. A. Wright (p).
C. A. Wright (p).
C. A. Wright (p).
C. A. Wright (p).

94. 11. 26. 1.

95. 1. 2. 1.
7. 7. 6. 1-4.
7. 7. 6. 5.

* Studied in much detail by Cavazza, Ricerche sui "Putorius nivalis"
e sui " Putorius ermineus " d' Italia (Bologna, Zanichelli), 1908, Studien
iiber die in Italien vorkommenden Wleselarten der Untergattung Arctogale
(Zool. Anzeiger xxxiv, pp. 582-603), 1909, Sulle Donnole e suU' Ermellino
in Italia (Boll. Soc. Zool. Ital. xviii), 1909, and Contributo alia conoscenza
della vita e delle abitudini della Donnola, Putorius nivalis, Linn. (ibid.
xix, pp. 65-82), 1910.

407

MusTELA NIVALIS IBERICA Barrett-Hauiilton,

1900. Putorius nivalis ihericus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 45, January, 1900. Type in British Museum.
1905. Putorius hibericus Seabra, Bol. da Direcc;.. Gen. da Agricultura,

Lisboa, viii, no. 2, p. 69.
1910. Putoriiis nivalis ihcrica Trouessart, Faune ]\Iamm. d'Europo, p. 83.

Type locality. — Seville, Spain.

Geographical distribution. — Iberian Peninsula and Balearic
Islands.

Diagnosis. — Intermediate in size between Mustela nivalis
nivalis and M. n. hoccamela, though nearer the former (hind foot
in adult males, 30 to 35 ; condylobasal length of skull in adult
male, 39 to 43 mm.) ; colour of upper parts a buflfy clay-colour,
not so dark as in the other races.

Measurements.- — Adult male and female from the neighbour-
hood of Silos, Burgos, Spain: l^ead and body, 225 and 184;
tail, 59 and 53; hind foot, 27 and 21 "8 ; ear from meatus, 15
and 13. Two adult males from Majorca, Balearic Islands : head
and body, 191 and 250 ; tail, 79 and 90 ; hind foot, 32 and 35.
Two adult males from Dehesa de Valencia, Spain : head and
body, 264 and 278 ; tail, 62 and 62 ; hind foot, 32 and 34.
Adult male from Seville, Spain (type): liind foot (dry), 31 •4.
For cranial measurements see Table, p. 411.

Specimens examined. — Twenty-seven, from the following localities : —
Spain : Pajdres, Leon, 1 ; Panticosa, Huesca, 1 ; Arrechavaleta,
Vitoria, 2 ; near Burgos, 1 ; Silos, Burgos, 1 ; Castrillo de la Reina,
Burgos, 2 ; La Granja, Segovia, 1 ; Villalba, Madrid, 2 ; Barracas, Castel-
lon, 2 ; Dehesa de Valencia, 4 ; Seville, 3 (including type) ; Jerez, Cadiz, 1 ;
Muro, Majorca, Balearic Islands, 1 ; Inca, Majorca, Balearic Islands, 2 ;
Mahon, Minorca, Balearic Islands, 1.
PoETUGAL : Alcochete, 1.

BemarJcs. — The Iberian weasel appears to be more nearly
related to Mustela nivalis nivalis than to M. n. hoccamela. It is
distinguishable from both of these by its paler colour.

9. Pajdres, Leon, Spain.

[N. Gonzalez.)
6. Panticosa, Huesca.

(N. Gonzalez.)

2 9. Arrechavaleta, Vitoria.

{N. Gonzalez.)
6, 9 juv. Burgos.

1 al. La Granja, Segovia.

5, 9. Villalba, Madrid.

[N. Gonzalez.)
6. Barracas, Castellon.

{N. Gonzalez.)

3 6. Dehesa de Valencia.

[N. Gonzalez.)

6, 9. Bormujo, Seville.

{Dr. A. Buiz.)

0. Thomas (p).
0. Thomas (p).
0. Thomas (p).
G. S. Miller (c)

8. 2. 9. 54.
8. 2. 9. 58.
8. 2. 9. 59-60.
8. 8. 4. 50-51.

M. de la Escalera (c). 8. 7. 30. 8.
0. Thomas (p). 8. 2. 9. 50-57.

0. Thomas (p).
O. Thomas (p).
Lord Lilford (p).

8. 2. 9. 55.
8. 2. 9. 51-53.
95. 9. 4. 12-13.

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CARNIVORA

9. Seville. {Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 10.

{Type of sub-species.)

6. Jerez. A. Chapmau (c&p). 9. 4. 24. 1.

1. Alcochete, Portugal. Lisbon Museum (e). 3.11.13.1.
c5. Muro, Majorca, Balearic 0. Thomas (p). 1. C. 1. 5.

Islands. {M. Riutort.)

2. Inca, Majorca. 0. Thomas and R. I. 0. 7. 1. 7-8.

Pocock (c & p).

9. Mahon, Minorca. 0. Thomas (p). 0. 8. 13. 1.

MUSTELA AFRICANA Desmarest.

1818. Musfela africana Desmarest, Nouv. Diet. d'Hist. Nat., xix, p. 376

(" Africa ").
1895. Putorius africanus Thomas, Proc. Zool. Soc. London, p. 128 (Malta

and Egypt).
1900. Putorius nivaUs africanus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 47, January, 1900.
1908. ? P[uforius~} nivalis var. corsicamis Cavazza, Ricerche sui "Putorius

nivalis" e sui "Putorius ermineus " d' Italia, Bologna, p. 37

(Corsica).
1910. Putorius (Ictis) nivalis suhpalmatus Trouessart, Faune Mamm.

d'Europe, p. 85.

Type locality. — Said to be Africa (no exact locality stated),
but more probably the Azores, as the type came to Paris through
the museum of Lisbon, Portugal.

GeograpMcal distribution. — Egypt, Malta, San Thome, Azores.

Diagnosis. — Like Mustela nivalis hoccamela but larger (hind
foot of male, 44 to 49 mm., condylobasal length of skull, 48 to
50 mm.), tail relatively longer (nearly half as long as head and
body) and brovi^n of sides usually spreading on undei'parts, the
line of demarcation very irregular.

Colour. — The colour does not differ noticeably from that of
3Iustela nivalis hoccamela. Upper parts wood-brown, in some
specimens clear and light, in others dark and strongly tinged
with raw-umber, the head usually concolor with body but
sometimes distinctly darker. Tail concolor with back above,
not so dark below, the pencil usually darker than rest of tail
and occasionally blackish. Underparts pale cream-buff or buffy
white, the light area on chest and belly much encroached on by
brown of sides, which occasionally passes completely across,
leaving only a few spots of white. Muzzle and upper lip entirely
brown, or with a trace of white in front.

Skull and teeth. — The skull resembles that of Mustela nivalis
hoccamela so closely that I can detect no characters others than
its larger size by which it can be distinguished. In size it fully
equals ordinary skulls of M. erminea ; but the much broader
rostrum as compared with interorbital region serves at once to
distinguish it. The teeth show no tangible characters by which
they may be differentiated from those of M. n. hoccamela except
that the carnassial is more robust and the small premolars in

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414

CAKNIVORA

both Jaws tend to be more crowded, so that their crowns are
usually set more obliquely to axis of tooth-row.

Measurements. — Two adult males from Terceira, Azores : head
and body, 255 and 266 ; tail, 105 and 116; hind foot, 44 and
44 ; ear from meatus, 19 and 18. Adult from Malta : hind foot,
44; ear from meatus, 19,

Specimens examined. — Five, from the following localities: — Malta, 1 ;
San Thome, 1; Terceira, Azores, 2; St. Michaels, Azores, 1.

Hemarhs. — This African species probably owes its presence in
Malta and on the Azores to introduction by man. So far as can
be judged from the description, the Corsican weasel to which
Cavazza has applied the name corsicanus, is the same animal.

1. Malta. C. A. Wright (p). 75. 4. 6. 1.

1. San Thome. Hon. W. Rothschild 4. 1. 1. 5.

6. Terceira, Azores. Hon. W. Rothschild 3. 6. 5. 25.

(W. B. Ogilvie-Grant.) (p).
6. Terceira. D. H. Chassereau 4. 3. 15. 1.

(c & p).
?. St. Michaels. Major P. A. Chaves 5. 1. 19. 1.

(c & p).

MUSTELA GALINTHIAS Bate.

1906. Putorius nivalis gaUnthias Bate, Proc. Zool. Soc. London, 1905, ii,

p. 319, April 5, 1906.
1910. Putorius (Ictis) nivalis gaUnthias Trouessart, Faune Mamm.

d'Europe, p. 14.

Type locality. — Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Similar to Mustela africana, but with brown of
sides not encroaching on underparts, the line of demarcation
straight. Skull not known.

Specimens examined. — Two, both from Crete.

Bemarhs. — Further material may show that the Cretan
weasel is not distinct from Mustela africana. The two specimens
now known are alike in colour and different from any of the
African or other skins of M. africana examined. Though with-
out skulls or measurements they evidently represent a weasel
quite equal to the African animal in size.

6. Malaxa, Crete. Miss D. Bate (c). 5. 12. 2. 23.

1. Crete. Miss D. Bate (c). 5. 12. 2. 24.

{Type of species.)

415

Sub-Genus LUTREOLA Warner.

1841. Lntrcola Wagner, Schreber's Stiugthiere, SuppL, ii, p. 239 (lutreola).
1843. Vison Gray, List. Spec. Mainni. Brit. Mus., p. 64 {vison).
1857. Fcetorius Blasius, Siiugetbiere Deutschlands, p. 219 (part).
1871. Hydromuslcla Bogdanow, T|)\4. 06m. Ectcctb. Ka.siiHCK. yiiiiBcpc I,
Mem. I, p. 167.

Tijj)e species. — Mustela lutreola Linnaeus.

Geographical distribution. — Northern portion of northern
hemisphere from eastern Germany to the Atlantic coast of Nortli
America.

Characters. — Form moderately slender ; tail bushy ; fur modi-
fied for aquatic life ; skidl without noticeably projecting mastoid
processes ; auditory bulh« sub-triangular ; premolars more prehen-
sive than in Ptitoriiis ; inner lobe of upper carnassial functioning
against apex of pHt^ ; anterior border of pm^ shearing against pos-
terior border oipm., ; point of p?w- opposed to posterior border of j^m.^.

Remarks. — Though well characterized in its extreme develop-
ment, as represented by the type species and the American forms,
the sub-genus Lutreola grades insensibly into true Mustela through
such Asiatic members of the genus as Mustela sihiriea, M. cani-
gula and others. So complete is this intergradation that it may
well be questioned whether the name Lutreola should be allowed
to remain in use. The only typical Old World species ranges
westward into south-western France.

MUSTELA LUTREOLA Linnseus.

1766. [_Mustela] lutreola Linnaeus, Syst. Nat. i, 12th ed., p. 66 (Finland).
1777. \Lutra'] minor Erxleben, Syst. Regni Anim., i, p. 451 (Renaming of

lutreola).
1792. M\^^stela'\ Lutra fulva Kerr, Anim. Kingd., p. 173 (Renaming of

lutreola).
1889. ? [Mustela lutreola'] var. alba De Selys-Longcbamps, Etudes de

Micromamm., p. 46 {nomen mulum).
1857. Foetorius lutreola Blasius, Siiugetbiere Deutschlands, p. 234.
1863. ? Putorius alpinus Og6rien, Hist. Nat. du Jura, iii, p. 59 (highest

portions of the Jura).
1879. Lutreola europn'a Homeyer, Zool. Garten, xxvi, p. 184, June, 1879

(Substitute for lutreola).
1910. Putorius (Lutreola) lutreola Trouessart, Faune Mamm. d'Europe,

p. 75.

Tyjje locality. — Finland.

Geographical distribution. — Northern Asia, westward to Fin-
land and south-western France, southward to eastern Roumania ;
limits of range not known. European distribution less extensive
than formerly.

Diagnosls.--Geneva\ characters as in the sub-genus Lutreola ;
colour throughout a rich dark brown, the region about the mouth
whitish, the tip of tail blackish ; head and body about 350 to

416

CARNIVORA

400 mru., condylobasal length of skull ranging about from 57
to 65 mm.

Colour. — Back, sides and underparts a rich glossy dai-k brown,
neai'ly the mars-brown of Ridgway, the hairs of underfur between
hair-brown and mars-brown, darker at tips than at base, the longer
hairs raw-umber, showing a faint greyish cast in certain lights ;
head slightly darker than back, especially on region in front of
eyes ; interramia and lips to level of middle of muzzle pad whitish ;
feet sepia ; tail similar to body, but becoming blackish at tip.

Fig. 83.
Mustela lutreola. Nat. size.

Skull. — The skull is about as lai*ge as in females of Mustela
putorius. In form it is narrower and less deepened than in
M. putorius, especially in interorbital region, the dorsal profile

417

Observations.

Teeth slightly worn.

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Number. Sex.

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3270 ,, 6

3678 „ 6

3679 ,, 9

Locality.

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418

C'ARNIVORA

less convex throughout and in particular less bent downward
anteriorly ; depth of rostrum at level of front of carnassial
barely equal to distance from carnassial to front of premaxillary
instead of appreciably greater ; orbit larger and more widely
open posteriorly than in 31. putorius, a peculiarity due chiefly to
the very slight development of postorbital angle on upper margin
of zygoma ; auditory bulla moderately inflated, irregularly almond-
shaped in outline, its width barely lialf its length, the meatus
slightly projecting but not tubular.

Teeth. — The teeth are less robust but more prehensive than
those of MusteJa jmtorius, the former peculiarity best shown by
the upper carnassial, the latter by the premolars seen in profile
when jaws are closed. In the upper carnassial the transverse
diameter behind inner lobe is distinctly less than in 31. putorins,
the anterior cingulum cusp is better developed, the main cusp is
lower, and the posterior cutting edge has a concave or nearly
horizontal posterior portion rather longer than the abruptly
sloping anterior portion (in M. putorius the cutting edge when
unworn slopes directly from apex of main cusp to base of
posterior cusp). Small premolars fitting closely against each
other when jaws are shut ; posterior lower premolar cutting
against anterior surface of upper carnassial and posterior surface
of pni^, instead of being widely removed from carnassial and
fitting distinctly inside of pm'^.

3Ieasurements. — Adult male and female from Laval, Mayenne,
France (skins) : head and body, 400 and 350 ; tail, 140 and 130 ;
hind foot, 59 and 53. Adult from Malcoci, Dobrudscha,
Roumania (skin): head and body, 380; tail, 130; hind foot, 52.
For cranial measurements see Table, p. 417.

Specimens examined. — Six, from the following localities : —
Feance : Vinsac, Gironde, 2 (Lataste) ; Soulac, Gironde, 1 (Lataste) ;
Laval, Mayenne, 2 (Lataste).

EouMANiA : Malcoci, Dobrudscha, 1 (Schliiter).

Sub-Genus PUTORIUS Cuvier.

1817. Putorius Cuvier, Regiie Animal, i, p. 147 (Type by tautonymy

Miistela putorius Linnfeus).
1840. Fceiorius Keyserling and Blasius, Wirbelthiere Europas, p. 66

[pxdoriios).
1857. Foetorius Blasius, Siiugethiere Deutschlands, p. 219 (part).
1877. Cynomionnx Coues, Fur-bearing Animals, p. 99 [Putorius nigripes

Audubon and Bachman).

Type sjjecies. — 3Iustela putorius Linnst^us.

Geographical distrihition. — Mediterranean i-egion and central
Europe from Great Britain eastward into China ; central United
States ; exact limits of range not known.

Characters. — Form moderately slender ; ta,il bushy ; fur not
modified for aquatic life ; skull with noticeably projecting

MUSTELA 419

angular mastoid processes ; auditory bu Has distinctly triangular in
outline ; inner lobe of upper carnassial functioning against
paraconid of lower carnassial ; interrelationships of teeth essen-
tially as in Miistela, but small premolars more robust and less
trenchant.

Bemarls. — The sub-genus Pntorivs, with three species in the
Old World and one in America, appears to be the most sharply
defined of the three groups into which the genus 3Ii(.sfela is now
divided. No intermediate forms are known connecting it with
either Liifrcula or true Musfela, and if none are fovmd the group
will probably be regarded as a distinct genus. Two of the Old
"World species occur in Europe, but only one comes within the
scope of the present work.

MUSTELA PUTORIUS Linnfeus.

(Synonymy under subspecies.)

(ileoiirfqyhlral distr'dmtlon. — Europe from Great Britain east-
ward, and from the Mediterranean coast to central Scandinavia.
Eastern limit of range not known.

Diagnosis. — Skull with wide brain-case and conspicuously
angular projecting mastoid region, the mastoid breadth decidedly
greater than distance from basion to palation ; auditory bullae
slightly longer than broad, triangular in outline ; size nearly
equal to that of the martens (head and body of adult male about
400 mm.) ; form slender, fur coarse, tail rather bushy, tapering :
colour bufty grey or yellowish overlaid with black, the under-
parts darker than back.

External characters. — Form nnich as in the meinbers of the
sub-genus Miistela, but appearing less attenuate on account of
the longer fur and somewhat bushy tail. Underfur rather dense
and woolly, about 2.5 mm. long in winter, 15 mm. in summer ;
longer hairs coarse and loose, very different in quality from the
underfur and not concealing it, their length at middle of back
about 40 mm. in winter, somewhat less in summer ; tail about
one-third as long as head and body, rather bushy, its underfur
evident, the longer hairs about 30 mm. in length at middle,
decreasing in length toward tip so that the tail tapers noticeably
from basal half outward. Feet moderately long, more robust
than in true Mustela, the plantar and palmar tubercles larger,
bare in summer, furred in winter, the soles always thickly
furred ; claws of front feet about 6 mm. in length, rather
strongly curved and partly retractile, those of hind feet about
4 mm. long, slightly curved, not retractile. Head somewhat
flattened, muzzle rather wide ; eyes small ; ear low, rounded,
never conspicuously overtopping fur, its height from crown
seldom exceeding 15 mm. Mamma" : / 4 — 4 = 8.

Colour. — Feet, tail, chest and intercrural region blackish, rest
of body with long black hairs producing a clouded effect over

2 E 2

420 CARNIVORA

buffy grey cjr yellowish under fur, the two elements of the
pelage distinct and not intimately blending, the black more pre-
dominant in winter than in summer, the exact shade of underf ur
ranging from a light buffy grey with pale ecru-drab bases to
the hairs to a clear ijuff-yellow throughout ; middle of belly
usually like back and sides and conspicuously lighter than inter-
crural region and chest ; chin, interramia and upper lip to
muzzle pad whitish tinged with buffy grey, this light area, some-
what dulled by admixture of brownish hairs, extending upward
posteriorly as a broad band running between ear and eye
(usually not reaching latter) and across forehead, where it curves
slightly forward ; face in front of light band dark brown to
muzzle ; ear whitish grey, the orifice covered by a tuft of dark
brown hairs.

Skull. — The skull is robust and strongly built, rather fiat,
with short rostrum, short, broad brain-case, and long, nearly
parallel-sided postorbital constriction. Dorsal profile nearly flat
from slightlv overhanging lambda to between postorbital
processes, then abruptly bent downward at an angle of about
30^, the slanting portion less than half as long as the horizontal.
Occiput squarely truncate, the condyles sometimes visible from
above, sometimes concealed by the lambdoid crest. Brain-case
ovate when viewed from above, its outline obscured by the
strongly developed angularly projecting mastoid region ; sagittal
crest low or obsolete except in fully adult individuals, joining
lambdoid crest at middle of a shallow median concavity in out-
line of latter ; lambdoid crest well developed, slightly over-
hanging. Floor of brain-case with slight median longitudinal ridge
and shallow lateral depressions ; a transverse depression in front
of lip of foramen magnum. Auditory buihe moderately inflated,
especially along inner edge, about three-quarters as wide as long ;
the general outline triangular with antei'ior border shortest, the
meatus not tubular ; anterior extremity of bulla not in contact
with hamidar, and separated from foramen ovale by an oblique
ridge which is marked oft' from glenoid process by a deep
conspicuous notch, the distance between bulla and foramen
about one-third that between foramina. Postorbital region form-
ing a conspicuous nearly parallel-sided neck between brain-case
and postorbital processes, a character peculiar to this species and
to Lutra Intra among Eui-opean Mnstelidse. Interorbital region
slightly broader than rostrum, moderately convex longitudinally,
a little more convex laterally ; postorbital processes short but
distinct, directed slightly backward. Rostrum short and heavy,
the width across canines e(|ual to distance fi'om gnathion to
anteorbital foramen. Orifice of anteorbital foramen moderately
large, over antei-ior root of carnassial, and wholly beneath
anterior rim of orbit, the plate dividing foramen from orbit fully
as wide as foramen. Zygomata compressed, moderately spreading,
widest apart posteriorly, the orbital process short but distinct,

MUSTELA 421

followed l)y a broad, shallow concavity and a rather abrupt
posterior convexity. Palate moderately broad, its width between
molars equal to a little less than twice greatest diameter of

Pig. 84.
Mustela puturius. Nat. size.

molar ; incisive foramina small, broadly ovate, oblique, in front
of canines and very close behind alveoli of incisors, the minute
median foramen between or behind posterior border ; posterior

422

CARNIVORA

extension of palate longer than broad, emarginate posteriorly ;
mesopterygoid fossa parallel sided except for its anterior
convexity, its total length nearly equal to that of posterior
extension of palate, the slender hamulars hooked outward.
Mandible robust, its lower margin faintl}" convex from broad,
ill-defined angvdar process to middle ; coronoid process high,
triangular, the anterior border slightly the longest.

Teeth. — Relatively to size of skull the teeth are robust and
strong, though not unusually so. Incisors as in Maries vmrte.%
but posterior shelf of i^ and P less developed, and P less strongly
contrasted in size with the two other teeth. Canines lower and
more robust than in Martes and enamel of lower tooth usually
less rugose. Anterior premolar both above and below (pm'^ and
pm.2) small, single-rooted, the crown area about eqvial to that of
outer upper incisor, the ill-developed cusp at front of crown, its
height scarcely equal to width of tooth. Middle ujjper premolar

(pm^) two-rooted, compressed, the
outline of crown flattened-elliptical
when viewed from below, triangular
when viewed from the side, the
apex slightly in front of middle, the
height a little less than length ;
cingulum very slightly developed,
though rather evident at anterior
and posterior borders of crown.
Posterior lower premolar (p'»4)
essentially like pjiii-^ but larger and
relatively higher, its crown less com-
pi'essed posteriorly. Middle lower
premolar (pm^) intermediate in size
and form between prii^ and piH^.
Upper carnassial essentially like that
of Martes foina, but with inner lobe usually even more reduced,
anterior border of main cusp less oblique, and posterior cusp
slightly larger. Lower carnassial with crown nearly 2}, times as
long as wide, the main trenchant portion of the tooth formed by
the paraconid and protoconid, which are essentially similar to
those of Martes, though a little more compressed ; no trace of
metaconid ; posterior heel relatively small, its width distinctly
less than that of base of protoconid, its surface crossed by a low
but evident longitudinal trenchant ridge, the outer surface of
which sheers against inner surface of paracone and metacone of
upper molar. Second lower molar terete, flat, with slight median
longitudinal ridge, its crown area about equal to that of heel of
carnassial. Upper molar irregularly paudurate in outline, the inner
section slightly larger than the outer, the constriction evident
though not deep ; protocone low, terete, at middle of inner
section of tooth ; paracone and metacone small, confluent, the
paracone slightly larger than protocone, the metacone much

Fig. 85.
Muxtela 2J"tortus. Teeth.

MUSTELA 423

smaller, the outer border of crown slightly notched in region
between cusps.

Bemarks. — In certain conditions of pelage Mustela piitorius
bears a superficial resemblance to Marten foina, though the two
animals may always be distinguished by the different form of the
ears. Two geographical races ai-e at present recognized, though
their status is far from well understood.

The ferret, Maries furo Linnteus,* though usually assumed to
be a domesticated variety of Mustela putorius, appears to be
related to the Asiatic M. eversmanni Lesson. f This is shown by
the deeply constricted postorbital region, the less triangular,
more inflated auditory bullse, and the smaller size of the carnassial
teeth both above and below, characters by which the few skulls
of ferrets that I have exandned may at once be distinguished
from those of Mustela putorius, but which render them practically
identical with those of the Eastern form.

Mustela putorius putorius Linnpeus.

1758. [Mustela] putorius Linnseus, Syst. Nat., i, 10th ed., p. 46.
1785. Mustela iltis Boddaert, Elenchus Animalium, p. 87 (Renaming of
jputorius).

1797. Mlusteld] furo-putorius Link, Beytrage zur Naturgesch., i, p. 83

(Based on a melanistic putorius popularly regarded as a hybrid
between /;t9-o djndi putorius).

1798. Viverra foctens Thunberg, Beskrifning pa Svenske Djur, p. 15

(Renaming oi piUorius).

1801. Ml^ustelci] plutorius] albus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, 1, 2nd ed., p. 782. Not Mustela foiyia alba I.e. p. 759
(Thiiringen, Germany).

1827. P\_utorius\ vulgaris Griffith, Cuvier's Anim. Kingd., v, p. 120 (Re-
naming of putorius).

1839. ? [Mustela putorius'] var. flavicans De Selys-Longchamps, Etudes de
Micromamm., p. 145 (nomen nudum).

1839. ? [Mustela putorius] var. vison De Selys-Longchamps, Etudes de
Micromamm., p. 145 [nomen nudum).

1843. Putwius fontidus Gray, List Spec. Mamm. Brit. Mus., p. 64 (Re-
naming of putorius).

1857. Fcetorius putorius Blasius, Saugethiere Deutschlands, p. 222.

1868. Putorius infectus Ogerien, Hist. Nat. du Jura, iii, p. 59 (Substitute
iov putorius) .

1904. Putorius putorius Barrett-Hamilton, Ann. and INIag. Nat. Hist.,
7th ser., xiii, p. 389, May, 1904.

1904. Putorius putorius manium Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xiii, p. 390, May, 1904 (Teufen, Appenzell, Switzer-
land). Type in British Museum.

1910. Putorius putorius and P. putorius manium Trouessart, Fauue
Mamm. d'Europe, pp. 76, 77.

Type locality. — Sweden.

Geograpliical distribution. — Central Europe from central Scan-

* Syst. Nat., i, 10th ed., p. 46.
t Man. de Mamm., p. 144, 1827.

424

CARNIVORA

dinavia to northern Spain and the Mediterranean coast,
westward to Great Britain ; eastern Hmit of range not known.

Diagnosis. — -Underfur bufFy grey or pale buff, rarely if ever
decidedly yellow.

Measurements. — ^Two adult males from Aberystwyth, Cardigan-
shire : head and body, 403 and 415; tail, 163 and 190; hind
foot, 61 and 62 ; ear from meatus, 27 and 30. Adult female
from the same locality: head and body, 360; tail, 140; hind
foot, 53; ear from meatus, 23. Two adult males from Ingelheim,
Rhei'nhessen, Germany : head and body, 400 and 440 ; tail, 1 60
and 170 ; hind foot, 63*6 and 65; ear from meatus, 30 and 31.
Adult female from the same locality : head and body, 348 ;
tail, 132; hind foot, 51*4; ear from meatus, 26. Adult male
from Teufen, Appenzell, Switzerland (tyjae of manium Barrett-
Hamilton) : head and body, 408 ; tail, 145 ; hind foot, 61 ; ear
from meatus, 25. Adult male from Merse, Siena, Italy : head
and body, 420; tail, 150; hind foot, 63. For cranial measure-
ments see Table, p. 426.

Specimens examined. — Sixty-seven, from the following localities : —

Scotland : Glencassley, Sutherlaudshire, 1 ; no exact locality, 8.

England: No exact locality, 2 ; Oundle, Northamptonshire, 1; BuUnose
Coppice, Northamptonshire, 1 ; Wishech, Cambridgeshire, 1 ; Swaffam Fen,
Cambridgeshire, 1; Cardiganshire, G; Aberystwyth, Cardiganshire, 6.

Sweden: No exact locality, 1 (skull; U.S.N.M.) ; Skane, 1 (Stockholm).

France: Scientrier, Haute-Savoie, 2 (Mottaz).

Gekmany : Ingelheim, Rheinhesscn, 6; Ummerstadt, Thiiringen, 9;
Magdeburg, Saxony, 2 ; Marxhcim, Bavaria, 1 ; Niesky, Silesia, 1.

Switzerland : Teufen, Appenzell, 1 (type of maninm Barrett-Hamilton) ;
St. Gallen, 2; Urniisch, St. Gallen, 1 (U.S.N.M.); Bedano, Ticino, 1
(U.S.N.M.).

Italy : Near Florence, 3 (B.M. and Mottaz) ; Merse, Siena, 1 ; Rome, 1 :
Viterbo, Rome, 5.

Spain : Near Burgos, 1 ; Palacios de la Sierra, Burgos, 1.

JRemarJcs. — ^The Swiss form, to which the name manium has
been given, appears to be identical with true Mustela j^utorins.*

6. Glencassley, Suther- Mrs. Flower (p). 92. 10. 21. 2.

landshire, Scotland.

6, 9, G juv. Scotland. Purchased (Rowland Not registered.
St. Ward).

1. Oundle, Northampton- Lord Lilford (p). 94. 6. 11. 4.

shire, England.

?. Swaffam Fen, Cam- R. Metcalfe (p). 11. 1. 3. 397.

bridgeshire.

6 St. Aberystwyth, Cardi- Hon. N. C. Roths- 0. 10. 12. 1.
gaushire, Wales. child (p).

c5, 2 9 St. Aberystwvth. Hon. N. C. Roths- 1.5.22.5-7.

child (p).

* The names Putorius typiis P. Cuvier and P. communis G. Cuvier
mentioned by Trouessart as synonyms of M. indorins I have been unable
to verify.

MUSTELA 425

6, 9. Aberystwyth. {Riiskin- G. Barrett-Hamiltou 11.1.2.105-106.

Butter field.) (p).

6 skeleton. Boneath, Cardigan- W. E. de Vinton 0. 10. 9. 1.

shire. (c & p).

2. England. Purchased. 39. 7. 15. 2-3.

3 i, 9. Ingelheim, Rhein- C. Hilgert (c). 8. 11. 12. 18-21.
hessen, Germany.

6,9. Ingelheim, Rhein- G. Barrett-Hamilton 11. 1.2. 107-108.

hessen. [Erlanqcr.) (p).

2 i, juv. Ummerstadt, Thiirin- Lord Lilford (p). 11. 1. 1. 94-95.
gen. {Schucliarclt.)

9. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 93.

{Woltcrstorff.) 0. 2. 8. 5.

9. Niesky, Silesia, Ger- Lord Lilford (p). 97. 12. 4. 19.

many. {Baer.)

6. Teufen, Appenzell, 0. Thomas (p). 2. 8. 4. 24.

Switzerland. (Zolli- {Type of Putorius p. manium

kofer.) Barrett-Hamilton.)

S. St. Gallen. {ZoUi- 0. Thomas (r). 4. 4. 5. 32.

kofci:)

9. Ilorence, Italy. E. Cavendish Ta.ylor 5. 5. 6. 9.

(c & p).

3 (J, 2 c? juv. Viterbo, Rome. G.Barrett-Hamilton 11. 1. 2. 30-34.

(C. Coli.) (p).

6 juv. Burgos, Spain. G. S. Miller (c). 8. 8. 4. 49.

MusTELA PUTORIUS AUREOLUS BaiTett-Hamilton.

1904. Putorius putorius aureolus Barrett-Hamilton, Ann. and IMag. Nat.
Hist., 7th ser., xiii, p. 389, IMay, 1904. Type in British Museum.

Type locaUty. — Ferrol, Province of Coruiia, Spain.

(geographical distrihution. — Southern and western portions of
the Iberian Peninsula.

Diagnosis. — Underfur usually yellowish to extreme base, the
exact shade sometimes as bright as buff-yellow.

Measurements. — Type : hind foot, 61 -5 ; ear, 19. For cranial
measurements see Table, p. 426.

Specimens examined. — Four, from the following localities in Spain :
Ferrol, Coruna, 1 (type) ; Seville, 1 ; no exact locality (probably vicinity of
Seville), 2.

BemarJcs. — The status of this form is at present very
unsatisfactory. Only two specimens are known with exact
localities. One of these is from the extreme north-west of Spain,
the other from the neighbourhood of Seville. Together with two
Spanish skins without complete history they differ from ordinary
Mustela putorius in the conspicuously more yellow colour of the
underfur. Two skins from the neighbourhood of Burgos are,
however, indistinguishable from German or English specimens.

9. Ferrol, Coruiia, Spain. Dr. V. L. Seoane (p). 94. 3. 12. 1.

(Type of sub-species.)
9. Seville. [Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 9.

2. Spain. Lord Lilford (p). 94. 6. 11. 2-3.

426

CARNIVORA

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428

CARNIVORA

Genus VORMELA W. Blasius.

1857. Fcetorius Blasius, Saugethiere Deutschlands, p. 219 (part).
1884. Vormela W. Blasius, Bericht der Naturforsch. Gesellsch. in Bemberg,
XIII, p. 9.

Tyije species. — Mustela sarmatica Pallas = M. peregusna
Gueldenstaedt.

Geographical distribution. — From south-eastern Europe to
central China.

Characters. — Skull narrow, somewhat flattened (depth of
brain-case a little more than half mastoid breadth), the dorsal
profile slightly arched posteriorly, rather strongly bent down
from orbital region forward, the zygomatic arches not specially
widened, and postorbital region not unusually narrowed (distance
from point of greatest narrowing to zygoma less than breadth of
postorbital constriction) ; rostrum rather short and broad, its
width about equal to that of interorbital constriction, the
distance from anterior rim of orbit to gnathion equal to one-half
length of brain-case ; auditory bulla moderately inflated, oval or
sub-triangular in outline, the meatus with a tubular rim, the
anterior extremity of bulla in contact with hamular and nearly
or quite reaiihing level of foramen ovale ; dental formula :
i t^, c lii, })m H, m ^0^ = 34 ; teeth relatively somewhat larger
than in Mustela, the canines, especially those of upper jaw,
relatively higher than in any of the other European Musteliclne ;
upper carnassial robust, the width of its cutting portion about
half length, the anterior cusp so developed that its greatest
height is neai-ly three-quarters length of outer border of
tooth ; posterior cusp greatly reduced, appearing as a mere
thickening of posterior margin of crown ; inner lobe of carnassial
much wider than in Mustela, its posterior border extending about
to middle of crown ; upper molar elongate-pyriform, the outer
portion only a little narrower than inner, the median constriction
slight, the axis of crown strongly oblique to median line ; the
protocone, paracone and hypocone small but distinct, the metacone
barely indicated ; lower carnassial essentially as in Maries, the
metaconid small though evident, but posterior heel smaller, its
area less than half that of cutting portion of crown ; external
form polecat-like ; fur rather soft and dense, varied with black,
brown, and whitish or yellowish ; tail about half as long as
body, bushy.

Itemarks. — This strikingly characterized genus is readily
distinguished from Mustela, apart from its peculiar external
appearance, by the structure of the lower carnassial and by the
relationships of the hamular, auditory bulla and foramen ovale.
Two species are known, one of which ranges as far west as
eastern Hungary.

VORMELA 429

VORMELA PEREGUSNA Gueldenstaedt.

1770. Pcrcgusna, nova Miistcl;r species, Gueldenstaedt, Nov. Comm. Acad.

Sci. Imp. Petrop., xiv, p. 441.

1771. Mustcla sarmatica Pallas, Reise durcli verschiedene Provinzen des

Russischen Reichs, i, p. 453 (along the Volga in southern Russia).
1857. Fcetorius sarmaticus Blasius, Saugcthiere Deutschlands, p. 226.
1910. Putorius sarmaticus Trouessart, Fauna Mamm. d'Europe, p. 77.
1910. Vormcla percgusna Miller, Proc. U.S. Nat. Mus., xxxviii, p. 385,

August 19, 1910.

Tjipe Jocality. — Banks of the River Don, southern Russia.

GeograpMcal distribution. — From central Asia west through
Asia Minor and southern Russia to Roumania, Bulgaria and
eastern Austria-Hungary (Bukowina).

Diagnosis. — Size about as in Mustcla putorius (head and body
in an adult male, 340 mm., condylobasal length of skull about
50 mm.) ; auditory bulla sub-triangular in outline.

External form. — -In general external characters (dry skins
only examined) the animal appears to agree closely with Mustcla
putorius, except that the ear is relatively larger, the tail is more
bushy and less tapering, occasionally appearing somewhat
flattened, and the claws on front foot are more slender and
compressed. Soles and palms thickly furred except on the pads,
all of which are naked ; palm with two posterior tubercles, the
outer somewhat smaller than in M. putorius, the inner about
half as large as outer. Mammffi : a 4-4, i 2-2 = 12.

Colour. — Ground colour of upper parts raw-umber, most of
the hairs with darker tip and lighter subterminal band, a few
blackish throughout, the combination producing a slight eiiect of
variegation. Spots varying from a whitish buff to bufF-yellow,
and arranged as follows : a large spot covering base of tail and
extending slightly on rump (often jjartly divided by a dark
streak along middle) ; a broad stripe extending obliquely down-
ward from middle of shoulder across side of body, and separated
from its fellow in median line above by a brown, irregularly
defined, or interrupted line ; about thirty-five small spots in
region between shoulder stripes and base of tail, their boundaries
indistinct and tending to coalesce into ti'ansverse bands, their
total combined area slightly less than that of brown background ;
a stripe extending from back of ear along side of neck nearly to
shoulder ; several small ill-defined spots on back of neck tending
to form a median longitudinal stripe ; underfur in dark area a
slaty-drab, in light areas similar to long hairs but paler, so that
the colour pattern becomes more sharply defined on areas where
the fur has been nearly worn oft'. Head, feet, legs and under-
parts varying from dark seal-brown to black. Lips and inter-
ramia whitish or buflfy. A Ijroad white or buffy crescent crosses
forehead above eyes and extends downward and backward to
sides of throat behind ears. Dark crown area emphasized by a

430

CARNIYOKA

suffusion of whitish behind it. Eai" white or bufl'y except at
extreme base, which is blackish. Tail black at tip, elsewhere
white set off by underlying black ; the individual hairs cream-
colour through basal third, then darkening rapidly through raw-
umber to black, the terminal third white.

Skidl. — The skull is smaller than that of Miistcla putoriits,
though resembling it in its general strength and massiveness,

Fig. 86.
Vonnela peregusna. Nat. size.

particularly in lateral view. Dorsal profile almost exactly as in
Miisfela puiorins, flat from level of anterior portion of orbit to
occipital region, the rostrum bent downward at an evident angle.
Depth of rostrum relatively great, as in J\I. jnUorius, and notice-
ably exceeding that of Musfela erminea. Postorbital constriction
abrupt, as in Mustela eversmanni, giving the skull when viewed
from above a very peculiar aspect as compared with that of

431

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432 CARNIVORA

M. imtoriiin. In fully adult individuals the postorbifcal breadth
is noticeably less than that of rostrum or interorbital region.
Viewed from below the most striking feature in general outline
of skull is the shallowness of the emargination between mastoid
process and base of zygoma. Tip of hamular in contact with
bulla, a peculiarity due principally to the general shifting forward
of bulla with regard to glenoid region.

Teeth. — As compared with those of Mustela jmtorlns and
MusteJa erminea the teeth are through-
out relatively heavier and the cusps
more elongated. Height of upper
canine slightly more than depth of
rostrum, instead of decidedly less than
rostral depth as in Mustela imtoriuH.
Upper carnassial with broader antero-
internal projection than in M. jytitorlus,
and upper molar set more obliquely to
main axis of skull. Lower carnassial
strikingly different from that of all
^jg g_ species of Mustela in the presence of

a well developed third cusp on anterior

\ nrmela jieregusna. , , , /. /. i j i , i <.

Teeth, x u. elevated portion oi tootli, the form ot

which is thus essentially as in the
martens. Interrelationships of small premolai's much as in
Mustela putorim.

Measurements. — Adult male and female from Malcoci, Do-
brudscha, Roumania (from well-made skins) : head and body, 380
and 350; tail, 160 and 160; hind foot, 45 and 40; ear, 25
and 23. For cranial measurements see Table, p. 431.

Specimens examined. — Three, from the neighbourhood of IMalcoci,
Dobrudscha, Roumania (B.M. and U.S.N.M.) ; also others from various
localities in Asia.

BemarJcs.—Vormela psregusna is so well characterized both
cranially, dentally and externally as to require no detailed com-
parison with other European carnivores.

6. Malcoci, Dobrudscha, Rou- Hon. N. C. Rothschild 10. G. 4. 1.
mania. {Schliltcr.) (p).

Sub-Family GULONIN.^.

Geographical flisirihution. — iSTorthern forested portions of the
Nortliern Hemisphere.

Characters. — Dentition essentially as in the Miistelinse but
much less trenchant, the small premolars opposite and not capable
of shearing action, the points of all but j>m^ and pm^ widely
separated when jaws are closed ; upper carnassial very robust,
its posterior cusp broad, almost 1^ at -topped, its height much more

GULO 433

than half that of main cusp ; skull robust and heavy ; external
form heavy, not in the least cat-like ; feet sub-plantigrade.

Remarks. — The sub-family Guloninse, consisting of the genus
Gulo alone, is well characterized by its peculiarities of skull,
teeth and external form. Though usually regarded as a near
relative of the MiisieJinff', the genus more probably finds its true
affinities in the African Mellivova.

Genus GULO Storr.

1780. Gulo Storr, Prodr. ]\Ieth. INIamm., p. 34.

1857. Gulo Blasius, Stiugethiere Deutschlands, p. 208.

Type species. — Mustela gulo Linnajus.

GeograpTiical disfrihiiiion. — ISTorthern forests of the Northern
Hemisphere ; in Europe confined to Scandinavia and northern
Russia.

Characters. — Skull heavily built, rather narrow but not so
high as in Martes (depth of brain-case slightly more than half
mastoid breadth), the dorsal profile strongly curved dov/nward
anteriorly, the zygomatic arches not specially widened, and
postorbital region not unusually narrowed (distance from point
of greatest narrowing to zygoma less than breadth of postorbital
constriction) ; rostrum short and veiy robust, its width about
equal to that of interorbital region, the distance from rim of
orbit to gnathion about two-thirds length of brain-case ; auditory
bulla inflated along inner border, elsewhere rather flat, the meatus
forming an ill-defined, gradually nari^owing tube, the transverse
diameter of bulla greatest ; paroccipital process robust, long,
standing out conspicuously behind bulla ; dental formula :
i !.;3, c jlj, pm jq, tu ^_., = 38 ; teeth relatively lower, wider and
more robust than in Mustela (their actual size conspicuously
greater) ; small premolars almost directly oi^posed, not fitting
between each other when jaws are shut, and only pm^ and pm^
capable of being brought in contact ; lower carnassial with crown
wider anteriorly than posteriorly, the metaconid absent, the area
of posterior heel only about one-tifth that of cutting portion of
crown ; external form short and heavy, almost bear-like, the
head moderately pointed, the ears small, nearly concealed by the
fur, the tail very short, not so long as head, densely bushy with
hairs much longer than itself ; legs short, feet large, digitigrade,
the moderately long claws partly retractile ; fur long, soft and
dense.

Itemarlis. — The genus Gulo is represented by two closely
related species, (t. lu,scu,s of the northei'n portions of America,
and G. gulo of the Old World.

2 F

434 CARNIVORA

GULO GULO Linnreus.

1758. IMustela] gulo Linnaeus, Syst. Nat., i, 10th ed., p. 45.

1816. Gtilo vulgaris Oken, Lehrbuch d. Naturgesch., iii, pt. 2, p. 1004 (Re-
naming of giilo).

1820. Gido borealis Nilsson, Skaud. Faun., i, p. 95 (Renaming of giilo).\

1820. Gulo arcticus Desmarest, Mammalogie, p. 174 (Renaming of gulo).

1829. Gulo ardos Kaup, Entw.-Gescb. u. Naturl. Syst. Europ. Thierw., i,
p. 68, described on p. 09 (Renamiug of gulo).

1857. Gulo borealis Blasius, Saugetbiere Deutscblands, p. 209.

1910. Gulo luscus Trouessart, Paune Mamm. d'Europe, p. 71. Not Urs7is

luscus Linnneus, 1758.

1911. Gulo gulo CoUett, Norges Pattedyr, p. 3S0.

Type locality. — Lapland.

Geographical distrihution. — Northern forests of the Old
World ; in Europe, confined to Scandinavia and northern Russia.

Diagnosis. — Largest of the European Mustclidve (head and
body, about 800 mm.; condylobasal length of skull, about 130
to 140) ; general colour dark brown with a lighter crescent on
forehead and another lighter area crossing rump and extending
along sides of body toward front legs.

External characters. — General form heavy, badger-like, the
apparent size of the animal as well as its shortness of limb
exaggerated by the great depth and peculiar quality of the fur.
Head short and thick ; ears densely haired both inside and out,
low and rounded, fully as wide as high, not appearing con-
spicuously above fur ; muzzle pad well defined, naked, the bare
region extending downward to region where upper lip folds
inward (the infolded portion of lip hairy). Feet broad and strong,
the sole and palm each with three naked or almost naked pads,
which are wholly concealed in winter by the dense growth of
long hairs with which the feet are elsewhere covered, though in
the summer pelage they may be partly exposed ; claws strong,
not completely retractile, those on hind foot slightly the longer.
Tail short, the vertebrse scarcely equal to hind foot, the entire
tail covered with long hairs, so as to form a dense brush, the
hairs at tip 70 to 140 mm. in length, according to season, and
often as long as vertebrae. Fur of a peculiar and characteristic
quality ; underfur soft and dense, its hairs 20 to 30 mm. long
on back according to season ; longer hairs rather coarse and very
numerous, their length about 50 mm. on back, but increasing
abruptly to twice as long on flanks and rump, thus adding greatly
to the heavy appearance of the animal ; long hairs on wiist and
sides of fore foot tending to become almost bristle-like in texture.

Colour. — Upper parts and belly a rich dark brown becoming
blackish on legs, feet and tail ; a broad light (wood-brown) stripe
crosses rump and extreme base of upper side of tail and curves
forward along sides to axillary region where it gradually
disappears ; crown to and partly including ears, and face to

GULo 435

between eyes, suffused with cream-buff, so that this region is
paler than any other part of the animal ; muzzle and cheek to
and surrounding eye in a stripe about 10 mm. wide, dark brown ;
claws light horn-colour.

SJcull. — The skull is larger and more massive than in any of
the other European Mitsfelidse, its general form somewhat inter-
mediate between that of Melesi and Maries. Dorsal profile with
slight anterior concavity over middle of short nasals, then rather

Fig. S8.
Gido gulo. Kediiced.

strongly convex to bi-egmal region, behind which it is straight
and nearly horizontal to distinctly overhanging lambda. Brain-
case elongate-ovate in outline when viewed from above, about
l\ times as wide as deep, the sagittal crest moderately developed
in old individuals and joining rather low lambdoid crests in a
noticeably backward projecting point, the entire occipital region
sufficiently overhanging to conceal condyles when viewed from
above. Floor of brain-case with no special peculiarities, the
basioccipital essentially as in Maries martes but relatively wider.

2 F 2

436

CARNIVORA

Auditory bullae inflated along inner border, elsewhere rather flat,
their general form elongate flask-shaped (transverse diameter
greatest), much like that in Meles }nch'i>, the nieatal tube rather
long but not sharply differentiated from main portion of bulla.
Postorbital region less abruptly contracted than in Meles meles
and Martes martes, its width slightly more than half that of
bi*ain-case. Interorbital region about as wide as rostrum,
smoothly convex both longitudinally and laterally. Postorbital
processes short but relatively better developed than in Martes.
Rostrum short and heavy, the width across canines equal to
distance from ajnathion to infraorbital foramen. Orifice of

riG. 89.

Gulo gulo. Eechiced.

anteorbital foramen relatively small, its actual size scarcely
greater than that in Martes martes, its situation over posterior
root of jm^ and noticeably in front of anterior margin of rather
small orbit, a peculiarity not shared by any of the other European
members of the family. Zygomata essentially as in Martes
martes, but less bowed upward, and very noticeably comj^ressed
and deepened, especially over glenoid region. Palate rather
liroad, its width between molars equal to nearly 2i times greatest
diameter of molar ; incisive foramina elliptical, slightly oblique,
between canines, the minute median foramen between anterior
borders ; posterior extension and mesopterygoid space essentially

GULo 437

as in Mdrfcs. Mandible very robust and massive, but with no
special peculiarities of form. It is distinguishable from that of
Mele>< iiieles, which approaches it in size, by the deeper, more
compressed ramus and relatively higher coronoid process.

Teeth. — Except for their much greater size the incisors and
canines resemble those of Maries marten. Small premolars both
above and below with crowns relatively lower and broader than
in Marten, and less distinctly triangular when viewed from the

I'lG. t)0.
Gulo ijulii. Reduced.

side, the teeth of the upper and lower jaws almost directly
opposed and not fitting between each other when jaws are closed,
the tips of pni-'- and p)ii_^ barely coming in contact, those of the
other teeth separated by a wide space ; prn^ with slightly raised
cingulum at posterior border, but without trace of the secondary
cusp present in Maries. Upper carnassial like that of Maries in
general outline, but posterior cusp relatively much larger, its
height distinctly more than half that of main cusp, its outer

438

CARNIVOEA

surface at first nearly llai, then sloping almost perpendicularly
to very rudimentary cingulum ; commissure relatively shorter than
in Maries. Lower carnassial slightly moi'e than twice as long as
broad, the sectorial portion of the tooth consisting of two equally
robust cusps, the lower the paraconid, whose anterior border
slopes backward at an angle of about 50", and the higher the
protoconid, the two connected by an abruptly angled commissure.
No trace of metaconid other than a barely noticeable thickening
in the ridge which extends from apex of protoconid downward
along inner side of cusp to cinguh>m ; a similar ridge at outer
side of protoconid and across posterior heel slightly external to

Fig. 91.
Gulo gnlo. Teeth. Nat. size.

middle. Crushing portion of crown reduced to a mere ledge
much wider than long and with scarcely half the area of base
of protoconid ; a faintly indicated outer tubercle. Second lower
molar subterete, flat, with faintly developed cusp and longi-
tudinal and transverse cross ridges, its crown area about equal to
that of pm^, and a little larger than that of posterior heel of vi.,.
Upper molar relatively small, its crown area scarcely more than
half that of carnassial, its greatest diameter about two-thirds
that of carnassial ; diameter of inner portion of crown slightly
greater than that of outer portion, the constriction between the
two portions not well defined : cusps essentially as in Maries
martes, but paracone and metacone tending to become confluent.

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CARNIVORA

Mca,siircincHf><. — Adult (summer pelage) from Sweden (from
skin) : head and body, 825 ; tail, 125 (pencil, 75) ; hind foot,
140±. For cranial measurements see Table, p. 439.

Specimens examined. — Six, from the following localities : —
Norway : Vaage, Gudbrandsdal, 1 (Genoa) ; Egersund, Stavanger,
1 skeleton and 1 skull.

Sweden: Frastvik, 1; no exact locality, 1 skull (U.S.N.M.).
Lapland : No exact locality, 1 skull (U.S.N.M.).

6 skeleton, Egersund, Norway. K. H. Schaanning (c). flO. 11. 26. 1.
? skull. \11. 10. 23. 1.

Family YIVERRID^.

1821. Viverriche Gray, London Med. Repos., xv, p. 301, April 1, 1821.

Geocirapliical didrUmtion. — Warmer portions of the Old World
(exclusive of Australia) ; north to the Iberian Peninsula and
southern France.

Characters. — Dentition essentially as in the MHnteVidse, but
vipper molars usually 2-2, and upper carnassial usually with
three outer cusps ; auditory bulla, when not rudimentary
(1 African genus), inflated, thin-walled, divided by a distinct
septum into two chambers ; form usually rather slender, the legs
moderately short ; size moderate ; feet digitigrade.

JRemarJiS. — The family Viccrridse is a tropical Old Woidd
group, containing about 25 genera, the ranges of two of which
extend into the Mediterranean region of Europe.

KEY TO THE EUROPEAN GENERA OF YIVEBBID^.

General form ferret-like ; ear broader than high ; body and
tail uniformly grizzled ; bony palate extending behind
molars in median line about half distance to hamulars ;
orbit nearly or quite enclosed by bone posteriorly (Mon-
goose) Mangos, p. 440.

General form cat-like; ear higher than wide ; body spotted;
tail ringed ; bony palate extending behind molars in
median line much less than half distance to hamulars ;
orbit widely opening into temporal fossa (Genet) Genetta, p. 446.

Genus MUNGOS Geoff'roy and Cuvier.

17'J5. Mitncjos Geoffroy and Cuvier, Magasin Encyclopedique, ii, p. 187

{mungo, p. 184).
1811. Herpestes lUiger, Prodr. Syst. Mamm. et Avium, p. 135 (misspelt

Herpertes in text, corrected in list of errata, p. 802) {ichneumon).
1824. Mangnsta Horsfield, Zool. Researches in Java (javanica).
1830. Martes Wagler, Nat. Syst. der Amphibien, p. 29 (Substitute for

Herpestes lUiger, preoccupied in botany). Not of Pinel, 1792.
1842. Mungo Lesson, Nouv. Tabl. Regne Anim., Mamm., p. 63 {iclmeumon).
1907. Mitngos Thomas, Ann. and Mag. Nat. Hist., 7th ser., xix, p. 119,

January, 1907.

Type species. — Viverra mungo Gmelin.

Geographical distribution. — Africa, southern Asia, and larger

MUNGOS 4-41

Malayan Islands ; in the Mediterranean region one species
inhabits the Iberian Peninsula.

Characters. — External form ferret-like ; the ear low and
wide ; the tail broad and flattened at base, tapering con-
spicuously through terminal half, the fur coarse, grizzled ; soles
and palms naked ; claws long, non-retractile ; skull robust,
deepened anteriorly ; orbit small, nearly or quite surrounded by
bone ; backward median extension of palate long ; auditory bulla
complete, its two divisions conspicuously contrasted in size and

form ; dental formula : i ^~'\ c ^ ^ pm '^ ■*, m "^ " = 40 ; cheek-teeth
' 0-3' 1-1' -^ 4-4' 2-2 '

trenchant.

BcmarJi^. — The widely dispersed genus Miuigos, which is one
of the most characteristic features of the carnivort)us fauna of
Africa and southern Asia, where about 60 forms have been dis-
covered, is unknown in Europe outside of the Iberian Peninsula.

MUNGOS WIDDRINGTONII Gray.

1842. Hcrpcstes tvlddringtonii Gray, Ann. and Mag. Nat. Hist., 1st ser., ix,

p. 50, March, 1842. Type in British Museum.
1909. Hcy2X'stcs ichneumon var. fcrruqinca Seabra, Bull. Soc. Portugaisc

Sci. Nat., II, p. 286, May, 1909 (Altemejo, Portugal).
1909. Hcrpcstes ichneumon var. dorsalis Seabra, Bull. Soc. Portugaise Sci.

Nat., II, p. 286, May, 1909 (Eibatejo, Portugal).

1909. Ilcrpestes ichneumon var. grisea Seabra, Bull. Soc. Portugaise Sci.

Nat., II, p. 286, May, 1909 (Ribatejo, Portugal).

1910. Herpeste.s ichneumon widdringtoni Trouessart, Fauna Mamm.

d'Europe, p. 89.

Tj/pe localifij. — Sierra Morena, Spain.

Geographical distrihution. — Iberian Peninsula, principally in
the south ; exact limits of range not known.

Diagnosis. — Similar to the African Mangos ichneumon, but
with larger; carnassial teeth (greatest diameter of upper carnassial
about 11 "8 instead of 10*8 in males, 11*2 instead of 10 "2 in
females), and more inflated auditory bulhe (lateral diameter of
swollen portion about 12 mm. instead of 10 mm.).

External characters. — General form slender and ferret-like,
the conspicuously tapering tail about as long as body without
head, the legs short, the head rather small and pointed, the ear
low and rounded, the fur remarkably coarse and harsh. Ear
scarcely rising above fur, its width decidedly greater than height
from meatus, its outline evenly I'ounded. Rostrum slender,
tapering, the nostril pad entirely naked, its surface nearly
smooth, the bare area extending downward as a narrow line
across middle of upper lip ; region from muzzle to and imme-
diately surrounding eye very scantily haired, in some specimens
almost bare ; upper eyelid with a fringe of erect black hairs
about 5 mm. long ; whiskers short and inconspicuous, scarcely
extending beyond eye when laid backward. Feet rather large

442

CARNIVORA

and elongate, witli lung, no!i-retractile claws, those on front feet
longest ; hallux and pollex short but with well developed claw
more than half as large as those on other digits ; sole and palm
completely iiaked, the sole with a heart-shaped tubercular mass
about 15 mm. wide occupying region immediately behind bases of
digits, the palm with a similar mass at base of fingers, and an
isolated roundish median posterior tubercle about 8 mm. in
diameter. Fur with dense somewhat woolly under portion, the
hairs of which are about 20 mm. long, and a very abundant
growth of coarse almost bristly longer hairs (these essentially
absent along entire median region of underparts), whose length
is about 50 mm. at middle of back, increasing to 75 mm. on
flanks, rump, and base of tail ; near middle of tail the length of
the longer hairs decreases rather abruptly to about 50 mm., the
hairs at the same time becoming somewhat appressed, bhus
giving the tail its characteristic tapering form ; pencil long and
full. Mamma? : a 2-2 = 4.

Colour. — The colour does not differ appreciably from that of
Moroccan and Egyptian specimens of Miukjos ichneumon. Head,
back, sides and tail (except the clear black tip) a coarse grizzle
of blackish and pale buff, each hair on body and tail buff at base
and with three buff annulations 3 to 5 mm. in length interposed
between the blackish tip (5 mm.) and the three blackish
intervening areas, the length of which varies from 6 to 10 mm.
Underfur dull tawny, the hairs slaty at base, the tawny
appearing everywhere irregularly at surface, though mostly
overlaid by the annulated bristly hairs. On body and tail the
black and buff are about equally noticeable, but on head the
annulations become much liner and the dark colour predominates,
often appearing as a clear blackish brown wash on face and
muzzle. Ears a light indefinite grizzled brown, somewhat paler
than the broccoli-brown of Ridgway. Underparts from chin to
base of tail a dark brown slightly or not grizzled, the exact
shade varying from a dark hair-brown to broccoli-brown. This
extends down inner side of hind legs to feet, where it becomes
nearly black, and involves entire foi'e leg except the uppermost
outer portion. Onfoi^efeet and outer portion of leg below elbow
it becomes blackish.

Shull. — The skull is narrow and rather high, mastoid breadth
less than occipital depth including bullae, with deep short rostrum
(depth at front of orbit greater than distance from orbit to tip of
premaxillary) and small orbit (vertical diameter less than depth
of rostrum at canine) completely encircled by bone in the adult
and with the circular form always clearly indicated by the long
drooping postorbital process and the distinct orbital process of
the zygoma rising to meet it. Anteorbital foraman small, higher
than wide, over space between jpm- and pm^. Median back-
ward prolongation of palate conspicuous, extending rather moi-e
than half way to hamulars, its posterior border squarely truncate

MDNGOS 443

or irregularly aiigular-emarginate, never with median projection ;
mesopterygoid space much less than twice as long as broad,
parallel-sided ; hamulars robust, the thickened extremities bent

riG. 92.

Mimgos ividdringtonii. Keduced.

444

CARNIVOEA

abruptly upward. Auditory bulla with anterior annular portion
small and flattened, the small meatus with short but evident
tube marked off from ring by a constriction ; posterior inflated
part of bulla large, higher than wide, extending noticeably below
level of palate, and projecting laterally beyond mastoid level ;
greatest transverse diameter of inflated portion about equal to
least breadth of basioccipital. Paroccipital process long, narrow
and scale-like, closely flattened against posterior surface of bulla,
less than half of which it covers. Lambdoid and sagittal crests
well developed, the latter extending across entire brain-case to
postorbital region. Mandible I'obust, the lower border bent
abruptly upward at level of m.^ ; coronoid process short and
broad, its width at level of condyle much greater than height
above condyle.

Teeth. — The teeth are robust and strong but not remarkably
large. Incisors and canines with no special peculiarities, the

cutting edge of upper incisors entire,
that of lower incisors obscurely bi-
lobed, the outer lobe largest. Ante-
rior premolar both above and below
simple, smaller than corresponding
outer incisor. Second and third
premolars triangular when viewed
from side, the two upper teeth with-
out true secondary cusps, but pm}^
sometimes with a slightly developed
posterior basal tubercle ; inner me-
dian projection well developed and
beaiing a small cusp in prii\ slightly
indicated in pju- • the two lower teeth
with longer base and less robust
main cusp, pjKg with slightly de-
veloped anterior basal cusp and a
small but distinct median cusp on
Fourth lower premolar like second
and third but larger, the crown wider behind and the cusp
on posterior cutting edge large and well developed. Upper
carnassial with large antero-internal lobe somewhat exceeding
posterior heel in area, its cusp robust ; antero-external corner
with large, occasionally almost cusp-like cingulum ; anterior
box'der of tooth slightly concave. Lower carnassial with the
three anterior cusps large, the outer exceeding the two inner,
which are of approximately equal height, the posterior heel low,
its area barely half that of anterior portion of tooth and about
equal to that of second molar, its postero-external border elevated
and trenchant. First upper molar with crown nearly twice as
wide as long, its anterior border conA^ex, its outer and posterior
borders concave; pi'otocone large, occupying entire inner border
of crown ; paracone and metacone small, sub-equal, the latter

Munyos widdringtonii.
Teeth. Nat. size.

posterior cutting edge.

445

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CARNIVORA

occupying postero-extex*nal corner of croMn, the former exceeded
by the large low parastyle, the basal area of which is nearly
equal to that of protocone. Second upper molar with crown area
only about one-third that of first, its antei'o-external extremity
fitting into convexity behind metacone of m^ ; in general structure
it is like m^ except that the projecting parastyle is absent and
the paracone and metacone are less distinct. Second lower molar
with crown slightly longer than wide, somewhat exceeding the
heel of vij^ in height, its borders sometimes with four distinct
cusps, the two on inner side largest, but more often with postero-
external and postei'o-internal cusps partly confluent.

Measurements. — Type (adult male) : hind foot, 95 ; ear from
meatus, 34*5. Immature male and adult female from Seville:
head and body, 530 and 510; tail, 430 and 330; hind foot,
90 and 87. Adult male from Goto Doiiana, Huelva (from skin) :
head and body, 550 ; tail, 450 ; hind foot, 95 * 5. Immature male
from Ferrol, Coruna ; hind foot, 8G.

Specimens examined. — Eight, from the following localities in Spain :
Ferrol, Coruila, 1 ; Sierra Morena, 1 (type) ; Seville, 3 ; Goto Doiiana,
Huelva, 1 ; Andalucia, no exact locality, 2.

Memarhs. — Among the smaller carnivores of Europe Mungos
ividdringtonii is recognizable by its conspicuously grizzled coloura-
tion and long, coarsely-haired, tapering tail. It is nearly related
to the African jlf. ichneumon, though suihciently distinguished by
its larger teeth.

6. Ferrol, Coruila, Spain. Dr. V. L. Seoane 94. 11. 3. 1.

(c & p).
6 Sierra Morena. Capt. S. E. Widdring- 42. 2. 26. 2.

ton (p). {Type of species.)
6, ?. Seville. {Dr. A. Ruiz.) Lord Lilford (p). 95. 9. 4. 5-6.

skull. Seville. Col. Irby (c & p). 85. 9. 1. 1.

6. Coto Doiiana, Huelva. A. Chapman (c & p). 10. 7. 14. 1.

ad & juv. Andalucia. Lord Lilford (p). 78. 7. 3. 2-3.

Genus GENETTA Oken.

1816. Genetta Oken, Lehrb. der Naturgesch., iii, pt. 2, p. 1010 {genetta).
1841. Odmiflurus Gloger, Gemeinn. Hand- u Hilfsbuch der Naturgesch., i,
p. 72 (genetta).

Type species. — Viverra genetta Linnaeus.

Geograpfiical distrihution. — Africa and Mediterranean region
of Europe.

Characters. — External form cat-like, the ear narrow and high,
the tail subterete, not conspicuously tapering, ringed with dark
and light bands, fur soft, spotted with black on a light ground ;
soles and palms densely furred except on pads ; claws short,
retractile ; skull slender, not specially deepened ; orbit large,
opening widely into temporal fossa ; backward median extension

GENETTA 447

of palate short ; auditory bulla coroplete, its two divisions not
strongly contrasted in size and form ; dental forruula : i '^', c ~,
pm ^-^, m -"^ = 40 ; cheek-teeth trenchant.

l" 4-4' •_.-!; '

Bemarks. — The genus Genetta, though practically confined to
Africa, extends its range into the Mediterranean region of Europe,
chiefly in Spain and the western portion of France. Many local
forms have been described, the status of which is imperfectly
understood. Three of these appear, on the basis of the insufficient
material seen, to be recognizable in Europe.

KEY TO THE EUROPEAN FORMS OF GENETTA GENETTA.

Spots in first three rows at side of dorsal stripe
tending to be confluent, and occupying more
space tlian light ground-colour in same region

(Southern France) O. <j. r/wdanica, p. 452.

Spots in first three rows at side of dorsal stripe

tending to be distinct, and occupying less space

than light ground-colour in same region.

Ground-colour of back a huffy grey ; largest spots

about 20 mm. in diameter (Central and southern

Spain) G. g. rjenetta, p. 451.

Ground-colour of back a smoky grey ; largest spots

about 30 mm. in diameter (Balearic Islands)... G. g. baleanca, p. 452.

GENETTA GENETTA Linmeus.
(Synonymy under subspecies.)

Geographical distribution. — Iberian Peninsula, Balearic
Islands, and central France, north to Department of the Eure,
east to the valley of the Rhone ; probably in western Africa
also ; limits of range not known.

Diagnosis. — General characters as in the genus ; size moderate
(head and body, about 550 ; tail, about 450 ; condylobasal length
of skull, about 90 mm.) ; colour light grey with black spots
arranged in longitudinal rows, the spots not tending to run
together except on back ; tail with distinct alternating rings of
black and light grey.

External characters. — General form cat-like but rather more
slender, the tail about as long as body without head, the legs
moderately long, the ear large, higher than wide, the fur fine
and soft. Head moderately tapering, the muzzle somewhat
pointed ; ear rising conspicuously above fur, its general outline
oval, the upper border evenly rounded ofl", the anterior border
with a projecting basal angle, the posterior border with an
angular emargination somewhat below middle, the emargination
covered by a small rounded lobe arising from outer surface of
ear. Muzzle pad entirely naked, its surface finely granular, the
bare area extending downward as a narrow line dividing middle
of uj^per lip ; whiskers well develoj^ed, reaching beyond base of
ear when laid back. Feet cat-like, the digits short, armed with

448

CARNIVORA

retractile claws, those of hind foot slightly the larger ; sole, palm,
and under surface of digits velvety-furred except on balls of toes
and surface of tubercles ; palm with large, cordate, obscurely
3-lobed tubercular mass at base of longer digits, a small roundish
pad at base of thumb, and two semi-confluent, elongate pads, the
outer nearly twice as large as inner, at posterior margin of palm ;
sole with similar trilobed mass at base of longer digits, an
elongate pad at base of hallux, and a linear-elongate pad on outer
side of sole, partly divided into two by longitudinal furrow, and
extending from level of base of hallux more than half way to
heel. Fur not peculiar in quality, the underfur dense, its hairs
about 25 mm. in length on back, the longer hairs soft, exceeding
the underfur by about 5 to 10 mm.; hairs on tail somewhat
longer than those of body, about 50 mm. at middle, rather more
at base and less at tip. Mammae : a 2-2 = 4.

Colour. — Ground colour throughout a light grey, with or
without an evident bufly or brownish tinge, the slaty bases of
the hairs sometimes appearing at surface, particulai'ly on under-
parts. Back and sides thickly spotted with black, the spots
always confluent along median line, where they form a narrow
stripe extending from shoulders to base of tail, those on sides
arranged in four or five longitudinal rows, of which the two
uppermost on each side usually show some tendency to run
together into stripes, while the lower ones are more widely
spaced ; when not crowded the spots are sharply and evenly
outlined, roundish or somewhat longer than broad, the diameter
of largest about 30 mm. Neck with narrow well-defined lateral
black stripe and one or two less sharply marked median stripes.
Base of ear black, the terminal half grey. Cheek and face grey,
with ill-defined median dark streak and faint supraorbital shade ;
muzzle with conspicuous blackish area at base of whiskers and
extending back to eye, the extreme anterior region at side of
pad and beneath it sharply contrasted light grey. Feet grey
above with a few small black spots, the under surface blackish
except for lighter region between pads ; black of hind foot
extending upward to cover much of inner surface of thigh. Tail
ringed with black and light grey, the extreme base light, the tip
either light or dark, the rings about equally wide throughout,
their number 8 to 1 0 of each colour ; black rings tending to run
together along median line above on basal third of tail.

Skull. — The skull is not unlike that of Hcrpestes widdriugtonii
in general form, though much less robust, not unusually
deepened, the orbit large, widely open posteriorly, its %ertical
diameter greater than depth of rostrum at canine, the auditory
bullse not much inflated ; mastoid breadth about equal to
occipital depth including bullae ; depth of rostrum at front of
orbit considerably less than distance from orbit to tip of
premaxillary ; postorbital processes short, not produced down-
ward, no orbital process of zygoma, the large orbit thus opening

GENETTA 449

widely into temporal fossa ; anteorbital foramen moderately large,
slightly higher than wide, over middle of pm^ ; median backward
prolongation of palate short, extending about 3 mm. behind
level of last molar, its posterior border double concave, with

> Fig. 94.

Genetta genetta. Keduced.

short but well developed median projection ; mesopterygoid
space a little more than twice as long as wide, parallel-sided ;
hamulars slender, with tapering, neai'ly horizontal extremities ;
auditory bulla with anterior annular portion rather large, slightly

2 G

450

CARNIVORA

«*

9

but evidently inflated, the large meatus without trace of tube ;
posterior inflated part of bulla moderate, much less high than
wide, extending downward about to level of palate, and laterally
barely reaching mastoid level, the greatest transverse diameter
of inflated portion about two-thirds least breadth of basioccipital ;
paroccipital process short and wide, applied to and completely
covering posterior surface of bulla, but not thin and scale-like
as in Mungos ividdringtonii ; lambdoid crest high ; sagittal crest
evident posteriorly, but low and inconspicuous over middle and
fore part of brain-case ; mandible slender, the entire lower
border convex to level of middle of coronoid process, the
convexity scarcely moi-e evident posteriorly than anteriorly ;
coronoid process with height above condyle about equal to width
at same level.

Teeth. — The teeth do not differ markedly from those of
Mungos widdringtonn except that they
are less robust, a peculiarity especially
noticeable in the lower premolars.
Lower incisors very slender, with biloba-
tion obscure. Anterior premolar both
above and below simple, actually as well
as relatively larger than in Mungos wld-
dringtonii, its size greater than that of
corresponding outer incisor, pm^ with
minute postero-basal cusp ; second and
third premolars triangular when viewed
from the side, the two upper teeth with-
out true secondary cusps, but both with
cingulum rather prominent before and
behind, and pni^ often with a minute
lobe near base of posterior cutting edge ;
inner median projection well developed
and bearing a small cusp in jJi/i^, slightly
indicated in jjm- ; the two lower teeth with base scarcely longer,
both with small but evident anterior and posterior cingulum
cusp, and prn^ with well developed median cusp on posterior
cutting edge ; fourth lower premolar Uke second and third
but larger, the crown less narrow behind, and the cusp on
posterior cutting edge better developed. Upper carnassial with
well developed, cusp-bearing antero-internal lobe, the area of
which is distinctly less than that of posterior heel ; antero-
external corner of crown with cusp-like cingulum; anterior
border of crown rather abruptly concave f lower carnassial with
the three anterior cusps large, the antero-external highest, the
antero-internal less elevated, the postero-internal barely half the
height of the others ; posterior heel with area considerably less
than half that of anterior portion of crown and not equal to that
of posterior molar, a low but evident cusp near middle. First
upper molar as in Mungos, but anterior border scarcely convex

Fig. 95.

Genetta genetta.
Teeth. Nat. size.

GENETTA 451

and posterior border nearly straight ; cingulum often with
indication of a minute cusp between protocone and paracone and
another between protocone and metacone ; paracone and meta-
cone not well defined from each other but tending to coalesce
to form a rather high commissure continued outward on
parastyle. Second upper molar leather more than one-third first
in area, its outer border extending to outer border of metacone
of the larger tooth, owing to the presence of a distinct lobe
representing parastyle. Second lower molar about as large as
heel of carnassial, its cusps varying from two to four, often
differing in the two jaws of the same animal.

Bemarhs. — Its slender form, elongate head, spotted body and
ringed tail distinguish this animal at a glance from all other
members of the European fauna. The material examined, while
inadequate to any final conclusions, indicates the existence in
south-western Europe of three local forms.

GeNETTA GENETTA GENETTA LinnteUS.

1758. IViverra] genetta Linnaeus, Syst. Nat., i, 10th ed., p. 45 (Spain).
1816. Viverra Genetta hispanica Oken, Lehrb. der Naturgesch., ii, pt. 3,

p. 1010 (Ronda, Malaga, Spain).
1816. Viverra Genetta gallica Oken, Lehrb. der Naturgesch., ii, pt. 3,

p. 1010 (alternative name for hispanica, suggested on the

supposition that the animal occurs in Prance also. Not Viverra

gallica Kerr, Anim. Kingd., p. 167, 1792, based on some Genet

from unknown locality).
1827. Genetta vulgaris Lesson, Man. de Mamm., p. 173 (Renaming of

genetta).
1830. ? [^Genetta'] comnmnis Burnett, Quart. Journ. Sci. Lit. Art, xxviii,

1829, p. 349 (Substitute for genetta). Nomen nudum.

1897. Genetta melas Graels, Mem. Real Acad. Sci., Madrid, xvii, p. 175,

pi. 2 A (Sierra Morena, Spain).

1898. [^Genetta'l genetta Trouessart, Catal. Mamm. viv. fors., p. 325.
1905. ? Genetta peninsulx Cabrera, Bol. Real Soc. Espafi. Hist. Nat

p. 266, May, 1905 (El Pardo, near Madrid, Spain).
1910. Genetta genetta melas, G. genetta peninsulx and G. afra Trou^sart,
Paune Mamm. d'Europe, pp. 88, 89.

Type locality. — Spain.*

Geo(jraphical distribution. — Central and southern Spain ;
limits of range unknown.

Diagnosis. — ^Spots in first three rows at side of dorsal stripe
tending to be distinct and occupying less space than light ground
colour in same region, the largest about 20 mm. in diameter ;
ground colour of back a distinctly bufty grey.

Measurements. — Adult male and female from neighbourhood
of Seville, Spain (approximate, from skin) : head and body, 550

* Linnaeus says : "Habitat in Oriente," an expression merely equivalent
to "foreign." His first reference is to Ray, whose first locality is Spain,
(see Thomas, Proc. Zool. Soc, London, 1911, p. 137).

452 CARNIVORA

and 560 ; tail, 445 and 470 ; hind foot, 81 and 84 ; ear, 46
and 43.

Specimens examined. — Nine, from the following localities in Spain :
near Seville, 5 (B.M. and U.S.N.M.) ; near Madrid, 2 ; Valencia, 1 skull
(U.S.N.M.); Gerona, 1 skull (U.S.N.M.).

Bemarks. — On the basis of the few specimens that I have
seen I am unable to recognize more than a single race of Genetta
from the Iberian Peninsula, apart from the somewhat doubtful
occurrence of the French form at the extreme north.

2 6,2 9. Seville, Spain. {Dr. A. Lord Lilford (p). 95. 3. 3. 5.

Ruiz.) 95. 9. 4. 2-4.

1. Madrid. Purchased (Parzu- 55. 11. 26. 3.

daki).

Genetta genetta balearica Thomas.

1902. Genetta genetta halearica Thomas, Ann. and Mag. Nat. Hist., 7th ser.,

X, p. 162, August, 1902. Type in British Museum.
1910. Genetta genetta halearica Trouessart, Paune Mamm. d'Europe, p. 88.

Type locality. — Inca, Majorca, Balearic Islands.

Geographical distribution. — Balearic Islands.

Diagnosis. — Similar to Genetta genetta genetta but general
colour paler, a light cream-buff with faint brownish tinge ; tips
of hairs of underf ur more whitish than in the continental Spanish
animal ; spots rather lai^ger (many of them attaining a diameter
of 30 mm.) but showing no peculiarities in arrangement.

Measurements. — Type (young-adult male) : head and body,
520 ; tail, 480 ; hind foot, 87. Adult male and female from the
same island : head and body, 580 and 550 ; tail, 440 and 420 ;
hind foot, 85 and 82. For cranial measurements see Table
opposite.

Specimens examined. — ^Four, all from the Island of Majorca.

6, 1 9. Majorca, Balearic Islands. 0. Thomas (p). 1. 6. 1. 1-4.

(Dr. Riutort.) (1. 6. 1. 3. Type of subspecies.)

Genetta genetta rhodanica Matschie.

1902. Genetta rhodanica Matschie, Verhandl. des V Intern. Zool. Congr.,

Berlin, 1901, p. 1139.
1910. Genetta genetta vulgaris Trouessart, Faune Mamm. d'Europe, p. 86

(not Genetta vulgaris Lesson).

Type locality. — Montpellier, Herault, France.

Geograpliical distribution. — South-western France from the
valleys of the Rhone and Loire to the Bay of Biscay ; perhaps
westward through the Asturias.

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GENETTA 455

Diagnosis. — Ground colour as in Gcnetta genctta genetta, but
black spots smaller (rarely attaining a diameter of 20 mm.),
more numerous, and showing a decided tendency to become
confluent. In region occupied by tirst three rows of spots at
each side of median stripe the black covers rather more space
than the light ground colour.

Measurements. — Adult female from Sainte-Cecile, Vaucluse,
France : head and body, 490 ; tail, 440 : hind foot, 85 ; ear, 53.
For cranial measurements see Table, p. 453.

Specimens examined. — Four, from the following localities in France :
Sainte-C^cile, Vaucluse, 1 ; near Nimes, Gard, 1 (Mottaz) ; near Biarritz,
Basses- Pyrenees, 2.

BemarJcs. — An immature individual from Ferrol, Coruiaa,
Spain, appears to be referable to this form rather than to
true genetta,

?. Sainte-Cecile, Vaucluse, France. M. Mourgue (c). 8. 7. 11. 1.
2 6. Biarritz, Basses-Pyrenees. Zoological Society 8. 1. 26. 1-2.

1. Ferrol, Coruna, Spain. Dr. V. L. Seoane 94. 11. 3. 2.

(c. & p.)

Family FELID^.

1821. Felidfe Gray, London Med. Repos., xv, p. 302, April 1, 1821.

Geograpliical distribution. — Entire continental region of both
Old and New Worlds, to the limits of tree growth ; Malay
Archipelago ; in Europe west to Great Britain.

Characters. — Larger cheek-teeth of a strictly trenchant type,
their crowns compressed and high, without the crushing surfaces^
the last upper premolar and first lower molar persenting the
extreme phase of carnassial modification, the inner lobe of upper
carnassial small, in front of middle of crown ; upper molar narrow,
trenchant, the main axis of its crown transverse to tooth-row ;
auditory bulla highly inflated, divided by a septum into two
chambers ; form usually slender, the legs moderately long ; size
moderate to large ; feet digitigrade ; toes, 5-4.

BemarJcs. — The family Felidse is one of the most widely
distributed and sharply differentiated groups of carnivores. At
present there is much difference of opinion as to the generic
status of its members, all of which bear the unmistakable stamp
of common descent. Two groups commonly regarded as genera
occur in Europe.

KEY TO THE EUROPEAN GENERA OF FELID^.

Small anterior upper premolar present, teeth 30 ; ear slightly

ornot tufted (Cats) Felis, p. 456.

Small anterior upper premolar absent, teeth 28 ; ear con-
spicuously tufted (Lynxes) Lynx, p. 470.

456 CARNIVORA

Genus FELIS Linnteus.

1758. Felis Linnseus, Syst. Nat., i, 10th ed., p. 41 (Type by tautonymy
Felis catus Linnseus).

1855. Catus Fitzinger, Wiss.-pop. Naturgesch. der Siiugith., i, p. 2G5
(catus by tautonymy).

1857. Felis Blasius, Saugethiere Deutschlands, p. 159.

1858. Catolyyix Severtzow, Rev. et Mag. de Zool., 2nd ser., x, p. 385 (catus).

Type S2jecies. — Felis catus Linnseus.

Geographical distribtdton. — Tropical and noi'thern temperate
portions of the Old World ; exact limits of range not known.

Characters. — Dental formula : * ||, c ti, pm pl, m ^^ = 30 ;
anterior upper premolar small but well developed, rarely
deciduous ; skull broad, rounded and deep, the zygomatic
breadth about three-quarters condylobasal length ; external form
slender, typically feline, the fur full and soft, not developing
special manes or tufts.

Remarks. — In the uncertainty now existing with regard to
the taxonomic rank of the various groups of cats the genus
Felis is here regarded in its most restricted sense with respect to
both characters and synonymy. As thus understood it contains
about twenty species, mostly of the Old World. Three of these
occur in Europe.

KEY TO THE EUROPEAN FORMS OF FELIS.

Fur of median dorsal area noticeably longer and

coarser than that of sides ; tail slender, its hairs

about 30 mm. in length ; no black stripes on

shoulders or back.

General colour of back and sides grey, the darker

markings practically absent (Sardinia) F. sarda, p. 468.

General colour of back and sides yellowish brown,

the darker markings evident though faint (Crete) F. agrius, p. 470.

Fur of median dorsal area not noticeably longer and

coarser than that of sides ; tail thick, its hairs

about 40 mm. long; two short black stripes

between shoulders, and a long median black stripe

on back F. silvesfris, p. 457.

Teeth enlarged (pni'^ and pm^ together 19 to 20 mm. ;
three lower cheek-teeth together 22 '4 to 23 '6

mm. ; w, 9*2 to 10 mm.). Southern Spain F. s. tartessia, p. 465.

Teeth normal (pni^ and pm* together 16*6 to 19 mm. ;

three lower cheek-teeth together 18 • 8 to 22 mm. ;

?Wi 8-0 to 8-8 mm.).

General coloiar light, approaching the smoke-grey

of Ridgway ; stripes on sides and legs obsolete

(Central Europe) F. s. sUvestris, p. 462.

General colour dark, approaching the broccoli-
brown of Ridgway ; stripes on sides and legs
well defined (Scotland) F. s. grampia, p. 464.

FELIS 457

FELIS SILVESTRIS Schreber.

(Synonymy under subspecies.)

Geographical distribution. — Central and southern Europe,
from Great Britain eastward into Asia Minor ; formerly occur-
ring everywhere throughout its range, but now restricted to the
wilder regions.

Diagnosis. — Size slightly larger than in the domestic cat, Felis
catus Linnjeus," and form appearing more robust owing to the
somewhat greater depth of the fur ; tail rather more than half
as long as head and body, its width at middle about 70 mm.,
abruptly, almost truncately, rounded at tip ; fur soft and full,
the hairs about 50 mm. long on sides, those along middle of back
about 10 mm. longer, but not noticeably different in texture ;
occiput and nape with four longitudinal black stripes ; two
short longitudinal black stripes on middle of shoulder ; a single
median black stripe from back of shoulder to base of tail ; sides
with two or more brownish cross stripes ; tail with black tip and
two to four complete or nearly complete black rings ; middle of
chest obscurely or not spotted ; ground colour of upper parts a
very pale cream-buff overlaid with whitish and slightly darkened
by the black annulations and tips of the longer hairs, the general
effect about smoke-grey.

External characters. — General form essentially as in the
domestic cat, but appearing heavier on account of the longer
fur, a peculiarity especially noticeable in the tail. Fore foot with
thumb much shorter than other digits, but bearing a well
developed claw ; third and fourth digits sub-equal and longest
(the third slightly exceeding fourth), the second and third
successively shorter ; palm with obscurely trilobed, somewhat
heart-shaped pad at base of longer digits ; a rounded pad on ball
of each digit, that on thumb small and inconspicuous, and a
noticeable, subterete, almost horny excrescence near wrist ;
surface of all tubercles smooth ; surface of palm elsewhere
velvety-furred. Hind foot with digits and pads essentially as in
fore foot except for absence of both hallux with its pad and
posterior horny excrescence.

Colour. — IJnderfur of back and sides a cream-buff distinctly
paler than that of Ridgway, the basal half of the hairs light
drab-grey. Longer hairs whitish from base to just below tips of
underfur, then blackish, followed by a whitish annulation about
7 mm. wide and a long black tip. The coincidence of the lower
dark area with the terminal portion of the under hairs makes
the latter appear at first sight brownish at tip. The general
effect of the ground colour is a buffy grey frosted with whitish.
This is very uniform throughout the head, back, sides and tail,

* For use of this name in place of Felis domestica Auct. see Pocock,
Proc. Zool. Soc, London, 1907, p. 149, June 12, 1907.

458

CARNIVORA

but on outer side of front legs there is usually a sooty tinge,
while the upper side of the feet becomes a nearly clear,
light, bufFy clay-colour. Underparts and inner surface of
hind legs cream-buff, becoming whitish or white on chin,
throat, fore part of chest and in intercrural region, and clouded
and blotched with blackish across middle of chest and on
inner side of fore legs, the amount of blotching on chest vari-
able ; soles and palms dark brown. Tail black at tip (about
70 mm.) and with two to four nari'ow (15 to 30 mm.) black or
brownish lings, usually narrower or incomplete below, and often
showing a tendency to become joined in median line above.
The dark markings on upper parts and sides are typically as
follows. Nape with four narrow longitudinal black stripes,
these extending to head where they tend to become confused
arid confluent, and disappearing at front of shoulder. Just
behind region where nape stripes disappear the shoulder has two
usually well defined longitudinal stripes about 60 mm. in length
and 10 mm. wide, lying 10 to 20 mm. apart. About 30 mm.
behind the shoulder stripes the median dorsal stripe begins, often
as a distinct spot, and continues to base of tail. The sides are
marked with several transverse, frequently obsolete stripes, of
which that starting at front of median dorsal stripe is usually the
best developed, while those behind it tend to become mere wavy
blotches, always preserving, however, when visible and when fur
is smoothed, their general transverse and parallel arrangement.*
In continuation of this system of transverse stripes, four or five
fairly distinct, narrow (5 to 10 mm.) bands usually cross outer
side of hind legs, and two or three narrower and less distinct
bands may be detected on front legs. Ear like ground colour of
back, the tip slightly blackish.

Skull. — The skull resembles that of Felis catus so closely that
it is impossible to find tangible characters by which to distinguish
it. In the larger races, however, a slightly greater size and
massiveness may be attained than in the domestic cat. General
aspect of skull broad, rounded and deep, the zygomata long and
very widely spreading (zygomatic breadth equal to about
three-quarters condylobasal length), the orbits nearly or quite
surrounded by bone, very large (diameter greater than that of
interorbital region, and area equal to or greater than that of
I'ostrum when skull is viewed from in front). Dorsal profile of
skull convex throughout, sometimes a little flattened in inter-
orbital region and along nasals, the latter occasionally terminating
in a slight anterior concavity ; nasals sloping very abruptly
forward and downward, the palatal depth at their posterior
extremity at least twice as great as that at front : from posterior

* There is never any trace of the spiral arrangement of the dark
markings characteristic of the typical form of the domestic Fdis cafiis as
described and figured by Pocock, Proc. Zool. Soc, London, 1907, pp. 143-168,
pis. VIII and ix.

FELIS 459

extremity of nasals the convexity of the dorsal profile is more
gradual and regular to slightly overhanging lambdal region ;
occiput obliquely truncate but not noticeably concave, the
condyles not visible when skull is viewed from above ; ventral
profile flat throughout. Brain-case broadly ovate or nearly
globular when viewed from above, the posterior outline distorted
in older specimens by the projecting lambdoid border, the
greatest width above zygomata slightly greater than depth at

7-ERZ( —

Fig. 96.
Felis silvestns.

middle, and about equal to distance from posterior border of
parietal to deepest portion of postorbital constriction ; sagittal
crest low, appearing first on interparietal, the only region where
it becomes a true crest, the muscular ridges in front of
interparietal at first separate and forming a lyre-like figure, but
later uniting in continuation of the median crest. Floor of
brain-case flat, without noticeable surface features ; auditory
bulUe large, highly inflated, longer than broad, formed mostly by

460

CARNIVORA

posterior chamber, the surface smooth, the aieatus large, not
tubular ; width of basioccipital at middle less than that of bulla
in same region ; mastoid and paroccipital processes small, the
latter thin and plate-like, both closely applied to surface of
bulla. Postorbital region abruptly contracted behind postorbital
process, the depth of the constriction increasing with age ;
postorbital process vei'y large, at first triangular in outline,
afterward becoming ligulate, its upper surface obliquely flattened.
Interorbital region much constricted, its least width considerably
less than that of orbit. Rostrum short and deep, the distance
from orbit to gnathion about equal to breadth over canines and
slightly less than palatal depth at posterior extremity of nasals ;
nasal branch of premaxillary broad, its width at front of nasal
often nearly equal to that of nasal in same region ; anteorbital
foramen moderately large, opening directly forward under
overhanging rim of orbit and above middle of first large
premolar. Palate broad, triangular in outline, its width
including molars about equal to median length, the median
posterior extension slight, but usually marked oS" at each side by
a lateral emargination extending in some instances nearly to
level of middle of carnassial ; incisive foramina moderately large,
elliptical in outline, slightly oblique, extending from level of
middle of canine to middle of space between canine and outer
incisor ; mesopterygoid space nearly parallel-sided, somewhat
longer than wide, its anterior border double-concave (encroached
on by blunt median spine of palate) ; hamulars slender, a little
converging ; external pterygoid process well developed, laminate,
angular, much smaller than hamular. Mandible rather slender,
the anterior border of coronoid rising at an angle of only about
50°, the distance from apex of process to posterior border of
molar about equal to that from latter point to front of
symphysis ; lower profile of ramus slightly convex, though with
anteiior and posterior flattened regions ; angular process straight,
rather short, a little below level of alveolar line.

Teeth.- — As a whole, and in comparison with that of European
Carnivores of other families, the dentition appears rather weak,
owing to the complete suppression of all crushing surfaces and
the consequent narrowness of the cheek-teeth. Incisors small
and weak, the crown area of i^, the largest tooth, about equal to
that of ^^ and i"^ together and scarcely more than one-eighth that
of canine ; upper incisors forming a continuous, straight or
faintly convex row separated from canine at each side by a well
marked diastema nearly as wide as base of canine ; i^ and z'^ sub-
equal, the second tooth slightly the larger, the crowns a little
compressed, slightly higher than wide when viewed from in
front, spatulate in outline, the cutting edge without cusps or
lobes, but a little higher at middle than at sides, posterior border
somewhat concave with faintly developed heel ; P with cutting
edge oblique, and crown more nearly terete. Lower incisors

FELIS 461

smaller than corresponding upper teeth, their crowns rounded
and flattened, with faintly indicated longitudinal groove.
Canines simple, without cingulum, rather large, the shaft curved
very slightly backward, that of upper tooth terete with a few
longitudinal grooves in enamel, that of lower slightly hollowed
on inner and posterior surface. Anterior upper premolar about
as large as outer incisor, though with crown not so high, its root
single, but often showing indications of longitudinal division.
Its usual position is a little behind middle of wide diastema
between canine and first large premolar. Other premolars
(except carnassial) essentially alike in form, two-rooted, the
crowns compressed, triangular when viewed from the side, some-
what longer than high, the base elliptical, slightly wider
posteriorly than anteriorly, that of pm^ wider than the others
and somewhat inclined to assume a concavo-convex outline ;
anterior border of pm'-^ simple, that
of pm^ and pm^ with a small basal
cusp (better developed in latter than
in former) ; posterior borders of all
three teeth with two secondary cusps,
the basal formed by the rudimentary
cingulum, the second on base of main
cusp ; pm^ slightly smaller than the
others. Upper carnassial about twice
as long as broad, the outline of its
crown almost an isosceles triangle,
though inner side is slightly longer fig. 97.

than outer ; inner lobe at anterior Feiis sUvestris. Teeth. Nat. size.
border of crown, scarcely breaking the

general contour, its cusp low but well developed, terete ; outer
or main cutting portion of tooth with three cusps, the anterior
low and very similar to that of inner lobe, the median and
posterior of the usual carnassial type, the posterior cutting edge
nearly horizontal and meeting anterior commissure at an abrupt
angle (about 50°). Upper molar small, transversely compressed
imperfectly two-rooted, the crown about half as long as wide
without definite structure though showing traces of an outer and
inner cusp. Lower molar consisting of a sub-equal paraconid and
protoconid, the latter slightly the larger, the anterior and
posterior borders of tooth nearly vertical, the upper border
formed by the two sub-equal commissures, both of which slope
downward toward the middle, where they meet at an angle of
about 50° ; outer surface of tooth convex longitudinally, inner
surface rather deeply concave.

Bemarks. — Externally Felis sihestris is distinguishable from
domestic cats with well developed markings of the "striped
tabby " type* by slightly greater size and by the somewhat

* For figures of the two main types of marking in domestic cats see
Pocock, Proc. Zool. Soc, London, 1907, pis. viii and ix.

462

CARNIVORA

longer fur, this giving the animal the appearance of mui'e robust
legs and less tapering tail. The skull and teeth differ only in
the greater average size of fully adult individuals, a character

Fig. 98.
Pelis silvestris (upper figure) and /'. cattis (lower figure). Ka.t. size.

most readily appreciable in the teeth. In twenty European
specimens of Felis silvestris and a like number of F. cafus the
extremes for certain dental measurements are as follows : —

Combined length of upper carnassial silvestris. catus.

a,ndp7n^ 16-6 to 20-0 .. 15-0tol7-8

Combined length of lower cheek-teeth 18-8 ,, 23-6 .. 18-0 ,, 20-4

Lowermolar 7-8 ,, 100 .. 6-6,, 8-6

Felis silvestris silvestris Schreber.

1758. [Felis] catus Linnseus, Syst. Nat., i, 10th ed., p. 42 (part; synonyms

only, the description refers to the domestic cat).
1777. Felis {Catus) silvestris Schreber, Siiugthiere, in, p. 397.
1777. \_Felis catus] ferus Erxleben, Syst. Regni Anim., i, p. 518.
1857. Felis catus Blasius, Saugethiere Deutschlands, p. 162 (Not of

Linnseus).
1896. Catus ferox Marfcorelli, Atti Soc. Ital. Sci. Nat., Milano, xxxv, p. 253,

January, 1896 (Accidental renaming of Catus ferns Brehm, Illustr.

Thierleben, i, p. 275, 1863).
1907. Felis sylvcstris Pocock, Proc. Zool. Soc, London, p. 150, June 12,

1907.
1910. Felis silvestris Trouessart, Faune IMamm. d'Europe, p. 98.

Tyjje locality. — Germany.
Geographical distrihution.-

-Central and southern continental

FELIS 463

Europe from northern Spain to northern Germany, and from
the Atlantic coast eastward at least to the Black Sea ; eastern
limits of range not known.

Characters. — Teeth not unusually large (combined length of
upper carnassial and pm^ 16 "G to 19 mm. ; three lower cheek-
teeth together, 18 '8 to 22 mm. ; lower molar, 8"0 to 8*8 mm.) ;
g^eneral colour light, the predominant hue approaching the smoke-
grey of Ridgn^ay ; dark markings on sides and legs tending to be
faint, brownish and ill-defined.

Measurements. — Adult male from northern Germany : head
and body, 545; tail, 310; hind foot, 135; ear from meatus, 63.
A specimen (not sexed) from Ingelheim, Rheinhessen, Germany :
head and body, 481; tail, 309; hind foot, 119. Adult from
near Athens, Greece : hind foot, 127. For cranial measurements
see Table, p. 466.

Specimens examined. — Fifteen, from the following localities : —

Spain : Province of Burgos, 3 (B.M. and U.S.N.M.).

Fbance : Manonville, Meurthe-et-Moselle, 1 ; Salavou, Haute-Marne, 1 ;
Caterille, Haute-Garonne, 1.

Germany : North Germany (no exact locality), 1 ; Ingelheim, Rhein-
hessen, 1; south Germany (no exact locality), 3 (skulls).

Austria-Hungary: Baranza, Hungary, 1 (skull).

Bulgaria: Varna, 1 (Andersen, skull).

Greece : Near Athens, 1.

Italy : Near Rome, 2.

Bemarks. — The wild cat of central Europe shows little
individual variation in colour, or in pattern and character of
markings. The two skins from Rome and that from Athens
appear to be identical with more northern specimens. In the
skull from Varna, Bulgaria, the nasals are not narrowed as in
the form occurring in Asia Minor and the Caucasus.

2. Prov. Burgos, Spain. S. and N. Gonzalez 8.7.7.10-11.

(c).
6. Manonville, Meurthe-et- Dr. E. Hamilton 95. 11. 9. 1.

Moselle, France. (Lomoiit.) (p).
6. Salavon, Haute - Marne. Dr. E. Hamilton 95. 11. 9. 2.

{Loniont.) (p).

i. Caterille, Haute-Garonne. O. Thomas (p). 8. 7. 15. 1.

{A. Robert.)

Lord Lilford (p). 95. 5. 1. 1.
Dr. A.Giinther (c). 59. 9. 6. 55-57.
G. Barrett-Hamil- 11. 1. 2. 104.

ton (p).
Dr. E. Hamilton 2. G. 3. 1.

C. W. L. Merlin 47. 7. 22. 2.

6. N. Germany.
3 skulls. S. Germany.

1. Ingelheim, Rheinhessen

Germany. {Erlanger.)
6. Baranza, Hungary.

1. Athens, Greece.

464 CARNIVOEA

Felis silvestris grampia Miller.

1907. Fdis grampia Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 396,

November, 1907. Type in British Museum.
1910. Felis grampia Trouessart, Faune Mamm. d'Europe, p. 99.

TyjJe locality. — Invermoriston district, Inverness, Scotland.

Geographical distribution. — Formerly throughout Great Britain;
now restricted to the wilder portions of Scotland.

Diagnosis. — Like Felis silvestris silvestris but general colour
darker, the predominant hue approaching the broccoli-brown of
Ridgway ; dark markings on sides and legs tending to be
extensive, blackish and well defined.

Colour. — Underfur of back and sides a light ochraceous-buff,
the basal half of the hairs mouse-grey. Light annulations of
longer hairs very nearly the cream-buff of Ridgway. Black tips
to longer hairs more noticeable than in Felis silvestris, and general
effect of ground colour distinctly browner, the light colour not
producing any frosted appearance. Upper side of feet and
inner surface of hind legs ochraceous-buff, becoming duller and
somewhat drab-tinged on under side of body. Intercrural and
pectoral white areas well defined and strongly contrasted with
surrounding colour. Black mottling on middle of chest con-
spicuous. Soles and palms blackish. Dark markings on tail, legs
and upper parts similar to those of Felis silvestris in arrangement,
but more definite in outline, particularly the transverse stripes
on outer side of forelegs and those on posterior half of body.

Skull and teeth. — The skull and teeth are not distinguishable
from those of typical Felis silvestris.

Measurements. — Type (young-adult male) : head and body,
534 ; tail, 338 ; hind foot, 127 ; ear, 67. Young-adult male and
female from the type locality : head and body, 534 and 534 ; tail,
280 and 290; hind foot, 126 and 123; ear from meatus, 67
and 68.

Specimens examined. — Twenty-two, from the following localities in Scot-
land : Glencassley, Sutherland, 2 ; Lochcarron, Eoss, 2 ; Aberdeenshire, 1 ;
Invermoriston district, Inverness, 5 ; Beauly, Inverness, 1 ; Fort William,
Inverness, 2 ; Knoydart, Inverness-shire, 2 ; Arisaig, Inverness-shire, 1 ;
Braulea Forest, Inverness-shire, 1 ; Balmachan, Inverness-shire, 1 ; Jardine
House, Dumfriesshire, 1 ; ^Inverness (no exact locality), 1 ; Scotland (no
exact locality), 2.

BemarJcs. — The British wild cat differs from true Felis
silvestris in darker general colour and more pi'onounced black
markings, characters in which it approaches F. s. tartessia. It
does not show, however, any tendency to the enlargement of
teeth so noticeable in the south-Spanish form.

skuU. Glencassley, Sutherland- E. R. Alston (p). 79. 9. 25. 81.

shire, Scotland.
6. Glencassley, Sutherland- Mrs. Flower (p). 92. 10. 21. 1,

shire.

FELIS 465

3 6, ?. Invermoriston, Inverness- A. H. Cocks (p). 4. 1. 25. 2-5.
shire. (4. 1. 25. 3. Type of subspecies.)

6 st. Invermoriston, Inverness- Hon. N. C. Roths- 1. 5. 22. 4.
shire. child (p).

1. Beauly, Inverness-shire. Lord Tweedmouth 5. 10. 12. 1.

2 6. Knoydart, Inverness-shire. Sir Herbert Maxwell 98. 12. 2G. 1.

(p). 99. 2. 9. 1.

1. Arisaig, Inverness-sbire. Dr. E. Hamilton (p). 2. 6. B. 4.

9. Braulin Forest, Inverness- G. W. Henderson (p). 6.12.18.1.

shire,

st. Balmachan, Inverness-shire. J. E. Harting (p). 83. 11. 13. 1.

st. Jardine House, Dumfries. Sir W. Jardine (p). 86. 7. 2. 13.

1. Scotland. 114b.

Felis silvestris tartessia Miller.

1907. Felis tartessia Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 397,

November, 1907. Type in British Museum.
1910. Felis tartessia Trouessart, Faune Mamm. d'Europe, p. 99.

Type locality. — Goto Donana, Huelva, Spain.

Geographical distribution. — Southern Spain ; limits of distri-
bution not known.

Diagnosis. — Like Felis silvestris silvestris but larger and with
disproportionately larger teeth. Colour darker, and black
markings essentially as in F. silvestris grampia.

Colour. — The colour is noticeably darker than in Felis
silvestris silvestris, rather nearly approaching that of F. s. grampia
but less brownish. Underfur more slaty at base than in the
related forms, about the grey No. 6 of Ridgway, its terminal
portion a dull cream-butf. Pale annulations on longer hairs
nearly as whitish as in F. silvestris silvestris. Inner surface of
hind legs a light ochraceous-buff ; rest of underparts as in
F. s. grampia. Dark markings well defined, their arrangement
and extent as in the British wild cat.

Shtll. — In fully adult males the skull is very large, apparently
exceeding that of any of the other members of the group. Its
form, however, is not peculiar.

Teeth. — The teeth of the Spanish wild cat are noticeably
larger than those of any of the other members of the Felis
silvestris group, or of the domestic cats. The difference is
particularly evident in the premolars both above and below and
in the lower molar (see measurements, p. 467). In form, how-
ever, the teeth show no peculiarities.

Measurements. — Type (adult female) from well-made skin :
head and body, 650 ; tail, 350 ; hind foot, 133. For cranial and
dental measurements see Tables, pp. 466 and 469.

Specimens examined. — Eight, from the following localities in southern
Spain : Sierra Morena, 1 ; Goto Donana, Huelva, 5 ; Andalucia (no exact
locality), 2 (skulls).

2 H

466

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468 CARNIVORA

BemarJcs.—The Aiulalusian form of Fcli>^ siJvcstris is well
characterized by its lai-ge teeth and rather dark colour.

1. Goto Donana, Huelva, A. Chapman (c & p). 7. 6. 4. 1.

Spain. {Type of subspecies.)

1. Goto Donana, Huelva. A. Chapman (r). 8. 3. 8. 1.

6, 9. Andalucia. {Chapman.) Dr. E. Hamilton (p). 2. 6. 3. 2-3.

FELIS SARDA Lataste.

1885. [Felis libyca^ var. sarda Lataste, Actes Soc. Linn., Bordeaux, xxxix,
p. 231 (p. 109 of reprint), September, 1885. Type in Lataste
collection.

1896. Felis mediterranea Martorelli, Atti Soc. Ital. Sci. Nat., Milano
XXXV, p. 266, January, 1896 (Sardinia).

1910. Felis ocreata sarda Trouessart, Faune Mamm. d'Europe, p. 101.

Type localiti/. — Sarrabus, Sardinia.

Geograjjhical distribution. — Sardinia, and according to Mar-
torelli portions of the mainland of western Italy.

Diagnosis. — Differs from Felis silvestris and resembles the
African F. ocreata in the shorter fur and more slender tail
(average length of hairs at middle about 30 mm. instead of
40 mm.) ; hairs of median dorsal line slightly elongated and
stiffened, forming a faint ridge or rudimentary mane ; dark
maikings obsolete, the back and sides greyish or brownish,
without definite stripes.

Colour. — Underfur drab-grey through basal half, the terminal
half of hairs ochraceous-buff. Pale annulations nearly white.
The pale annulations and black tips form a clear grey grizzle in
which the ochraceous-buff scarcely appears except when fur is
disarranged. Pale annulations on elongated dorsal hairs dull
buff, giving this region a brownish cast noticeably different from
sides, but not producing any suggestion of the clearly defined
stripe characteristic of the members of the silvestris group. No
stripes on nape, shoulders or sides, but the position of the
typical markings faintly indicated by shadings in the grizzle.
Dark markings on legs as in the silvestris grou^. Back of ear
yellowish clay-colour, the extreme tip black. Feet pale, dull
buff above, black below. Inner surface of hind legs bright
ochraceous-buff, fading toward buff on chest and anterior portion
of belly. Dark mottling on chest moderately distinct. Tail
with well defined black tip, but with other markings obscure,
the general effect essentially like mid-dorsal area.

Skull. — The skull closely resembles that of Felis silvestris,
except that in the few specimens examined the auditory bulla3 are
higher and more inflated anteriorly, a difference which may
prove to be purely individual.

Teeth. — The teeth are larger than in true Felis silvestris,
nearly equalling those of the Spanish form.

Measurements. — Type (adult male), from skin : head and

469

DENTAL MEASUREMENTS OP FELIS SILVESTBIS
AND F. SARD A.

Locality.

Number.

Sex.

Pm.3

and

ptn*

together.

Tliree
lower
cheek-
teeth.

Lower
molar.

F. silvestris silvestris.

France : Manouville, Muerfche-1
et-Moselle . . J

95.11.5.1

6

18-2

—

—

Salavon, Haute-Marne

95. 11. 5. 2

6

18-2

21-2

8-8

Caterille, Haute - Ga- 1
ronne . . ./

8. 7. 15. 1

6

17-6

20-2

8-0

Germany : southern Germany .

1143cZ

6

16-6

18-8

8-0

)) ))

1143/

6?

17-2

—

— .

Austria-Hungary : Baranza

8. 7. 7. 10

6

18-8

21-8

8-4

Bulgaria: Varna

Andersen

6

17-8

21-0

8-2

Greece : near Atlieus

47. 7. 22. 2

—

19-4

22-4

8-8

F, silvestris grampia.

Scotland : no exact locality

5. 10. 12. 1

—

17

6

20-8

8-0

)) »)

1146

—

18

0

20-4

8-0

Sutherland

79. 9. 25. 81

6?

18

4

21-6

8-4

Port William .

99.2.9.1

6

17

4

21-0

8-0

Inverness-shire

Zool. See.

6

18

0

21-0

8-2

,1

4. 1. 25. 2

6

18

6

21-2

8-0

)j

4. 1. 25. 3*

6

16

8

21-0

8-0

)»

4. 1. 25. 4

6

18

0

21-0

80

>>

98. 12. 26. 1

6

17

6

21-0

8-0

)>

4.1.25.5

?

16

6

19-8

7-8

))

6. 12. 18. 1

9

16

8

19-0

7-8

F. silvestris tartessia.

Spain : Sierra Morena

--

— .

19

0

22-4

9-2

Goto Douana, Huelva .

7. 6. 4. 1*

6

19

8

23-6

9-4

?> )j •

2. 6. 3. 2

6

19

8

—

—

)> )i

2. 6. 3. 3

?

19

8

23-4

10-0

)j ))

8. 3. 8. 1

—

20

0

22-8

9-2

F. sarda.

Sardinia : Sarrabus

88. 12. 1. 1

S

19-4

22-2

9-0

Type.

470

CARNIVORA

body, 600; tail, 300 ; hind foot, 124. For cranial and dental
measurements see Tables, pp. 466 and 469.

Specimens examined. — Sixteen, all from Sardinia (B.IM., Turin, Genoa,
and Lataste).

6. Ogliastra, Sardinia. {Dr. 0. Thomas (p). 0. 5. 2i. 1.

Woltcrstorff.)

6. Sarrabus. '" Marquis G. Doria(p). 88.12.1.1.

1. Sardinia. Purchased (Linnsea, 86. 7. 10. 1.

Berlin.)

FELIS AGRIUS Bate.

1906. Felis ocrcata agrius Bate, Proc. Zool. Soc, London, 1905, ii, p. 317,

April 3, 1906.
1910. Felis ocreata agrius Trouessart, Paune Mamm. d'Europe, p. 102.

Type locality. — Crete.

Geographical distribution. — Crete.

Diagnosis. — Similar to Felis sarda, but paler and more yellow-
in general colour, and dark markings of shoulders and back less
obsolete.

Colour. — The elements of the colour are essentially as in
Felis sarda, but the ochraceous-buff of underf ur is not so dark,
and the black tips and annulations of the longer hairs are not so
extensive. As a result, the general colour is a yellowish grey,
made up chiefly of a mixture of the ochrac(}ous-buff and the
whitish sub-terminal annulations, very slightly darkened by the
black. While there are no black markings on body or legs,
the longitudinal shoulder stripes and dorsal stripe, as well as
the transverse lateral stripes, are clearly indicated, when fur is
smooth, by brownish shades. Tail with black tip and two or
three black sub-terminal rings. Outer surface of ear dull buffy
clay-colour, becoming blackish at tip. Feet buffy above, darker
and somewhat clouded with blackish below. Inner surface of
hind legs and middle of chest light ochraceous-buff"; belly
slightly darker than in Felis sarda, the mottling obsolete.

Skull and teeth not known.

Specim,ens examined. — Two skins, both from Crete.

2. Kanea, Crete. Miss D. Bate (c). 5. 12. 2. 14-15.

(5. 12. 2. 15. Type of species.)

Genus LYNX Kerr.

1792. Lynx Kerr, Anim. Kiugd., Systematic catalogue inserted between

pages 32 and 33. (Type by tautonymy Lynx vulgaris Kerr =

Felis lynx Linnaeus.)
1821. Lynceus Gray, London Med. Repos., xv, p. 302, April 1, 1821 {Fdis

lynx Linnoeus).
1829. Pardina Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,

p. 53 {Felis pardina Temminck = Lynx pardelhis Miller).

LYNX 471

1834. Lynchus Jardine, Naturalists' Library, Mamm., ii, p. 274 {Felis lynx
Liniiffius).

1857. Lynx Blasius, Saugethiere Deutschlands, p. 161 (Sub-genus of Felis).

1867. Ccrvaria Gray, Proc. Zool. Soc, London, p. 276 (Lyncus pardinus =
Lynx pardellus Miller ; type fixed by Miller and Rehn, Proc.
Boston Soc. Nat. Hist., xxx, p. 199, December 27, 1901). Not of
Walker, 1866.

1903. Eucervaria Palmer, Science, N.S., xvii, p. 873, May 29, 1903 (Substi-
tute for Gervaria Gray).

Tijj}c species. — Felis lynx Linna?us.

Geographical distribution. — Northern forested portions of both
Old and New Worlds ; in Europe south to the Mediterranean
coast and west to the Atlantic (perhaps within historic times to
Great Britain).

Characters. — Similar to Felis but without small upper pre-
molar, form heavier and less typically feline, ears tufted.

Bemarks. — As in the case of the genus Felis the limits of the
genus Lynx are not clearly understood. As here restricted the
group includes about a dozen named, but for the most jaart
imperfectly known forms peculiar to the Northern Hemisphere.
Two of them occur in Europe.

KEY TO THE EUROPEAN SPECIES OF LYNX.

Length of lower cheek-teeth together about 85 mm. ;
skull relatively long and low, the interorbital
convexity moderate ; back and sides never thickly
spotted, usually without dark markings ; cheeks
not conspicuously whiskered (Northern and cen-
tral Europe) L. lynx, p. 471.

Length of lower cheek-teeth together about 30 mm. ;
skull relatively short and deep, the interorbital
convexity abrupt ; back and sides thickly spotted
(Iberian Peninsula) L. jjcirdcllits, p. 475.

LYNX LYNX Linmi^us.

1758. [Fells'] lynx Linnaeus, Syst. Nat., i, 10th ed., p. 43 (Sweden).

1792. Lynx \mlgaris Kerr, Anim. Kingd., Systematic catalogue inserted

between pp. 32 and 33 ; described, p. 157.
1792. Lynx vulgaris alba Kerr, Anim. Kingd., Systematic catalogue inserted

between pp. 32 and 33; described, p. 157 (Forests of Sweden).
1792. ? Lynx vulgaris melina Kerr, Anim. Kingd., Systematic catalogue

inserted between pp. 32 and 33 ; described, p. 157 (Banks of the

Volga near Gazan, Russia).
1798. -P[eZis] borealis Thunberg, Beskrifning pa Svenske Djur, IMamm.,

p. 14 (Heavy forests of northern Sweden).
1798. Felis kattlo Schrank, Fauna Boica, i, p. 52 (Bohemia).
1820. Felis lyncula Nilsson, Skand. Fauna, i, p. 14 (Wooded and moun-
tainous regions of Scandinavia).
1825. Felis lupulinus Thunberg, Denkschr. k. Akad. Wissensch., Miinchen,

IX, p. 189 (Northern Scandinavia).
1825. Felis vulpinus Thunberg, Denkschr. k. Akad. Wissensch., Miinchen,

IX, p. 192 (Near Upsala, Sweden).

472

CARNIVORA

1829. Felis virgata Nilsson, lilum. Fig. Skand. Fauna, pis. .3 and 4
(Wooded and mountainous portions of Sweden).

1857. Felis lynx Blasius, SJiugetbiere Deutschlands, p. 173.

1870. Lynx ccrvaria Fitzinger, Sitzungsber. kais. Akad. Wissensch., Wien,
Math.-Naturwiss. Classe, lx, Abth. i, p. 208 (Not Felis ccrvaria
Temminck, 1827).

1904. [Lynx] lynx Trouessart, Catal, Mamm., Suppl., p. 276.

1910. Lynx lynx Trouessart, Faune Mamm. d'Europe, p. 103.

Type locality. — Near Upsala, Sweden.

Geographical distribution. — Wooded poi'tions of Europe from
the extreme north to the Alps and Pyrenees, and from the
Atlantic coast eastward. Now practically exterminated except
in the wilder portions of the Scandinavian Peninsula.

Diagnosis. — Length of lower cheek-teeth together about
35 mm. ; skull relatively long and low, the interorbital convexity
moderate ; back and sides never thickly spotted, usually without
dark markings ; cheeks not conspicuously whiskered ; underfur
dense and woolly.

External characters. — Form heavier and more dog-like than in
Felis silvestris, the legs relatively longer and feet more robust,
the tail decidedly less than half as long as head and body. Ear
conspicuously tufted at tip, and hairs on side of head below ear
and behind angle of jaw somewhat elongated. Digits and
tubercles as in Felis silvestris, the claw on thumb especially well
developed, the horny excrescence on palm near wrist wider at
base ; hairy covering of both palm and sole woolly rather than
velvety in texture. Tail shorter than hind foot. Underfur
evei'ywhere densely woolly.

Colour. — Upper parts and sides varying from yellowish brown
to brownish grey, the back usually frosted by white hair-tips
except along median region, and sometimes rather thickly
sprinkled, especially in immature individuals, with small black
spots or short streaks tending to arrange themselves in about five
longitudinal rows, but spotting never so conspicuous as in Lynx
pardellus. Head like back, the face with a few indistinct
longitudinal dark streaks ; a whitish eye ring about 5 mm. wide
interrupted in front and behind ; median portion of upper eyelid
black ; a small black spot on side of head below ear ; inner
surface of ear whitish ; outer surface like crown on basal half,
but posterior border and tip, including pencil, black, the inter-
mediate region whitish ; contrast of whitish area sometimes
heightened by a dai'k shade along its lower border. Underj^arts
ranging from whitish to pale bluff". Feet intermediate between
back and belly ; front of foi-earm and of thigh usually with a few
black or dark brown specks ; tail and posterior surface of thigh
somewhat darker than upper parts, often with a slight rusty tinge,
the tip of tail black. Undei'fur usually brownish or buffv' to
extreme base, but occasionally grey.

Skull. — The skull of Lynx lynx differs in general aspect from

LYNX 473

that of Felts silvestris ia much greater size and massiveness
(condylobasal length of skull in adult males exceeding 140 mm.)

Fig. 99.
Lynx lynx, x J.

and relatively wider interorbital region (interorbital breadth
fully equal to diameter of orbit) ; the zygomata are, however

474

CARNIVORA

somewhat less abruptly spreading. Dorsal profile convex
throughout, somewhat more so in front than behind, essentially
as in Fclis silvestrls. Other general features so nearly as in the
wild cat as to require no detailed description. A few details,
however, are peculiar. Auditory bullae relatively smaller than in
FeJis silvestris, the space between them fully as wide as bulla,
the surface less closely applied to mastoid and paroccipital
processes. Postorbital process seldom, if ever, becoming ligulate,
the apex remaining sharply pointed. Anteorbital foramen about
as large as in Felis silvestris, therefore relatively smaller, its
position slightly more posterior, over posterior root of pm^ or
space between roots of pm^ and carnassial. Nasal branch of
premaxillary very narrow, its width at front of nasal barely
one-fifth that of nasal in same region. Incisive foramina
opposite middle of canine, their anterior border not reaching
level of space between canines and incisors. Lateral emargina-
tion of posterior boi'der of palate very slight, extending scarcely
to level of front of molar. External pterygoid process reduced
to a mere ridge. Mandible with coronoid process relatively
shorter than in Felis silvestris, the distance from alveolus of
molar to extremity of process about equal to that from back of
molar to middle of canine ; angular process rather noticeably
bent inward.

Teeth. — ^In general structure, number and arrangement
of cusps the teeth exactly resemble those of Felis silvestris,
except that the posterior border of m^ is distinctly oblique and
armed with a minute though evident basal cusp slightly above
the rudimentary cingulum. All the cheek-teeth are i-elatively
longer, lower, wider, and less trenchant than in the wild cat,
and the cingulum at posterior border of lower premolars is fre-
quently less distinct, scarcely forming a cusp.

Measurements. — Adult from Valais, Switzerland (skin) :
head and body, 800± ; tail, 110± ; pencil, 30. For cranial
measurements see Table, p. 478.

Specimens examined. — Thirteen, from the following localities :—
Norway : No exact locality, 3.

Sweden: No exact locality, 2 (B.M. and U.S.N.M.).
Lapland: No exact locality, 1 (Genoa).

Switzerland : Near Geneva, 3 (Geneva) ; Alpes Vaudoises, 1 (Geneva) ;
Valais, 1 (Geneva).

Italy : Piedmont Alps, 2 (Genoa).

MemarTcs. — The material seen is insufficient to form the basis
of any discussion of the local races of the European Lynx.

1. Sweden. Purchased. (Brandt.) 51. 11. 8. 16.

St. Norway. Zoological Society's 55. 12. 24. 273.

Museum.
1. Norway. Zoological Society's 58. 5. 4. 63.

IMuseum.
1. Norway. Zoological Society's 69. 10. 19. 16.

Menagerie.

475

LYNX PARDELLUS Miller.

1827. Fells pardina Temminck, Monogr. de Mamm., i, p. 116 (Near

Lisbon, Portugal). Not Lynx pardina Oken, 1816.
1907. Lynx pardclla Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 398,

November, 1907 (Goto Douana, Huelva, Spain). Type in British

Museum.
1910. Lynx {Eucervaria) pardclla Trouessart, Faune Mamm. d'Europe,

p. 105.

Type locality. — Goto Donana, Huelva, Spain.

Geographical distribution. — -Iberian Peninsula.

Diagnosis. — Length of lower cheek-teeth together about
30 mm. ; skull relatively short and deep, the interorbital
convexity conspicuous and abrupt ; back and sides always
densely spotted ; cheeks noticeably whiskered.

External characters. — Form less robust than in Lynx lynx,
but not differing in essential details. Underfur not densely
woolly. Elongated hairs on sides of head below ears forming
conspicuous pointed tufts 50 to 80 mm. in length, separated from
each other along median line by a narrow area of ordinary fur.

Colour. — Underfur pale buff at base, the hairs becoming pale,
dull tawny on distal half. Longer hairs bufFy at base, black at
tip, each with a whitish sub-terminal annulation about 5 mm.
long. Ground colour above a fine mixture of the whitish and
dull tawny, very uniform throughout body, neck, outer surface
of fore leg and entire hind leg. On face it becomes greyish, and
on feet there is usually a wash of cream-buff. In the more usual
type of coloration the entire back and sides are closely sprinkled
with rounded black spots, mostly about 10 mm. or less in
diameter, the spots tending to become confluent so as to suggest
two longitudinal dorsal stripes, and also though less clearly
several transverse stripes on sides and shoulders. Although it is
impossible to count the spots along back, there are evidently more
than twenty-five between base of tail and most distinctly indicated
transverse stripe on shoulder. On legs the spots become slightly
larger, and on neck smaller and less distinct. In a less usual
colour phase the spots are much larger and less numerous, many
of those on back 20 mm. in diameter, the number between tail
and most distinct shoulder stripe only about a dozen. Under-
pai'ts creamy-white, heavily blotched with black on belly ; throat
more buffy, its spots very small ; interramial space nearly white.
A V-shaped black mark, open in front, extends from hinder
portion of interramial space to ends of whiskers. Inner surface of
fore legs like chest ; a heavy black baud at elbow and a faint one
at wrist and at middle of upper arm. Whiskers mixed black
and white. Ear and its long pencil black, the conch buffy along
the anterior rim and at base, the posterior rim greyish at base,
then with a black line 5 mm. wide jDarallel to edge. A broad,
sub-triangular grey area on back of ear. Inner surface clothed

476

CARNIVORA

with long creamy-white hairs. Tail like back above, clear,
light buffy below, the entire tip (about 40 mm.) black; a few
black spots on upper surface which tend to arrange themselves
in three transverse bands.

Fig. 100.
Lynx pardellus.

Skull. — The skull of Lynx pardellus differs conspicuously from
that of L. lynx and Fells silvestris in the high, flattened inter-
orbital region and anterior portion of brain-case, and abruptly

LYNX 477

sloping rostrum, the two surfaces forming an evident angle in
dorsal profile ranging from 40° to 50°. Aside from this very
noticeable peculiarity in general form the skull agrees with that
of L. lynx in all of the special features in which the latter
differs from that of the wild cat. The ridge rejDresenting the
external pterygoid process appears to be somewhat better
developed, however, than in L. lynx. Temporal ridges rarely if
ever uniting to form a sagittal crest in front of region of
intei'parietal.

Teeth. — The teeth are intermediate in form between those of
Lynx lynx and Felis silvestris. Cingulum at posterior border
of lower premolars obsolete as in L. lynx, but posterior border of
m^ nearly as vertical as in Felis silvestris, and without secondary
cusp or evident cingulum. Crowns relatively a little narrower
and more trenchant than in Lynx lynx, but less so than in the
wild cat, their length relatively as great as in the lynx.

Measurements. — Two males from Goto Doiiana, Huelva, Spain
(skins): head and body, 880 and 930; tail, 125 and 130; hind
foot, 170 and 180; ear from meatus, 73 and 80; pencil, 30
and 55. For cranial and dental measurements see Tables,
pp. 478, 479.

Specimens examined. — Eighteen, from the following localities in Spain
Old Castile, no exact locality, 1 ; Sierra Morena, Cordova, 3 ; near Seville, 3
Jerez de la Frontera, 1 ; Coto Dofiana, 8 ; Andalucia, no exact locality, 1
no exact locality, 1.

jRemarJcs. — Specimens in the two colour phases differ so
noticeably from each other as to suggest the existence of distinct
forms ; and as yet I have seen no skins which could not at once
be referred to one or the other. The two occur together, how-
ever, and there are no correlated peculiarities in the skulls and
teeth.

9. Old Castile, Spain. Lord Lilford (p). 94. 6. 11. 1.

1. Sierra Morena, Cordova. Capt. S. E. Widdriug- 42.2.26.1.

ton (c & p).

2 6. Sierra Morena, Cordova. Lord Lilford (p). 89. 8. 27. 1.

94. 6. 11. 1.

1. Seville. Lord Lilford (p). 74. 10. 7. 1.

6. Seville. {Dr. A. Ruiz.) Lord Lilford (p). 95. 9. 4. 1.

1. Jerez de la Frontera. A. Williams (c & p). 7. 12. 10. 1.

2. Coto Dofiana, Huelva. J. P. Gassiot (c & p). 72. 10. 26. 1-2.
head, st. Coto Dofiana, Huelva. A. Williams (c & p). 3. 3. 16. 1.

2 c5, 2 9. Coto Dofiana, Huelva. A. Chapman (c & p). 4. 12. 12. 1-2.

8. 3. 8. 2-3.
(4. 12. 12. 2. Ty2?e of species.)
1. Coto Dofiana, Huelva. B. F. Bucli (c & p). 7. 6. 4. 2.
St. Andalucia. B. P. Buck (c & p). 6. 9. 16. 1.

1. Spain. Lord Lilford (p). 74. 9. 4. 1.

Note. — A lynx from Nuoro, Sardinia, has recently been
described by Mela (Boll. Soc. Zool. Ital., Roma, 2d ser., ix,
p. 48, 1908) as Lynx sardinise. The more important characters

478

CARNIVORA

•AVOJ-tJlOO^

Ai o

3 ^

-^iOOC^CMOCMOtMQO

CD t- t-

•A\O.T-mOO^

ooofMtM-^oqoooo

lO '^ '^

■aiqipuupf

CO 05 05

•q^pBajq

piO^SBJf

mdap [i3J')so}£

o

O

o

iO

lO

CO

8

CO

^

o

o

o

CO

CM

CM

(M
CM

00

o

o

lO

CM

'SiCOOOOOCMOOCM

CMC300O3COiH<MCqiOCM

00000000000302000001

O O O O (M O -^

•q^jdap le'ivi'ei

JO mdaa

■^(MOCOCOOOCO

Tji O O

C33 CO >0

•S8UIUB0 J8A0

mpBajq iBj^soy:

00 ^ CM

OCOOOOCDOCM

•iionorj^suoo
IB^iqjo.ia^ni

o '* 00

<M CO 00 '

T-H T-H O ■
CO CO CM I

^ O oq

t- T-l (M
<M CO CO

•iiO!C(ou;suoo
IBiiqio^sod

CO "^ "^

o o
6 6

O00O^->*OCM-*

•q^p^ajq
oi^biuo3j(z

•qiSnat
IBSBqo[jfpuoo

CD '^ O)

ocooocoooqo

00 o o

CO 00 CO
'ei o ■<*<

9-H

o o

CM (M

-H

113 I

•^ 1-^ ir-i

CO ^ t~ g ^ J?

in > CO g: CO ^

: ?3

(M CM "^ CO

O

r-tCCCQr-l (BtM ^O <1>
^ COO <^0 "^O

rH oq ■^^ rH "^^ ^ 00 ;:;; _

cDcdCf5.d2cdc6<j,-<»g

-*■ CTJ !P . -^- t-: 00 '"' QO' ""I
CT, CO <^ t- g2 ^-

n

>,

J

a

<D

■^

ts

en

m

' o3

a
in 03 <o

^ -S S3

'-'< "Tli gj c3 „
.— I .S a> CD

a> CD CD

479

DENTAL MEASUREMENTS OP LYNX LYNX AND
L. PAEDELLUS.

Locality.

Number.

Se.K.

Upper

car-

nassial.

Pm3

and

pm*

together.

Lower
cheek-
teeth.

Lower

car-
nassial.

Lynx lynx.

Sweden

I 1034 \
\ U.S.N.M./

6?

18-6

30-8

37-6

16-0

...

B.M.

juv.

—

29-0

—

15-6

Switzerland

( 668.32 1
\ Geneva j

6?

19-4

31-0

37-2

15-0

„

( 223.37 1
\ Geneva j

9?

18-6

29-4

—

—

. .

i 603.64 )
\ Geneva j

juv.

17-0

—

—

14-0

Lynx pardellus.

Spain : Old Castile

94. 6. 11. 1

?

14-4

23-8

27-0

12-4

Sierra Morena

42. 2. 26. 1

juv.

15-4

26-0

30-8

13-6

.,

89. 8. 27. 1

6

16-0

26-2

30-8

13-0

near Seville .

95. 9. 4. 1

6

14-8

25-0

30-0

12-8

Jerez

7. 12. 10. 1

6?

14-0

23-6

28-4

12-4

Goto Donana .

72. 10. 26. 1

6?

14-6

24-4

29-6

12-8

r, ■ ..

7. 6. 4. 2

6

14-4

24-0

28-8

11-8

»

8. 3. 8. 2

6

16-0

27-0

32-6

13-8

,. ..

4.12.12.2*

9

15-0

25-0

29-2

12-2

)> )>

8. 3. 8. 3

?

14-2

24-2

28-8

12-4

„

152618

6?

14-6

24-6

28-8

12-8

.,

152619

6

16-2

26-4

30-8

13-4

Type.

480 CARNIVORA

mentioned are as follows : Length of head and body, 1000 mm. ,
tail, 330 ; height, about 350 ; cheeks distinctly whiskered, and
ears tufted ; back reddish, the dorsal area almost forming a band
(fascia) ; sides reddish gi-ey ; head, neck, shoulder and flanks
with reddish brown or greyish brown spots ; legs with transverse
tawny (fulvo) stripes ; tail with black tip and three black
sub-terminal rings ; head with black stripe on each side beginning
at mouth (fauci) and passing back through eye to side of neck ;
ear tawny inside, reddish outside ; tending toward black below,
the pencil (about 30 mm. long) reddish tending toward black ;
underparts and inner surface of legs dirty white with a reddish
tinge.

481

Order RODENTIA.

1758. Glircs Linnaeus, Syst. Not. i, 10th ed., p. 56 (part).

1827. Rodcntia Griffith, Cuvier's Animal Kingdom, Mamm., iii, p. 61.

Gcograjjliical distribution. — Continents and larger islands of
the entire world, New Zealand and the Antarctic excepted.*

Characters. — Terrestrial (occasionally semi-aquatic) placental
mammals, incapable of true flight (a parachute-like membrane
present in certain groups) ; feet unguiculate ; dentition consisting
of —j- functional scalpriform incisors (a second pair of minute,
terete, non-trenchant incisors present in upper jaw in the
Duplicidentata) growing from a persistent pulp, and a row of
from two to six tuberculate or laminate cheek-teeth, the crowns
of which show no distinct traces of tuberculo-sectorial structure ;
canines absent ; a wide diastema between incisors and cheek-
teeth.

BemarJcs. — The members of the order Bodentia are imme-
diately recognizable by the peculiarities of their dentition, no
near approach to which is found in any other group of living-
mammals. They are all relatively small animals, the largest
member of the group, the South American Hydrochoerus lujdro-
choeris, attaining a weight of only about 50 kg., while the largest
European species, the beaver, is less than half this size. In
structure they show variety exceeded by the Insect ivora only,
while in number of genera and species they far surpass all other
groups of mammals. About thirty families are recognized among
living rodents, eleven of these occur in Europe.

KEY TO THE EUROPEAN FAMILIES OF RODENTIA.

Upper incisors 2-2, the enamel extending to pos-
terior surface ; incisive foramina very large,
confluent ; bony palate much shorter than
mesopterygoid space ; fibula articulating with
calcaneum (Hares and Rabbits) Leporid.r, p. 484.

Upper incisors 1-1, the enamel coiifined to the front
surface ; incisive foramina moderate or small,
distinct ; bony palate longer than mesopterygoid
space ; fibula not articulating with calcaneum.

* IMembers of the genera Mus and Oryctolagus are now artificially estab-
lished on many of the remote islands from which the order was naturally
absent.

2 I

482

Anterior portion of zygomatic arch formed chiefly

by the jugal bone ; infraorbital foramen small.

Skull without postorbital processes ; molars

rootless ; tail broad and Hat (Beavers) Castoridx, p. 9-47.

Skull with postocbial processes ; molars rooted ;
tail terete.
Fore and hind leg not joined by a fold of

skin (Squirrels) Sciurichv,-p. 897.

Fore and hind leg joined by a fold of skin

acting as a parachute (Flying Squirrels) . Petaurisiidn-, p. 940.
Anterior portion of zygomatic arch not formed
chiefly by the jugal bone ; infraorbital fora,-
men large.
Jugal bone not supported by long zygomatic
process of maxillary : mandible with
angular part arising from outer side of
alveolus of incisor ; tibia and fibula dis-
tinct ; body covered with long quills
(nasal and frontal regions of skull in
European genus conspicuously inflated)

(Porcupines) HystricicLv, p. 542.

Jugal bone supported by long zygomatic pro-
cess of maxillary ; mandible with angular
part arising from under side of alveolus
of incisor ; tibia and fibula joined ; body
not covered with quills.
Lachrymal and malar bones in contact ; ante-
orbital foranaen much wider below than
above, its inner border with conspicuous,

nearly closed secondary canal Zapodidx, p. 535.

Lachrymal and malar bones not in contact ;
anteorbital foramen not wider below than
above, its inner border without secondary
canal.
External form excessively modified for
underground life ; skull wedge-shaped,
the supraoccipital sloping forward to
middle of brain-case, its area much
greater than that of parietals (Mole-
rats) SpalacidcV, p. 887.

External form not specially modified for
underground life ; skull not wedge-
shaped, the supraoccipital vertical or
nearly so, confined to ba'ck of brain-case,
its area much less than that of parietals.
Cheek-teeth (in European genera) | ;

cfecum absent (Dormice) Muscardinidse, p. 549.

Cheek-teeth (in European genera) § ;

caecum present Muridie, p. 591.

Molars prismatic, hypsodont, usually
rootless, their crowns flat (Voles

and Lemmings) MicroUnie, p. GIO.

Molars tuberculate, brachyodont,
rooted.
Tubercles on crowns of upper molars
arranged in two longitudinal

series Cn'cetinoe, p. 592.

Tubercles on crowns of upper molars
arranged in three longitudinal
series Murinx, p. 791.

DUPLICIDENTATA 483

ARTIFICIAL KEY TO EUROPEAN MEMBERS OP
RODENT FAMILIES.

(Based exclusively on the teeth.)

Incisors ^^ ; cheek-teeth *,'~'- Leporichr, p. 484.

Incisors \l\ ; cheek-teeth not more than ^•
Cheek-teeth '^^.

Crowns of mandibular cheek-teeth with con-
spicuous median concavity Sciuridic, p. 897.

Crowns of mandibular cheek-teeth without

conspicuous median concavity Petauristidie, p. 940.

Cheek-teeth less than ^j^.
Cheek-teeth ^-

4-4

Cheek-teeth brachyodont, rooted Muscardinid»' , p. 549.

Cheek-teeth hypsodont, rootless.

Enamel foldings transverse only Castoridx, p. 947.

Enamel foldings transverse, oblique and

longitudinal Ilystricidn', p. 542.

Cheek-teeth less than jj^.

Cheek-teeth '^^,^ Zapodidtv, p. 535.

Cheek-teeth ijl^.

Enamel foldings sigmoid Spalacidie, p. 887.

Enamel foldings not sigmoid MuridcV., p. 591.

Cheek-teeth hypsodont, prismatic MicroUnx, p. GIO.

Cheek-teeth brachyodont, tubercular.
Tubercles of upper molars arranged

in two main rows Cricetlnx, p. 592.

Tubercles of upper molars arranged

in three main rows Murinx, p. 791.

Sub-Order DUPLICIDENTATA.

1811. Duplicidcntata lUigcr, Prodr. Syst. Mamm. et Avium, p. 91

GeograpMcal distribution. — Essentially that of the order
Bodentia except that it does not include Madagascar or Australia.
One genus now almost universally distributed in the warmer por-
tions of both hemispheres as the result of artificial introduction.

Characters. — -Upper incisors 2-2, their enamel covering
extending to posterior surface, the second tooth minute, sub-
terete, without cutting edge ; distance between mandibular
tooth-rows much less than that between maxillary tooth-rows,
only one pair of rows capable of opposition at the same time, the
motion of the jaws in mastication consequently lateral ; premolars
^-^ ; incisive foramina very large, confluent posteriorly ; bony
palate reduced to a narrow bridge lying mostly between the
premolars ; facial portion of maxillary cribriform or incomplete ;
fibula articulating with the calcaneum.

Bemarhs. — The well defined sub-order'"" Duplicldentata contains

* Not improbably an order distinct from the Rodentia, as there is
reason to believe that the resemblances between the two groups are the
result of convergence rather than of relationship.

2 I 2

484 RODENTIA

two families, the Leporidse and Ochotonidse. Only the former,
including the hares and rabl)its, is rcjaresented in Europe.

Family LEPORID^.

1821. Lcporichv Gray, London Med. Repos., xv, p. 304, April 1, 1821.
1857. Leporina Blasius, Siiugethiere Deutschlands, p. 409.
1897. Lagidm Schulze, Helios, xiv, p. 82.

Geographical distribution. — As in the sub-order Dnplicidentata.

Characters. — Palatal bridge formed chiefly by the maxillary
bone ; supraoccipital processes present ; auditory bulhe rather
small, not inflated with spongy tissue ; clavicle rudimentary ;
hind legs elongated.

BemarJcs. — The members of the family Leporidde are now
arranged in ten genera,* two of which occur in Europe.

KEY TO THE EUROPEAN GENERA OF LEPOBIDJE.

Mesopterygoid region narrow, the width of space imme-
diately behind palate much less than least longi-
tudinal diameter of palate ; postorbital processes
(except in domestic races) slender, not distinctly
triangular in outline (Rabbits) Orijdolagus, p. 484.

Mesopterygoid region broad, the width of space imme-
diately behind palate greater than least longi-
tudinal diameter of palate ; postorbital processes
robust, their outline distinctly triangular (Hares) Leipus, p. 495.

Genus ORYCTOLAGUS Lilljeborg.

1758. Lepiis Linnaeus, Syst. Nat., i, 10th ed., p. 57 (part).

1857. Lepus Blasius, Siiugethiere Deutschlands, p. 410 (part).

1867. Cunicultis Gray, Ann. and Mag. Nat. Hist., 3rd ser., xx, p. 225

(Not of Brisson, 1762, or Wagler, 1830). Sub-genus of Leims.
1874. Orydolagus Lilljeborg, Sveriges og Norges Ryggradsdjur, i, p. 417

(Sub-genus of Lepus).
1899. Oryctolagus Major, Trans. Linn. Soc, London, 2nd ser., Zool., vii,

p. 514, November, 1899. Genus (part).
1904. Onjctolagus Lyon, Smithsonian Miscell. Coll., xlv, p. 402, June 15,

1904 (Genus),

Type species. — Lepus cuniculus Linnseus.

Geographical distribution. — ^Northern Africa and southern
and central Europe ; now artificially introduced and established
in many portions of the warmer region of both hemispheres.

Characters. — Externally similar to Lepus, but young at birth,
blind and essentially naked. Skull differing from that of Lepus
in the slender, never distinctly triangular postorbital process,

* For the most recent general work on the classification of the group
see Lyon: Classification of the Hares and their allies. Smithsonian
Miscell. Coll., XLiv, pp. 321-447, June 15, 1904.

ORYCTOLAGUS

485

the posterior liiiil) of which (except in certain domestic forms)
is alwcays free from cranium, persistence of sutures surrounding
interparietal, and the strongly narrowed choanpe, the width of
which, immediately behind palate, is noticeably less than least
length of palate. Teeth essentially as in Lepus, but incisors
both above and below less deeply implanted, and course of root
of upper tooth not visible on outer surface of premaxillary.
Cheek-teeth consisting of enamel cylinders tilled with cement
and dentine. In m-^ the cylinder is simple, its cross-section
elliptical. In the anterior upper premolar it is also simple, but
the anterior border is indented by longitudinal grooves appearing
as re-entrant angles on worn surface of crown. In all the other
cheek-teeth the cylinder is divided into two sections by a
re-entrant fold arising from inner side of maxillary teeth and
from outer side of mandibular teeth, in the former nearly
crossing the crown, in the latter completely crossing it, though
the two sections remain joined at their inner extremities
except in m.^, in which they are distinct throughout ; * anterior
section of first lower premolar indented by longitudinal grooves
on its anterior surface.

Hcmarhs. — The genus OrycioJagus contains the domestic
rabbits and the wild rabbit of central and southern Europe. In
addition to the moz'e technical characters of the group its
members are readily distinguishable from the other Leposidse,
with which they are naturally associated, by the blind, helpless
condition of the young at birth. f

ORYCTOLAGUS CUNICULUS Linnpeus.
(Syuouymy under subspecies.)

Geographical distribution. — Northern Africa and southern
and central Europe, the details of distribution considerably
modified by human agency.

Diagnosis. — Size smaller than in the European species of
Lepus (the Sardinian Lepus mediterraneits excepted), the hind
foot in dry specimens seldom exceeding 90 mm., the occipito-
nasal length of skull in largest individuals seldom more than
80 mm., ear from crown (dry) usually 70 to 80 mm., tail, including
pencil, about as long as hind foot, its under surface and edge
white, its upper surface blackish, usually grizzled with light
brown ; general colour above a grizzle of black and light bi-own ;
a distinct, dull buffy nape patch ; ear essentially concolor with
back though more finely grizzled, without distinct colour pattern,
the outer surface narrowly edged with black at tip ; underparts

* The structure of the teeth may be most clearly understood by
examining the roots.

t This character recurs in the American Sylvilagns and perhaps ako
in some of the less known genera.

486

RODENTIA

and inner surface of legs whitish except for dark inguinal
patches and collar.

External characters. — The external form is typically leporine,
the hind legs noticeably elongated, the ears long and the eyes
large ; soles of feet covered by a dense brush of somewhat
elongated and stiftened hairs which nearly conceal the large,
nearly straight claws ; digits, 5-4 ; front foot with third digit
longest, fourth and second successively shorter but not con-
spicuously so, fifth with tip of claw barely reaching base of
claw of fourth, first much shorter than the others, only the
claw projecting from integument of foot, its tip falling decidedly
short of level of base of claw of fifth. Mamm?e : p 1 — 1,
a 2-2 = 6.

Colour. — Upper parts a coai'se grizzle of cream-buff and black,
the cream-buff in excess on sides, the black usually in excess on
back, especially across loins. On rump the ground colour becomes
paler and on outer surface of legs slightly darker and with an
evident tinge of clay-colour, though in neither region forming
any decided contrast with surrounding parts. On parting the
fur of the back it is seen to have five colour bands : (1) a broad
grey (about Ridgway No. 6) basal area (13 mm.), the extreme
base somewhat paler ; (2) a brownish band (6 mm.) vai-ying in
exact colour between russet and light clay-colour and occupying
terminal portion of underfur ; (3) blackish (2 mm.), not sharply
defined below ; (4) cream-buff (3-4 mm.) ; (5) black (2-4 mm.),
the last three occupying terminal portion of longer hairs. In
addition to the underfur and ordinary longer hairs there are
others about 40 mm. in length entirely black on back (except for
grey basal area), and black with a cream-buff sub-terminal ring on
sides. These longest hairs are never conspicuously different from
the general fur as they are in Lepus. Nape patch a clear bi'own
intermediate between russet and clay-colour. Head essentially
like back but more finely grizzled, the cheeks not evidently
different from back, the pale eyering barely indicated. No light
spot between eye and muzzle or between eye and ear. Muzzle and
region from which whiskers spring a clear dull buffy clay-colour,
not strongly contrasted with rest of head. Ears not noticeably
contrasted with head or back, their colour pattern nearly obsolete,
though a faint trace of that so conspicuous in the hares is
indicated by a slightly darker line along middle of posterior inner
surface, and by a slight greyish tinge on basal half of posterior
outer surface. The black tip is reduced to a mere ill-defined rim,
sometimes obsolete and never more than 5 mm. wide, strictly
confined to posterior surface. Collar and inguinal patches con-
color with sides, the inguinal patches sometimes nearly meeting
in median line. Rest of underparts together with inner side of
legs buffy white or pale cream-buff. On hind legs the whitish
area extends over dorsum of foot to extreme tip of toes, though
sometimes suffused with the buffy brown of sides of feet. On

OKYCTOLAGUS 487

front legs it does not extend beyond wrist, the dorsal surface of
which is occasionally marked with whitish or white.

Skull. — -General form of skull slender, typically leporine, the
rostrum produced, the occipital region strongly bent downward.
Dorsal profile essentially straight from tip of nasals to front of
brain-case, then slightly convex and strongly deflected to posterior
edge of occiput ; ventral profile nearly parallel to dorsal profile,
but downward curve beginning further forward, about at level of
anterior extremity of postorbital process ; occiput flatly truncate
at right angles with axis of brain-case, therefore obliquely to
main axis of skull. Brain-case short-ovate in outline when
viewed from above, its anterior extremity sharply defined by
very deep postorbital constriction, its postero-lateral outline
somewhat distorted by slightly projecting mastoid and auditory
region. Interparietal rhomboidal or ligulate, its length usually
about half width. No sagittal crest ; lambdoid crest represented
by a low ridge which curves abruptly backward at level of outer
extremities of interparietal to form the outline of a sharply
defined, somewhat elevated, bony shield lying on dorsal surface of
supraoccipital and extending backward to edge of abrupt trunca-
tion ; the general outline of this shield is squarish, but the
posterior border is occasionally concave and the lateral borders
may be convex. Posterior aspect of brain-case a broadly
rounded arch slightly wider than high, the conspicuous, elongated
paroccipital processes descending below level of lip of large
foramen magnum, and closely applied to posterior surface of
auditory bulla}. Floor of brain-case marked transversely by
persistent suture between basioccipital and basi sphenoid ; behind
this suture the basioccipital widens slightly, then narrows
abruptly at front of condyles, its median region with wide
longitudinal groove ; auditory bulla flask-shaped, moderately
inflated, with well developed straight neck, the main axis of
bulla nearly parallel with posterior surface of occij^ut, though
directed slightly outward, the sub-circular meatus plainly visible
when viewed from above ; transverse diameter of bulla about
equal to that of basioccipital at region of greatest width between
bullce ; height including tube neax'ly twice transverse diameter.
Interoi'bital region flat, somewhat depressed, its width about
equal to that of rostrum a little in front of middle ; postorbital
process slender, crescentic, bent downward at each extremity, the
posterior limb decidedly longer than anterior limb though rarely
coming in contact with brain-case ; anterior limb occasionally
fusing irregularly with frontal, zygoma rather slender, its upper
edge narrow throughout and not flattened or everted along
anterior half as is usually the case in the European species of
Lepus. Rostrum slender, its depth to anterior rim of alveolus of
first cheek-tooth noticeably less than distance from latter point
to front of incisive foramen, nearly parallel-sided when viewed
from the side, tapering slightly when viewed from above or

488

Fig. 101.
Ori/etolayus cunicuhis. Nat. size.

ORYCTOLAGUS

489

below ; nasals flattened posteriorly, convex laterally in front,
their combined posterior margin deeply emarginate ; nasal branch
of premaxillary extending nearly to posterior border of nasal.
Outer margin of incisive foramen nearly straight, the outline of
the two together narrowly cuneate, the greatest combined l)readth
about one-third length. Least longitudinal diameter of palate
about two-thirds or three-quarters distance between alveoli of
anterior premolars. External pterygoid plate well develo^Ded.

Teeth. — First upper incisor with root extending somewhat
more than half way from aveolus to suture between premaxillai-y
and maxillary, its course not distinctly indicated on outer surface
of bone ; shaft of tooth about one-third greater in lateral
diameter than in antero-posterior diameter, its section nearly a
parallelogram in outline, though slightly wider internally than
externally ; anterior face of tooth with deep, simple groove lying
slightly nearer to internal than to external border ; posterior
face with wide, shallow longitudinal concavity, the beveled edge
abruptly angled near middle. vSecond incisor elliptical in cross
section, flattened antero-posteriorly, the area of its shaft in cross
section about one-sixth that of first tooth. Lower incisor witli
root extending back nearly or quite to level of front of alveolus
of anterior molar, its shaft essentially like that of first upper
incisor except that anterior groove and posterior concavity are
absent and beveled edge is nearly flat Anterior upper
premolar with crown narrowly elliptical in outline, the anterior
border with three re-entrant angles, the second of which is
deepest, usually extending to about middle of crown. Anterior
lower premolar irregularly squarish in outline, narrower anteriorly
than posteriorly, with two deep, rather wide grooves on outer side,
a narrow re-entrant angle at middle of anterior border and a
slight projection at middle of inner border ; anterior section of
tooth decidedly larger than posterior section, its posterior enamel
border forming a conspicuous, irregularly folded transverse ridge
slightly behind middle of crown. Posterior upper molar
essentially like second incisor. Posterior lower molar consisting
of a larger elliptical, transversely flattened anterior section, and
a smaller posterior subterete section, the enamel of each section
forming a complete tube, the area of crown about one-third that
of second molar. Second, third, fourth and fifth upper molariform
teeth essentially alike, though slightly decreasing in size from
before backward, the crown narrowly elliptical, the inner
and outer margins notched or flattened (varying according to
wear), the re-entrant enamel fold extending from inner border
nearly across crown and dividing it into essentially eijual halves,
its anterior border crenulate. Second, third and fourth lower
molariform teeth essentially alike in size and form, the crowns
much longer than in the corresponding upper teeth, each con-
sisting of two narrowly elliptical sections with very narrow,
almost pointed extremities, the anterior section decidedly larger

490

RODENTIA

than the posterior, the inner half of anterior border of smaller
section broadly applied to middle of posterior border of larger
section, the posterior enamel wall of anterior section forming a
high, simple transverse ridge completely crossing crown.

Remarks. — The well-known European rabbit needs no special
comparisons with other members of the fauna. In different
parts of its range it is represented by two local races.

KEY TO THE SUBSPECIES OF OBYCTOLAGUS CUNIGULUS.

Occipitonasal length of largest skulls, 78 to 82 mm. ;

hhid foot, 83 to 93 mm. (Central Europe) 0. c cunicultis, p. 490.

Occipitonasal length of largest skulls, 71 to 77 mm. ;

hind foot, 72 to 82 mm. (Mediterranean region,

Azores, &c.) 0. c huxleyi, p. 491.

Oryctolagus cuniculus cuniculus Linnreus.

1758. [Lcxnis] cuniculus Linnteus, Syst. Nat., i, 10th ed., p. 58.

1837. ? Lepiis vernicularis Thompson, The Athenaeum, p. 468. Nomen

nudum (Ireland).
1843. ? Lcpus vennicula Gray, List Spec. Mamm. Brit. Mus., p. 128.

Nomen nudum.
1857. Lcpus cuniculus Blasius, Siiugethiere Deutschlands, p. 426.
1867. Cuniculus fodiens Gray, Ann. and Mag. Nat. Hist., 3rd ser., xx,

p. 225, September, 1847 (Substitute for cuniculus). Type in

British Museum.
1904. 0[_ryctolagiis~] cuniculus Lyon, Smithsonian Miscell. Coll., xlv,

p. 406, June 15, 1904.
1910. Oryctolagus cuniculus Trouessart, Faune Mamm. d'Europe, p. 215.

Type locality. — Germany.*

Geographical distribution. — Central Europe north of the
Mediterranean region ; west to Ireland.

Characters.— iiize large (occipitonasal length of largest skulls,
78 to 82 mm. ; hind foot, 83 to 93 mm.) ; black of upper parts
usually producing an evident clouded effect.

Measurements. — Adult male and female from Clandeboye,
Belfast, Ireland : head and body, 420 and 408 ; tail, 47 and 63 ;
hind foot, 87 and 85 ; ear, 68 and 65. Adult male and female
from Tidmarsh, Berkshire, England : head and body, 408 and
392 ; tail, 63 and 64 ; hind foot, 92 and 86 ; eai', 70 and 70.
Adult female from Lezayre, Isle of Man : head and body, 437 ;
tail, 66 ; hind foot, 91 ; ear, 70. Two adult females from
Ingelheim, Rheinhessen, Germany : head and body, 407 and 418 ;
tail, 68 and 62 ; hind foot, 90 and 87 ; ear, 70 and 73. For
craneal measurements ses Table, p. 492.

Specimens examined. — Forty-six, from the following localities : —
Ireland : Belfast, 8 ; Clandeboye, Belfast, 1 ; Dumdrum, Tipperary, 1 ;
Kilmanock, Waterford, 2; no exact locality, 1 (skull).

* Based mainly on the domestic rabbit and on Gesner's account of the
rabbit (wild and tame) of Germany. That the wild rabbit of Germany was
considered by Linnaeus as the typical animal is indicated by his statement :
"Habitat in Europa australi."

ORYCTOLAGUS 491

Scotland. — Shetland Islands, 1 (skull).

England : Craigmoor, Isle of Man, 1 ; Lezayre, Isle of Man, 1 ; Kilnsea,
Yorkshire, 1 ; Leeds, Yorkshire, 1 (U.S.N.M.) ; Sandringham, Norfolk, 2 ;
Pangbourne, Berkshire, 2 ; Croydon, Surrey, 2 ; Orleston, Kent, 1 ; Kent, 1
(skull); Ditchling, Sussex, 4 ; Seatown, near Bridport, Sussex, 3 ; Ditchling,
Sussex, 4; Tidmarsh, Berkshire, 2 ; near London, 1 (U.S.N.M.); Orleston,
Kent, 1 ; Kent, no exact locality, 1 (skull) ; Barrow, Suffolk, 1 (U.S.N.M.) ;
no exact locality, 1 (U.S.N.M.).

Germany : Ingelheim, Rheiuhessen, 2 ; Strassburg, 2 ; south Germany, 3
(skulls).

Memarhs. — The wild rabl^it of Ireland, England and central
Europe appears to be very constant in size, allowance being made
for individuals that show evidence of crossing with domestic
stock.

6. Belfast, Down, Ireland. W. Thompson (p). 37. 7. 8. 25.

(5, 9. Belfast, Down. Marquis of Dufferin 5. 2. 22. 1^2.

and Ava (p).

?. Dumdrum, Tippcrary. Major P. J. Waldron 5. 1. 15. 1.

(c & p).

6, 9. Kilmanock, Waterford. G. Barrett-Hamilton 'J. 12. 15. 5.

(p). 11. 1. 2. 109.

skull. Ireland. C. Darwin (p). 68. 2. 19. 92.

skull. Shetland Islands, Scot- 0. Darwin (p). 68. 2. 19. 96.

land.

9. Lezayre, Isle of Man. C. H. B. Grant (c & p). 11. 1. 3. 448.

6, 9 St. Sandringham, Norfolk, H.M. King Edward 96. 5. 9. 1-2

England. VII. (p).

skull. Cambridgeshire. J. Baker (p). * 519. c.

{Type of Cuniculus fodicns Gray.)

6, 9. Pangbourne, Berkshire. Sir James Clark, Bt. 97. 2. 14. 1-2.

(c & p).

(5,9. Croydon, Surrey. C. H. B.Grant (c &p). 11.1.8.398-899.

i, 3 9. Ditchling, Sussex. Guy DoUman (c & p). 8. 9. 25. 1-4.

6. Orleston, Kent. P. N. Garrard (c & p). 90. 1. 5. 1.

skull. Kent. C. Darwin (p). 68. 2. 19. 98.

2 9. Ingelheim, Rheinhesscn, C. Hilgert (c). 8.6.15.10-11.

Germany.

3. S. Germany. Dr. A. Giinther (c). 59. 9. 6. 44-46.

Oryctolagus cuniculus huxlevi Haeckel.

1874. Lepus huxleyi Haeckel, Hist, de la creation des etres organises

d'apres les lois naturelles, p. 130 (Porto Santo).
1906. Oryctolagus cuniculus cnossius Bate, Proc. Zool. Soc, London, 1905,

II, p. 322, April 5, 1906 (Dhia, off Candia, Crete). Type in British

Museum.

1910. Oryctolagus cuniculus cnossius Trouessart, Faune Mamm. d'Europe,
p. 215.

Type locality. — Island of Porto Santo, Madeira.

Geographical distribution. — Mediterranean region; introduced
on the Azores, Madeira, and Salvage Islands, and probably
elsewhere.

Characters. — Like Lepus cuniculus cuniculus but smaller
(occipitonasal length of largest skulls, 71 to 77 mm. ; hind foot,
72 to 82 mm.) ; ear relatively longer (equal to that of the larger

492

RODENTIA

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494 KODENTIA

animal) ; colour more finely grizzled, the back showing less
tendency to be distinctly clouded with black.

3Ieasurements. — Adult from Porto Santo : hind foot, 72.
Two males from Terceira, Azores : hind foot, 73 and 76. Adult
male from Seville, Spain : hind foot, 75 ; two females from the
same locality : hind foot, 72 and 72. Adult male and female
f rom Coto Doiiana, Huelva, Spain : hind foot, 73 and 80. Adult
male and female from San Cristobal, Minorca, Balearic Islands :
head and body, 356 and 340 ; tail, 40 and 45 ; hind foot, 82 and
80 ; ear from crown, 70 and 70. Average and extremes of five
males from Poulx, Gard, France : head and body, 371 (347-397);
tail, 55 (45-63) ; hind foot, 77 • 8 (76-80 • 2). Adult female from
Island of Dhia, off Candia, Crete (type of cnossius) : head and
body, 341; tail 65; hind foot, 77 "6; ear, 70. For cranial
measurements see Table, p. 493.

Specimens examined. — Seventy, from the following localities : —

Porto Santo : Five.

AzoEES : Terceira, 2 ; San Miguel, 4.

Spain : Coto Donaua, Huelva, 4 ; Seville, 4 ; Silos, Burgos, 3 ; Castrillo
de la Reina, Burgos, 1 ; Selva, Majorca, Balearic Islands, 1 ; Inca, Majorca, 2 ;
Alcudia, Majorca, 1 ; San Cristobal, Minorca, 2.

Prance: Poulx, Gard, 5; near Nlmes, Gard, 14 (Mottaz) ; Digne,
Basses- Alpes, 8 (Mottaz).

Italy : Island of Capraja, Genoa, 5 (B.M. and Genoa) ; Palermo,
Sicily, 1 (U.S.N.M.).

Sardinia: Assuni, 7 (U.S.N. M.).

Crete : Dhia Island, off Candia, 1 (type of cnossius).

Eemarhs.- — In colour the Mediterranean rabbit nearly agrees
with the central European form, but when series of skins are
compai"ed it is at once seen that the smaller animal is the more
grey and finely grizzled of the two, its back seldom distinctly
clouded with black, a condition that is usual in true cunicnivs.
The actual elements of the colour are essentially the same in the
two, but in huxleiji Ihe sub-terminal annulations of the longer
hairs are slightly paler than in the central form. Except for
the differences in size the skull and teeth agree with those of the
larger animal.

The failure to distinguish between this race and that of
central Europe has been responsible for much speculation as to
the probable origin within historic times of the small i-abbits of
Porto Santo, which happened to be compared by Darwin with
British examples of true cuniculus. Some of Darwin's specimens
from this island are still in the British Museum. As might have
been anticipated they prove to be exactly similar to the common
Mediterranean form.

5. Porto Santo, Madeira C. Darwin (p). 68.2.19.93,104,

Islands. 107, 111, 117.

1. San Miguel, Azores. F. du Cane Godman 73. 3. 21. 1.

(c & p).

c5, 9. San Miguel. Major F. A. Chaves G. 3. 25. 1-2,

(c & p).

LEPUS 495

9. San Miguel. Major F. A. Chaves G. 3. 25. 3.

(c & p).

2. Goto Donana, Huolva, J. P. Gassiot (c & p). 72. 10. 26. 3-4.
Spain.

6, ?. Goto Dofiana, Huelva. A. Chapman (c & p). 2. 8. 3. 1.

2 6,29. Seville. {Dr. A. Buiz.) Lord Lilford (p). 95.3.3.15-18.

1. Silos, Burgos. S. & N. Gonzalez (c). 8. 7. 7. 26.

9. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 123.

9. Castrillo do la Reiua, G. S. Miller (c). 8. 8. 4. 122.

Burgos.

6. Selva, Majorca, Balearic 0. Thomas (p). 1. 3. G. 3.

Islands. [M. liiutorl.)

2 9. Inca, Majorca. 0. Thomas (p). 1. 3. G. 5 G.

{M. Riutort.)

6. Alcudia, Majorca. 0. Thomas (p). 1. 3. C. 4.

{AI. Riutort.)

6, 9. San Cristobal, Minorca. 0. Thomas and R. I. 0. 7. 1. GG-67.

Pocock (c & p).

3 6,19. Poulx, Gard, France. 0. Thomas (p). 8. 8. 10. 135-138.

(C. Mottaz.)
1. Capraja Island, Tuscany. Genoa Museum (e). 7. 2. 28. 4.
9. Island of Dhia, Crete. Miss D. Bate (c). 5. 12. 2. .35.

{Type of O. cnossms Bate.)

Genus LEPUS Linna3us.

1758. Lepus Linnaeus, Syst. Nat., i, 10th ed., p. 57 (type by tautonymy,
L. timidus Linnaeus).

1828. ? Lagos Brookes, Catal. Anat. and Zool. IMus. of Joshua Brookes,

p. 54 (Nomen nudum).

1829. Chionohatcs Kaup, Entw.-Geseh. u. Natiirl. Syst. d. Europ. Thier-

v^^elt, I, p. 170 {variabilis and borealis).
1857. Lepus Blasius, Stlugethiere Deutschlands, p. 410 (part).
18G7. Eulagos Gray, Ann. and Mag. Nat. Hist., 3rd ser., xx, p. 222,

September, 18G7 {mcditerrancus a-ndjudn'se).
1899. Lepus Major, Trans. Linn. Soc, London, 2nd ser., Zool., vii, p. 514,

November, 1899.
1899. Eulepus Acloque, Faune de France, Mammiferes, p. 52 {curopnncs

and variabilis).
1904. Lepus Lyon, Smithsonian Miscell. Coll., xlv, p. 389, June 15, 1904.

Type species. — Lepus timidus Linnseus.

Geographical distribution. — Europe, Asia, North America and
Africa.

Characters. — Skull with bony palate short, its length at
narrowest region never more than two and one-half times that
of first upper molar ; width of choante greater than least length
of palate, and about four times that of first molar ; sutures of
interparietal obliterated in adult ; postorbital processes broad
and triangular, with distinct anterior and posterior limbs ; first
upper premolar with deep median re-entrant angle, on each side
of which is a smaller re-entrant angle of varying depth ; anterior
portion of anterior lower premolar with a narrow re-entrant
angle on its front face and a broad re-entrant angle on external
aspect ; second to fifth upper cheek-teeth alike, the re-entrant

496 RODENTIA

angle extending from inner face about three-quarters of distance
across crown, the adjacent edges of the fold closely approximated
and finely crenulate ; posterior section of second, third and
fourth lower cheek-teeth about four-fifths as wide as anterior
half; last upper molar a small elliptic cylinder; third lower
molar with anterior segment elliptical in section, the posterior
segment smaller and nearly terete, the enamel of the two
cylinders normally separate throughout. Externally charac-
terized by the soft pelage, well developed tail, and long feet
heavily clothed with hair that nearly conceals the nails. Young
at birth active, the eyes open, and the body completely furred.

Bemarhs. — As now restricted the genus Lepus^ contains about
eighty named species. Eight of these occur in Europe, most of
them represented by several geographic forms.

KEY TO THE EUROPEAN FORMS OP LEPUS.

Tail, including pencil, much shorter than hind
foot, its upper surface white or clouded with
brown or grey, never with clear black median
area ; zygoma relatively deep anteriorly, the
distance from anterior termination of groove
on outer surface to front edge less than least
depth; root of upper incisor extending to
suture between premaxillary and maxillary ;
pelage usually changing to white or whitish
in winter (distribution northern and Alpine)
Varying Hares.
General colour of upper parts in summer dull

russet (approaching the wood-brown and

russet of Ridgway) ; winter pelage never

entirely white (Ireland) ;... L. hibcrniciis, p. 5,31.

General colour of upper parts in summer greyish

brown (approaching the broccoli-brown and

hair-brown of Ridgway) ; winter pelage

usually white or pale grey L. iimidiis, p. 522.

Occipitonasal length of adult skulls ranging
from 95 mm. to 103 mm. (Scandinavian

Peninsula) L. t. tinmlus, p. 526.

Occipitonasal length of adult skulls ranging
from 85 to 93 mm.

Ear from crown 80 to 90 mm. (Scotland) L. t. scoticus, p. 529.

Ear from crown 90 to 100 mm. (Alps) L.t. varronis, p. 528.

Tail, including pencil, about as long as hind foot,
its upper surface with conspicuous clear
black median area ; zygoma relatively shallow
anteriorly, the distance from anterior ter-
mination of groove on outer surface to front
edge usually equal to or greater than least
depth ; root of upper incisor not extending to
suture between premaxillary and maxillary ;
pelage never changing to white and rarely to
grey in winter (distribution central and
southern) Ordinary Hares.
Hind foot less than 105 mm. ; legs without white

markings, the inner surface scarcely lighter

than outer (Sardinia) L.meditcrraneics,T^.bl^.

LEPUs 497

Hind foot more than 110 mm. ; legs usually with
white markings, the inner surface always
noticeably lighter than outer.
Outer side of thigh much brighter than back,
its colour often nearly the cinnamon-
rufous of Ridgway ; dorsal surface of foot

and wrist marked with pure white L. granatcnsis, p. 515.

Ear from crown about 'J5 mm. (Basses-
Pyrenees) L. granate nsis ilu rissius,

p. 518.
Ear from crown, 105 to 115 mm.

Buffy tints pale, approaching the cream-
buff of liidgway (Central and south-
ern Spain) L. (jianalensia granatensis,

p. 516.
Buffy tints rich, approaching the ochra-
ceous-buff of Ridgway (North-western

Spain) L. i/ranatensis gallxciiis,

p. 517.
Outer side of thigh not brighter than back,
its colour ochraceous, buify, or greyish ;
dorsal surface of foot and wrist without
white markings.
Predominating brownish tints ; rump
usually bluish grey in strong contrast
with back.
Rump scarcely greyer than back ; black
area on outer side of ear extending
about 40 mm. below tip ; skull with
unusually slender rostrum (Parnassus

region, Greece) L. imrnassius, p. 519.

Rump noticeably greyer than back ; black

area on outer side of ear extending

20 to 30 mm. below tip ; rostrum

normal.

Size small, hind foot 119 to 12G, occipi-

tonasal length of skull 90 to 96

(Crete and Cephalonia) L. creticus, p. 512.

Size medium or large.

Rump very conspicuously grey ;
hind foot 135 to 155 mm. ; occi-
pitonasal length of skull 97 to
105 mm. (Roumania to the Pelo-

ponesus) L. eurojixus ti-anssylvanicits,

p. 509.
Rump less conspicuously grey ; hind
foot 125 to 140 mm. ; occipito-
nasal length of skull 95 to 103 mm.
(South-eastern France to Corfu) L. e. meridiei, p. 506.
Predominating buffy tints ; rump buff or
buffy grey, scarcely or not contrasted
with back.
Cheeks conspicuously whitish ; general
colour of upper parDs a light cream-
buff ; size very large, hind foot about
155 mm. (Eastern Germany and

eastward) L. e. hybridus, p. 508.

Cheeks greyish or brownish ; general
colour never a light cream-buff ; size
large or medium, hind foot 113 to
147 mm.

i08 RODENTIA

Hind foot 124 to 147 mm. ; occipifco-
nasal length of skull U3 to 102 mm.
(Central).
Buffy tints rather pale, approaching
the cream-buff of Ridgway (Ger-
many, Denmark, France, except

southern portion) L. e. europxus, p. 502.

Buffy tints rich, approaching the
ochraceous - buff of Ridgway

(England) L. e. uccidentalis, 50i.

Hind foot 113 to 125 mm. ; occipito-
nasal length of skull 8'J to 94 mm.
(Southern).
Nape patch well differentiated ; base
of ear conspicuously buffy be-
hind ; underfur not yellowish

(Pyrenees) L. c. 2^ji'eiiatcus, 506.

Nape patch scarcely indicated ; base
of ear not conspicuously buffy
behind ; underfur yellowish
(Corsica, Sicily, southern Italy) L. e. corsicamis, p. 507.

LEPUS EUROPXUS Pallas.
(Synonymy under subspecies.)

Geoyrcqjhicdl distrihution. — Central Europe from Great Britain
to Russia and from the Baltic south to the Pyrenees, Italy and
Greece.

Diagnosis. — Tail moderately long, its length including terminal
liairs about equal to that of hind foot, its upper surface never
white or grizzled, always with conspicuous black median area ;
ear long, extending decidedly beyond nostril when laid forward ;
general colour of upper parts a coarsely grizzled buffy or ochra-
ceous brown ; outer side of thigh not contrasting in colour with
back ; buffy and ochraceous tints occasionally replaced by pale
smoke-grey in a rarely assumed, special winter pelage ; ear with
conspicuous black terminal area on outer surface ; size rather
large, the hind foot usually more than 130 mm., though less in
some southern races ; root of upper incisor short, not extending
to suture between premaxillary and maxillary ; height of posterior
upper premolar measured from crown to upper surface of root
capsule usually less than alveolar length of tooth-row.

External characters. — Except for the relatively longer limbs,
ears and tail, the external form is essentially as in Oryctolagus.
Digits similarly 5-4, their proportional lengths as in the smaller
animal, but claws somewhat more flattened. Mammye : p 1 — 1,
a 2 - 2 = 6.

Colour. — Upper parts butiy, nearly clear on sides, underlaid
with blackish on back, the two colours forming a coarse mixture
varying considerably according to condition of pelage, but buffy
normally in excess. On sides the black becomes less, disappear-
ing entirely on legs, collar and along edge of white ventral area,
llurap usually lighter than either back or sides, sometimes

LEPus 499

conspicuously grey. Sides with evident though not very notice-
able sprinkling of hairs 60-70 mm. in length, the basal half dark,
the terminal half whitish. Tail white below and around entire
edge,* the dorsal surface with a broad clear black median stripe.
Nape clear bufFy, with or without a greyish white suffusion.
Head essentially like body but with the darker and lighter
colours more finely blended. An evident (sometimes conspicuous)
lighter area between eye and muzzle and another between eye
and base of ear. A slightly defined grey eyering, below and
behind which there is often a noticeable tawny area. An
ochraceous or tawny spot at base of whiskers. Ear buffy (rather
paler than buff of body) with the following colour-pattern : inner
surface cleai- buffy sometimes tinged with grey, the buffy clearer
and brighter just below tei'minal black rim, and much darker and
somewhat grizzled over an area about 40 mm. long and 10 mm.
wide along median portion of outer edge (though not involving
extreme rim, which is very pale bufl" *or even whitish from base
to black terminal portion), the grizzled region further emphasized
by a small pallid area at its inner base ; anterior outer surface a
fine grizzle of the same elements as those of back, the edge
conspicuously fringed with clear bufiy except at black-rimmed
tip ; posterior outer surface bufiy at extreme base, grey or
buffy grey at middle, and black at tip, the black area extending
conspicuously below rim of ear and forming a definitely outlined
j)atcla 25-35 mm. long, and 10-15 mm. wide. Underparts,
except collar, white, this colour extending to inner surface of
legs, which, though occasionally much tinged with buffy, are
always noticeably paler than outer surface (cf. L. mediterr emeus).
A clear buffy or ochraceous patch in inguinal region, the two
patches sometimes meeting in median line. Feet butty, darker
on the toes, paler proximally, but never marked with pure white
(cf. L. granatensis) ; soles an indefinite buffy grey.

The grey winter coat, when it occurs, differs from the usual
pelage merely in the substitution of a light drabby grey for all
the buffy and ochraceous tints, the black remaining unchanged.
The colour pattern is therefore in no way modified, though the
general appearance of the animal is so altered as to have given
rise to the belief that individuals in this pelage represented
hybrids between Lepus europse'm and some form of varying hare.|

Skull. — The skull of Le^ms europseus resembles that of L.
iimidus more closely than would be anticipated from the con-
spicuous external differences in the animals. Ordinarily they
may be distinguished by size, as in L. europseus the occipitonasal
length is usually more than 93 mm. (92 to 105 mm.), while in
L. timidus (except in the large L. t. timidus) it is seldom more

* The absence of the white edging at tip of tail is nearly always an
indicatioii that the extremity has been broken off.

t Such individuals may be distinguished from hybrids by the length
and colour of the tail, and by their cranial characters.

2 K 2

500 RODENTIA

than 92 mm. (8G to 93 mm.). Brain-case relatively narrower
and deeper than in Lepu.s tiiiiidiis, and parietal region less
flattened, the dorsal protile when viewed from behind arching
abruptly and conspicuously above level of zygomatic roots.
Anterior half of frontal nearly flat, the orbital rims rising slightly
but abruptly above general level of interorbita) region. Posterior
extremity of malar and relative length of suture between malar
and zygomatic process of squamosal as in L. timidus. Anterior
portion of zygoma usually though not always differing from that

Fig. 102a.
Lepus europsevit. Nat. size.

of L. timuliis in the greater width and relatively less depth of
the elevated region in front of lateral groove ; the least distance
from groove to anterior edge of zygoma usually equal to or greater
than depth at same region. Auditory bulla relatively a little
larger than in Lepus timidus, its ventral protile a little less
convex and anterior border less rounded off, a pecularity not
easily described but which when once seen is readily appreciated.
Teeth. — In general the teeth resemble those of Orydolagus
ciinieulus, diff'ej'ing in a few slight details only. Upper incisor
with groove on anteiior face decidedly nearer internal border

LEPUS 501

than external border ; posterior face with longitudinal concavity
barely indicated ; root as short as in Oryctolagus, bvit whole
course of tooth noticeably indicated on outer surface of bone.
Anterior upper premolar with lateral re-entrant angles on
anterior margin variable in development, one or both occasionally
obsolete. Other maxillary teeth as in Oryctolagus, but posterior
border of re-entrant fold sometimes crenulate ; roots of maxillary

Fig. 102b.
Lepus eitropcewt. Xat. size.

teeth not extending so far into orbit as in Oryctolagus and in
Lepus timldiis, the distance from crown of third premolar to
upper surface of its root capsule usually less than alveolar length
of tooth-row. Anterior lobe of anterior lower premolar usually
narrower than in Oryctolagus, but broader than in Lepus timidus ;
posterior section of this tooth as well as that of the succeeding
molariform teeth (including «<,) relatively larger than in
Oryctolagus.

502

RODENTIA

BemarJcs. — Lepus europseus is readily distinguishable among
the hares of continental Europe by its large size, long ear with
distinct colour pattern on dorsal surface, absence of white
markings on feet, and lack of noticeable contrast between colour
of sides of body and outer svirface of thigh. It is a plastic
species, readily becoming differentiated into local races. Seven
of these are now known to occur west of Russia.*

Lepus europ^us europ.eus Pallas.

1777. [Lepus] timidus Erxleben, Syst. Regni Anim., i, p. 325 (Not of

Linnaeus, 1758).

1778. Lepus curopa^us Pallas, Nov. Spec. Quaclr. Glir. Ord., p. 30 (Bur-

gundy, France).
1783. Lepus timidus Schreber, Saugtbiere, pis. ccxxxniA and ccxxxiiin

(Germany). Not of Liunseus, 1758.
1789. Ijcpus curopiKUs Scbreber, Saugtbiere, iv, p. 865.
1801. L^[_epiis\ t^imidus'] alhus Becbstein, Gemeinn. Naturgescb. Deutscb-

lands, I, 2nd ed., p. 1096 (Tbiiringen, Germany).
1801. L^cpus'] t[imidus'] flaviis Becbstein, Gemeinn. Naturgescb. Deutscb-

lands, I, 2nd ed., p. 1096 (Tbiiringen, Germany).
1801. L[epus'] t[;imidu.s'] nigcr Becbstein, Gemeinn. Naturgescb. Deutscb-

lands, I, 2nd ed., p. 1097 (Tbiiringen, Germany).
1820. Lepus viedius Nilsson, Skand. Fauna, i, p. 224 (Zealand, Denmark ;

based on specimens in grey winter pelage).
1857. Lepus timidus, b. IMitteleuropaiscbe Form, Blasius, Silugetbiere

Deutscblands, p. 417.
1859. ? Lepus campicola Gervais, Zool. et Paleont. Francaises, 2nd ed.,

p. 47 (Nomen nudum based on tbe common bare of France).
1867. ? [Lepus timidus'] coronatus, rufus, cinereus, nigricans, maculatus,

albus and niger 'Fitzinger, Sitzungsber. kais. Akad. Wissenscb.

Wien, Matb.-Naturwiss. Classe, lvi, p. 161 (Austria). Nomina

nuda.
1875. [Lepus timidus] var. argenteogrisea Konig-Wartbausen, Verzeicbniss

der Wirbeltbiere Oberscbwabens, Saugetbi/3re, ]). 85 (Ulm, Wiirtem-

berg, Germany). Probably based on an individual in grey winter

coat.
1910. Ljcpus europieus and L. medius Trouessart, Fauna Mamm. d'Europe,

p. 219.

Tij23e locality. — Burgundy, France. f

Geographical distribution. — Central Europe from Germany to

* Tbe Lepus europxus carpathorum of Hilzbeimer (Zool. Anzeiger, xxx,
p. 512, August 14, 1906) from tbe Carpatbians may represent an eigbtb
race. Tbe difficulty of clearly understanding tbe various local forms of
European bares is greatly increased by tbe frequency witb wbicb tbe
animals are transported from one region to anotber for tbe purpose of
restocking exbausted bunting grounds. Lj. e. tyansylvanicus bas tbus been
taken to Denmark, and L. e. occidenfalis to Switzerland. Otber instances
of tbe same kind will undoubtedl}' be found.

t Pallas refers to Daubenton's account of tbis animal for measurements
(Nov. Sp. Quadr. Glir. Ord., p. 2), and mentions no definite locality.
IPolonia and Pannonia are, be says (p. 5), inbabited by anotber form.
Daubenton's description (Buffon, Hist. Nat., vi, pp. 264-299) was based on
Burgundian specimens.

LEPUS 503

the Atlantic coast and from Denmark to central France. This
is probably the form recently introduced in southern Sweden.

Dia(jn(mii. — Bufty tints pale, approaching the cream-buff of
liidgway ; rump usually not grey, always with evident darker
area continuous with black tail-stripe.

Colour. — Underfur (l^O mm.) silvery white at base (occasionally
with a slight buffy tinge), becoming abruptly dark sepia slightly
beyond middle of hairs and blackish at extreme tips ; longer
hairs (30 mm.) whitish at base, dark brown at level of dark
portion of underfur, then with a cream-buff annulation 5-7 mm.
long and a shorter blackish tip. Clear area along sides cream-
buff, darkening to a light, yellowish ochraceous-butf in inguinal
region and on outer side of fore leg. Collar light ochraceous-
buff. Rump greyer than back, but usually so suffused with
cream-buff" as to form no marked contrast, its median area
always slightly darker than lateral portions.

Shull and, tectJi, — In the typical race the skull attains nearly
the maximum size for the species, the occipitonasal length
frequently exceeding 100 mm., but apart from this, neither skull
nor teeth show any special features worthy of note.

Measurements. — Two adult males and an adult female from
Haslev, Zealand, Denmark : head and body, 660, 680 and 640.;
tail, 80, 82 and 74; hind foot, 136, 147 and 138; ear from
crown (fresh) : 138, 131, and 125. Adult male from Ingelheim,
Rheinhessen, Germany: head and body, 571; tail, 98; hind
foot, 135 ; ear from crown (dry), 120. Two adult males from
Rheinthal, 8t. Gallon, Switzerland : head and body, 570 and 570 ;
tail, 100 and 85 ; hind foot, 136 and 137 ; ear from crown (dry),
117 and 115. Two adult females from Werdenberg, St Gallon,
Switzerland : head and body, 600 and 605 ; tail, 90 and 95 ;
hind foot, 145 and 144; ear from crown, 114 and 118 For
cranial measurements see Table, p. 510.

Specimens examined. — Twenty-seven, from the following localities : —

Denmark: Haslev, Zealand, 5; Zealand (no exact locality), 1.

Belgium: Slype, West Flanders, 1.

France : Near Paris, 1 ; Etupes, Doubs, 1 (Mottaz).

Germany: Brunswick, 3; Ingelheim, Rheinhessen, 1; Burg, near
Magdeburg, 1; south Germany, 6 (skulls).

Austria-Hungary: Salzburg, 1 (U.S.N.M.).

Switzerland : Flawil, St. Gallen, 1 (U.S.N.M.) ; Rheinthal, St. Gallen, 2
(U.S.N.M.); Werdenberg, St. Gallen, 2 (U.S.N.M.); Wittembach, St.
Gallen, 1 (U.S.N.M.).

RemarJcs. — The ty2:)ical form of Lepus europseus is characterized
by large though not maximum size, and rather light, strongly
yellowish colour. It is probably the most extensively distributed
of the western European forms. The pale winter coat is
occasionally assumed.

5. Haslev, Zealand, Den- O. Helms (c). 8. 2. 15. 1-5.

mark.
1. Zealand. Stockholm Museum (e). 40.0.2.71.

504

RODENTIA

1. Slype, Flanders, Belgium. Andrew van Iseghem 2.11,3.1.

?. Pans. A. Forsyth Major (p). 97. 2. 19. 1.

3. Brunswick, Germany. G. Barrett - Hamilton 8. 9. 29. 1-3.

6. Ingelheim, Rheinhessen. G. Barrett - Hamilton 11.1.2.110.

{Hilgert.) (p).

0. S. Germany. Dr. A. Giinther (c). 59. 9. G. 38-43.

Lepus europ^us occidentalis de Winton.

1898. Lcpus europseus occidentalis de Winton, Ann. and Mag. Nat. Hist.,
7th ser., i, p. 152, February, 1898. Type in British Museum.

1906. L{epus] e\_uropi£Us] occidentalis Hilzheimer, Zool. Anzeiger, xxx,
p. 512, August 14, 1906.

1910. Lepus europcTUs occidentalis Trouessart, Faune ]\Iamm. d'Europe
p. 220.

Tyj)e locality. — Herefordshire, England.

Geographical distrihution. — England, the Isle of Man and the
lower, more cultivated portions of Scotland, north to the Orkney
and Shetland Islands, the northern limits of the range much
extended artificially ; introduced in Ireland* and Switzerland.

Diagnosis. — Similar to Lepus europseus europseus, but buffy
tints rich and dark, approaching the ochraceous-bufF of Ridgway.

Colour. — Underfur as in L. europseus europseus, except that
the dark portion is more nearly black. Longer hairs with the
black tips better developed and the sub-terminal annulations a
dark ochraceous-bluff, showing in certain lights a decided tinge
of tawny. This tawny becomes clear and tinged with rufous
along sides and on inguinal patches, though it is duller and with
a decided clay-colour cast on collar. Rump as in L. europseus
europseus. Grey winter pelage : all the bufty tints replaced by
light grey, the exact shade of which is very uniform in the six
specimens examined. It is somewhat paler than the smoke-grey
of Ridgway and distinctly more blue, somewhat approaching the
grey No. 7, though not so dark. A very slight cream-bufF wash
may be detected on cheeks, sides of neck and of shoulders, and
on collar.

Skull and teeth. — The skull averages slightly smaller than in
L. europseus europseus, the occipitonasal length rarely attaining
100 mm. The teeth, however, are fully as large as in the
continental animal.

Measurements. — Two adults from the Isle of Man : head and
body, 538 and 548 ; tail, 67 and 89 ; hind foot, 131 and 134 ;
ear from crown, 105 and 110. Type (adult female) : head and
body, 570 ; tail, 86 ; hind foot, 135 ; ear from crown, 101. Two
adult females from Merton Hall, Norfolk : head and body, 575
and 583 ; tail, 80 and 90 ; hind foot, 141 and 135 ; ear from
crown, 120 and 98. Adult female (grey pelage) fi'om Pangbourne,

See Barrett-Hamilton, The Irish Naturalist, March, 1898, pp. G9-76.i

LEPUS

505

near Reading, Berkshu^e : tail, 75; hind foot, 135; ear from
crown, 99. For cranial measurements see Table, p. 510.

Specimens examined. — Thirty-six, from the following localities : —

Scotland : TuUoch, Inverness, 1 (grey pelage) ; Raith, Fifeshire, 1 ;
near Kilwinning, Ayrshire, 1 (grey pelage).

England : Isle of Man, 3 ; Bangor, Carnarvonshire, 2 ; Healey, North-
umberland, 1 ; Swithland Hall, Leicestershire, 1 ; Merton Hall, Norfolk, 3 ;
Barrow, Suffolk, 1 (U.S.N.M.) ; Sandringham, Norfolkshire, 2; Moor-
hampton, Hereford, 2 ; South Leigh, Oxfordshire, 1 (grey pelage) ; Hilling-
don, Middlesex, 1 ; Paugbourne, near Reading, Berkshire, 1 (grey pelage) ;
Marley Common, Haslemere, Surrey, 1 ; Weston Sands, Somerset, 2 ;
Stokenham Kingsbridge, Devonshire, 1 (U.S.N.M.) ; Gasford Castle,
Armagh, 2 ; south-east coast, Ireland, 1.

Ireland: Gasford Castle, Armagh, 2; south-east coast, 1.

Switzerland : Near St. Gallen, 4 ; Utzwil, St. Gallen, 1 (U.S.N.M.).

Remarks. — Though closely I'elated to Lcpna eurojjseus europixus,
this form is readily distinguishable by its darker, browner, general
colour. The pale winter coat is rarely observed.

1. Dingwall, Inverness-shire, Sydney Dennis (c &p). 8.11.14.1.
Scotland.
Raith, Fifeshire. W. R. Ogilvie-Graut 93. 12. 23. 2.

(c&p).
Kilwinning, Ayrshire. Hon. G. J\Iontgomerie 92. 2. 15. 1.

(c * p).
Isle of Man. P. M. C. Kermode 95. 1. 19. 1.

(c & p).
Bangor, Carnarvonshire, G. W. D. Assheton 97. 3. 8. 1.
Wales. Smith (p). 3. 1. 28. 1.

Healey, Northumberland, Rev. H. H. Slater 0.2.24.1.

England. (c & p).

Swithland, Leicestershire. Earl of Lanesborough 92. 1. 29. 1.
(c & p).
6, 2 ?. Thetford, Norfolkshire. Lord Walsingham (p). 98. 2. 11. 1-3.
2 St. Sandringham, Norfolk- H.M. King Edward 9G. 5. 9. 3-4.
shire. VII. (p).

Moorhampton, Hereford- W. E. de Winton 98.2.17.1.

shire. (c & p). (Type of subspecies.)

Moorhampton, Hereford- W. E. de Winton 3. 1. 26. 1.

shire. (c & p).

South Leigh, Oxfordshire. A. J. Butler (c & p). 93. 9. 7. 1.
Pangbourne, Berkshire. Capt. Saw bridge 0.11.16.1.

(c & p).
Haslemere, Surrey. S. J. Wigley (c & p). 6. 9. 18. 1.

Weston Sands, Somerset- Tomes Collection. 7. 1. 1. 175-176.

shire.
England. Purchased (Lead- 88. b.

beater).
Gasford Castle, Armagh, Hon. L. Powys (c&p). 76. 1. 25. 1-2.

Ireland.
South-east coast, Ireland. Miss E. Hope. 6. 4. 12. 1.

<5,2 ?. St. Gallen, Switzerland. O. Thomas (p). 2. 8. 4. 53-54.

(E. H. Zollikofer.) 4. 4. 5. 46-47.

6.
6.
6.
2.
1.
6.

506

RODENTIA

Lepus europ^us pvrenaicus Hilzheimer.

1906. L\_epus] cliiropams] pyrcnaictts Hilzheimer, Zool. Anzeiger, xxx,

p. 512, August 14, 1906.
1910. Lepus citropaius pyrcnaictis TrOuessart, Fauna Mamm. d'Europe,

p. 220.

Type locality. — -Bagneres [de Luchon 1], Pyrenees, France.

Gaograpliical disfrilmtion. — Pyrenean region, south-eastern
France.

Diagnosis. — Essentially like Lepus europeeus europseiis, but
smaller (hind foot, 113 to 125).

Specwiens examined. — Five, from the Department-'Of Ariege, France,
and the neighbouring portion of Andorra.

Remarhs. — The five specimens, though imperfect, do not
appear to be referable to any of the other forms of Lepus
europseus. They agree sufficiently well with pyrenaicas as
described by Hilzheimer.

2 9,1. Ax -les-Thermes, Ariege, V. Builles (c & i>). 10.9.20.1-3.
France.

Lepus europ^us meridiei Hilzheimer.

1859. ? Lepus mcridionalis Gervais, Zool. et Paleont. Francaises, 2nd ed.,
p. 47 (Nomen nudum, based on the hare of Languedoc and
Provence).

1906. Llepits] el^uropxus'] meridiei Hilzheimer, Zool. Anzeiger, xxx, p. 512,
August 14, 1904 (Aveyron, France).

1910. Lepus europ/EUS meridiei Trouessart, Faune Mamm. d'Europe, p. 220.

Type locality. — Department of Aveyron, France.

Geographical clistrihution. — South-central and south-eastern
France, northern Italy, Corfu.

Diagnosis. — Size as in Lepus europseus europseus ; colour
browner and less buffy than in the typical form, and rump
distinctly bluish grey, though less conspicuously contrasted with
back' than in the large L. e. transsylvanicus.

Measurements. — Four males from the neighbourhood of
Nimes, Gard, France, average and exti'emes : hind foot, 146*6
(140-152). Two males from Barcelonnette, Basses-Alpes,
France: head and body, 528 and 540; tail, 112 and 95; hind
foot, 138 and 140. Two males from Siena, Italy : head and
body, 500 and 550; tail, 80 and 90; hind foot, 130 and 133;
ear from crown, 116 and 118. Two females from the same
locality : head and body, 550 and 558 ; tail, 88 and 92 ; hind
foot, 130 and 134 ; ear from crown, 119 and 123. For cranial
measurements see Table, p. 511.

Specimens examined. — Twenty, from the following localities : —
Prance : Near Nimes, Gard, 7 (U.S.N.M. and Mottaz) ; near Digne,
Basses-Alpes, 3 (Mottaz) ; St. Paul, near Barcelonnette, Basses-Alpes, 2.

LEPUS 50»

Italy: Ceresole cl'Alba, Turin, 1 (Turiu) ; Poiiezza, Como, 1 (Ghidini) ;
Siena, 4.

Greece : Corfu, 2.

BemarJcs. — It is with some hesitation that I have referred
the hares of northern Italy and of Corfu to this form. In the
absence of more complete material it seems, however, the most
satisfactory course to pursue.

2 6. Barcelonnette, Basses-Alpes, O. Thomas (p) . 8. 8. 10. 133-134.

France. (C. Mottaz.)

2 i. Siena, Italy. {S. Biogi.) Dr. E. Hamilton. 98. 10. 2. 18-19.

6. Corfu, Greece. (C.J/oH«,J.) J.I. S. Whifcaker (p). 8.10.1.51.

6. Corfu. C. Mottaz (c). 8. 11. 3. 23.

Lepus europ^us corsicanus de Winton.

1898. Leims corsicanus de Winton, Ann. and Mag. Nat. Hist., 7th ser., t,
p. 155, February, 1898. Type in British Museum.

1906. L[eptis'] m[eclitcrraneus'] corsicanus Hilzheimer, Zool. Anzeiger, xxx,
p. 513, August 14, 1906.

1910. Lepus corsicanus Trouessart, Faune Mamm. d'Burope, p. 224.

Ty^^e localiti/. — Bastia, Corsica.

Geographical distribution. — Corsica, Sicily and Italy (Rome).

Diagnosis. — Size small, as in Lepus exiropeeus pijrenaicus (hind
foot, 114 to 125 mm., occipitonasal length of skull about 90 to
94 mm.) ; general colour more yellow than in true europxus, the
underfur conspicuously buff.

Colour. — The colour does not differ appreciably from that of
Lepus europseus europjseus, except that it is throughout somewhat
brighter and more yellowish, and the colour pattern of the ear
and the tawny markings below eye and at base of whiskers are
better defined ; nape patch scarcely indicated and without
special colour. Underfur with three distinct colour bands ; the
basal grey and terminal blackish as in L. europseus, but with an
intermediate band about 6 mm. in width of dull buff, thrown
into strong contrast by the black. Inguinal patches hght
tawny. Eump dull ochraceous-buff, the bases of the hairs
bluish grey (about grey ISTo. 7 of Ridgway).

Shdl and teeth. — Except for their reduced size the skull and
teeth resemble those of Lepus europseus europseus.

Measurements. — Type (adult male) : hind foot, 119 ; ear from
crown, 105. Two males from Marsala, Sicily: hind foot, 114
and 120; ear from crown, 103 and 110 mm. Adult male from
Rome, Italy : hind foot, 125 ; ear from crown, 127. For cranial
measurements see Table, p. 511.

Specimens examined. — Four, from the following localities : —
Corsica.: Bastia, 1 (type).
Sicily : Marsala, 2.
Italy : Rome, 1.

508 KODENTIA

BemarJcs. — Tlie conspicuously yellowish underfur of this hare
is perhaps its most striking characteristic. With the material
at hand I am unable to distinguish between the Corsican form
and that of Sicily and Rome.

1. Bastia, Corsica. Lord Lilford (r). 78. 7. 3. 4.

[Type of subspecies.)
2 6. Marsala, Sicily. J. I. S. Whitaker (p). 98. 2. 9. 1-2.

S. Rome, Italy. (C. Coli.) G. Barrett-Hamilton (p). 8. 9. 30. 1.

Lepus europteus hybridus Desmarest.

1822. Lcpus hybridus Desmarest, Mammalogie, p. 349 (Central Russia).
1842. Lcpus aquilonins Blasius, Amtl. Bericht xix Vers. Naturf. u. Aerzte,

Braunschweig, p. 89 (Central Russia).
1850. Lepus mcdius Middendorff, Bull, de la Classe T?hys.-Math. de I'Acad.

Imp. des Sci. Nat. de Saint-P6tersbourg, ix, p. 218 (part).
1857. [Le^ms fhnidus] c. Nordostliche Form, Blasius, Siiugethiere Deutsch-

lands, p. 417 (part).
1898. Lcpus curopnus (typical) de Winton, Anu. and Mag. Nat. Hist.,

7th ser., i, p. 150, February, 1898.
1906. L[epus'] vi\_cdius'\ acpuilonius Hilzheimer, Zool. Anzeiger, xxx, p. 511,

August 14, 1906.
1910. Lcpus medius aquilonius Troucssart, Fauue Mamm. d'Europe, p. 217.

l^ype hcality. — Central Russia. Based on the Russak of
Pallas (Nov. Sp. Quadr. Glir. Ord., p. 5).

Geographical distribution. — Central Russia, westward into
eastern Germany.*

Diagnosis. — Size greater than in Lepus ewopseus europssus
(hind foot about 15 mm.); colour in summer pelage essentially
as in the typical form but lighter, the cheeks strongly suffused
with dull white, the rump buffy grey with slightly darker
median area ; grey winter pelage habitually assumed.

Colour. — Though the actual elements of the colour of the body
are essentially as in Lepus europseus europveus, the general effect
is noticeably lighter, owing to the greater length of the sub-
terminal cream-buff annulations. The rump is also more
evidently grey in contrast with the back. White of interramial
region spreading conspicuously so that cheeks are practically
whitish throughout (except for a dull ochraceous-buff area under
and behind eye and another at base of whiskers), in strong
contrast with the grizzled buffy of crown, face and muzzle.
Grey area on back of ear nearly white and forming a very
conspicuous whitish rim to outer border. Inguinal patches
clear, light ochraceous-buff. Collar somewhat duller and paler.
Otherwise as in L. europstus europseus.

* While I have seen no German specimens of this hare there can be
little doubt that it is the East Prussian animal referred to by Altum
(Porstzoologie, 2nd ed., i, p. 181) as being not infrequently greyish white
in winter. The same author gives rather extended observations on the
occurrence of the grey winter coat in the hares of other parts of Germany.

LEPUS 509

Measurements. — Adult froui Litluuuiia : hind foot, 155; ear
from crown, 125.

Specimens examined. — One, from Lithuania.

Hemarhs. — This race appears to be well characterized by its
large size, rather light colour of summer pelage, and the
frequency with which the grey winter coat is assumed.

1. Lithuania. {T. Barcy.) Branicki Museum (b). 9i. 8. 7. 30.

Lepus europ^us transsylvanicus Matschie.

1901. Lepus transsylvanicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 236 (Taslau, Roumania).
1906, L{epus'] e\iiropiKUs\ transsylvaticits Hilzheimer, Zool. Anzeigcr xxx,

p. 512, August 14, 1906.
1910. Lepii.s curopn'us transsylvaticus Trouessart, Faunc Mamm. cl'Europe,

p. 221.

2'//j^e locality. — Taslau, Roumania.

Geogi-aphical distribution. — From Roumania southward through
the Balkan Peninsula to the Peloponesus. Exact limits of range
unknown.

Diagnosis. — Size nearly as great as in Lepus europseus
hyhridus (hind foot about 150 mm., occipitonasal length of skull
in largest individuals 105 mm.) ; general colour dark and
brownish, much as in L. eurojjisus meridiei, the rump bluish
grey in strong contrast with back ; black area on posterior
surface of ear normal in extent.

Measurements. — Adult male from Visoko, Bosnia, and from
Herzegovina : hind foot, 150 and 148 mm. ; ear from crown, 120
and 121 mm. For cranial measurements see Table, p. 511.

Specimens examined. — Eight, from the following localities : —

Bosnia : Visoko, 1.

Herzegovina: Mostar, 3 ; Dragajica, 1.

Albania : Coast opposite Corfu, 2.

Greece : Patras, 1.

BemarJcs. — While I have not seen Roumanian specimens of
this hare, the Balkan skins agree so essentially with the original
description that there seems little doubt that they represent the
same form. Though readily distinguishable from Lepus europseus
europaeiis its resemblance to L. e. meridiei, particularly to north
Italian skins of this form, is rather close. The Balkan-
Roumanian animal appears, however, to be sufficiently dis-
tinguished by its large size.

S. Visoko, Bosnia. Dr. 0. Reiser (c & p). 7. 10. 15. 3.

i, 9. Surmancipolje, Horze- Dr. 0. Reiser (c & r). 7. 10. 15. 1-2.

govina.
2 <5. Opposite Corfu, Albania. J. I. S. Whitaker (p). 8. 10. 1. 49-50.

(C. Mottas.)

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512

RODENTIA

LEPUS CRETICUS Barrett-Hauiiltoii.

1903. Lcpiia crdlctis Barrett-Hamilton, Aun. and Mag. Nat. Hist., 7th ser.,
XI, p. 126, January, 1903. Type in British Museum.

1905. Lcpus curopn'us crcHcus Bate, Proc. Zool. Soc, London, 1905, ii,
p. 322, April 5, 1906.

1910. Lepus creticus Trouessart, Faune Mamm. d'Europe, p. 222.

Type localiti/. — Crete and Cephalonia.

Geographical, distribution. — Crete.

Diagnosis. — General colouration light brownish essentially as
in Lepus curopseus transsylvanicus and L. p>arnassiiis, but paler,
the rump conspicuously grey ; size rather small, about equal to
that of L. europseus corsicanus (hind foot, 117-127 mm.; occipito-
nasal length of largest skulls about 95 mm.) ; cheeks grey,
noticeably grizzled ; ear scantily haired, very slightly buff-tinged.

Colour. — Upper parts a mixture of blackish and a very pale,
dull cream-buff, the general effect a coarsely grizzled brown,
becoming conspicuously grey on rump, but scarcely paler on
sides of body than on back. Underfur silvery grey at base.
Rump crossed by evident dark median area continuous with
black of tail. Nape patch wood-brown tinged with russet in
rather noticeable contrast with surrounding parts. Collar
varying from wood-brown to dull ochraceous-buff, usually grizzled
with buffy white, the former predominating. Head a fine grizzle
of same elements as body colour, the pale markings before and
behind eye silvery whitish though not very extensive. Cheeks
essentially like crown, distinctly grizzled and Avith an evident
greyish cast, the rusty markings obsolete. Muzzle and lips dull
cream-buff. Anterior outer surface of ear sepia finely and
inconspicuously grizzled with cream-buff and fringed along
border with bluish or buffy white ; posterior outer surface a
light bluish grey (about grey No. 10 of Ridgway) throughout,
except for the usual blackish (seal-brown) terminal area ; inner
surface sparsely sprinkled with fine bluish grey hairs except on
the brownish area along posterior margin ; a faintly indicated
buffy sub-terminal area ; posterior margin whitish except near
tip. Feet a dull buffy brown. Underparts (except collar) and
* inner surface of legs white, slightly tinged with pale buff
Inguinal patches dull ochraceous-buff.

ShuU and teeth. — The skull is of medium size, the occipito-
nasal length in old individuals about 96 mm. In foi'm it is
scarcely distinguishable from that of L. europjseus. Anterior
portion of zygoma essentially as in L. europseus, but showing
a decided tendency to become deeper. The teeth show no
peculiarities.

Measurements. — Average and extremes of three specimens
from Crete : hind foot, 124*3 (123-126); ear from crown, 116
(115-117). Average and extremes of six adults from Cephalonia :

LEPUS 513

head and body, 490 (483-495) ; tail, 92 (80-100) ; hind foot,
122-5 (117-127). For cranial measurements see Table, p. 520.

Specimens examined. — Twelve, from the following localities : —
Crete, 5.
Cephalonia, 7.

Remarks. — While resembling the brown races of Lepus
europseus the Cretan hare is readily distinguishable by its smaller
size and relatively somewhat larger ears.

2. Crete. H. O. Jones, R.N. (c). 99. 2. 14. 1-2.

1. Crete. H. 0. Jones, R.N. (p). 2. 11. 9. 1.

{Type of species.)

i. Kanea, Crete. Miss D. Bate (c). 5. 12. 2. 34.

i, 4 9. Mt. Vuno, Cephalonia, J. I. S. Whitaker (p). 8. 10. 1. 52-56.
Greece. (C. Mottaz.)

6,9. Mt. Vuno, Cephalonia. C. Mottaz (c). 8.11.3.20-21.

LEPUS MEDITERRANEUS Wagner.

1841. Lepus 7nediterraneus Wagner, Miinch. Gelehrt. Anzeiger, p. 439,
March 17, 1841.

1857. ILepus timidus'] a. Siideuropaische Form, Blasius, Sjiugethiere
Deutschlands, p. 417 (part).

1898. Lepus mcditerranetis de Winton, Ann. and Mag. Nat. Hist., 7th
ser., I, p. 154, February, 1898.

1906. L[epus'] m[_editerraneus'\ tijpicus Hilzheimer, Zool. Anzeiger, xxx,
p. 512, August, 1906.

1910. Lepus viediterraneus Trouessart, Fanne Mamm. d'Europe, p. 224.

Typ)e locality. — Sardinia.

Geographical distribution. — Sardinia.

Diagnosis. — Smallest of the European hares (hind foot, 93 to
103 ; occipitonasal length of skull, 81 • 4 to 87 mm.) ; colour much
as in Lepus granatensis, but white of underparts encroached on by
ochraceous-buff of sides, and not extending on legs, the inner
and outer surfaces of which are not evidently contrasted.

Colour. — Back essentially as in Lepus granatensis granatensis,
but elements more intimately blended, and ground colour a more
yellowish cream-buff, this slightly in excess of black. Underfur
with dull buff sub-terminal area (5 mm.) and black tips (4 mm.)
well defined, the bases of the hairs bluish grey (about the pearl-
grey of Ridgway). Sides with wide and noticeable clear buffy area
(the exact shade darker and more yellow than the ochraceous-buft
of Ridgway), strongly incroaching on white of underparts, and
continuous with inguinal patches and inner svirface of legs, on
fore legs becoming slightly paler, and on hind legs more yellow.
On outer surface of legs and dorsal surface of feet this colour
darkens so as to form a conspicuous contrast with cream-buff of
back, though there is no evident line of demarcation between the
two surfaces of the legs. Rump slightly more buflfy than back.

2 L

514 RODENTIA

Tail with the usual black and white areas. Nape patch rather
well defined, concolor with outer surface of legs. Head essen-
tially like back but more finely grizzled ; cheeks like crown or
somewhat sufi'used with ochraceous-buff, the pale eye-ring and the
spot before and behind eye obsolete ; a clear dark ochraceous-buff
patch at base of whiskers ; muzzle dull buffy or greyish, forming
no distinct contrast with surrounding jDarts. Ear dark and dull,
the colour pattern ill-defined, and the narrow whitish rim of
outer border thrown into unusually sti'ong contrast ; anterior
outer surface and dark area on inner surface prout-brown slightly
grizzled with dull buffy ; grey of posterior outer surface extending
nearly to base of ear ; black area normal ; no grey on inner
surface, this region, except on dark area, everywhere ochraceous-
buff like fringe of inner border. Collar dull ochraceous-buff
irregularly washed and varied with cream-buff. Interramial
region very dull white or pale ochraceous-buff. White of under-
parts limited to a narrow area, not more than 50 mm. across,
extending from posterior portion of chest to inguinal patches,
which may or may not completely separate it from the white of
intercrural region and base of tail. Anterior portion of chest
w^hitish ochraceous-buff like inner surface of fore legs.

Shull and teeth. — The skull of Lepus mediterraneus is imme-
diately distinguishable among European hares by its extremely
small size, the occipitonasal length in a fully adult male being
only 83 • 6 mm., thus scarcely exceeding that of Orcytolagus
cuniculus. In form and in the proportions of the different parts
it shows, however, no marked peculiarities, agreeing, in most
respects, with the large Lepus europseus, though the auditory bullse
more nearly approach the form occurring in L. granatensis.
Anterior portion of zygoma remarkably variable in form. Among
the four skulls examined two have this region essentially as in
L. europseus, one more nearly approaches the timidus type, while
in the fourth, although fully adult, the groove is so shallow and
ill-defined as to be practically absent, a condition that I have not
seen in other European hares. Teeth peculiar in their small size
only.

Measurements. — Adult male and female : hind foot, 103 and 96 ;
ear from crown, 114 and 109. Two other specimens, sex not
noted : hind foot, 93 and 95 ; ear from crown, 97 and 99. For
cranial measurements see Table, p. 520.

Specimens examined. — Six, all from Sardinia.

Beinarks. — This very distinct species is at once recognizable
among European hares by its small size and dull colour.

2. Sassari, Sardinia. Genoa Museum (e). 97. 2. 27. 1-2.

6,9. Cagliari, Sardinia. (Ma22a.) Genoa Museum (e). 98.5.12.1-2.

515

LEPUS GRANATENSIS Rosenhauer.

(Synonymy under subspecies.)

Geographical distribution. — Iberian Peninsula and Balearic
Islands. Exact northern limits of range not known.

Diagnosis. — Size rather small, about as in Lepiis corsicamis,
and L. creticus (hind foot, 115-120; occipitonasal length of
skull about 80-90 mm.) ; colour differing from that of all the
other European hares, except the much smaller L. merliterraneus,
in the bright, reddish colour of the outer surface of the thighs,
which forms the marked contrast with buffy of back ; inner
surface of legs normally white, the line of demarcation very
sharply defined ; doi'sal surface of feet and wrists marked with
pure white ; lengthened whitish hairs on sides A'ery conspicuous.
Mammse as in L. europseiis.

Colour. — Upper parts bufFy underlaid on back with blackish,
the two colours forming a coarse mixture, varying considerably
according to condition of pelage, but the bufty always in excess.
On sides the black gradually disappears, leaving a narrow, nearly
clear bufFy line bordering white of underparts. Rump usually
concolor with back, but sometimes tinged with grey. Sides
with a very conspicuous sprinkling of hairs about 50 mm. in
length, their basal portion blackish, the terminal half white.
Tail as in Lepus europseus. Nape patch usually small and ill-
defined. Head essentially like body but with the darker and
lighter colours finely blended. Cheeks and crown alike ; eye-ring
about as in L. europseus, but pale area before and behind eye
barely indicated, and no distinct rusty suffusion below eye ;
muzzle clear dull buffy ; a darker area at base of whiskers. Ear
as in Lepus europseus, but colour pattern more strongly defined
and black tei'minal area usually, though not always, more
extensive ; outer border conspicuously rimmed with whitish.
Outer surface of thigh conspicuously brighter than body, its
colour approaching cinnamon-rufous and very sharply contrasted
with the pure white of inner surface of leg. Outer surface of
fore leg not so bright as that of thigh but forming a strong
contrast with sides of body. Inguinal patch dull ochraceous-
rufous. Collar a dark, scarcely grizzled buffy. Rest of under-
parts a peculiarly snowy white, the white area unusually wide and
well defined, extending down hind leg and covering dorsal surface
of foot to base of claws, and usually though not always extending
similarly on front leg to wrist where a white spot is invariably
present. Soles an indefinite buffy brown.

Skull and teeth. — The skull of Lepus granatensis is rather
small, the occipitonasal length seldom exceeding 90 mm. In
form it differs from that of the other European hares in the less
deflection of the brain-case, and consequently less strong convexity
of the dorsal profile, a character not easily defined but readily

2 L 2

516 RODENTIA

appreciable on inspection, and evidently of much importance in
a group showing such remarkable uniformity in cranial characters.
The auditory bulla? are relatively larger than in the other species,
so that, when skull is viewed from below, the area of each bulla
appears to be about equal to that of flat portion of basioccipital
in front of condyles. Anterior portion of zygoma strictly as in
L. europseus. Teeth rather small but not showing any special
peculiarities of structure.

BeniarJcs. — The Iberian hare is at once distinguishable from
the other European members of the genus by the unusually
bright colour of the outer surface of the thighs and the presence
of white markings on the feet. The form of the skull is also
characteristic. Three local races are at present known.

Lepus granatensis GRANATENSis Rosenhaucr.

1856. Lc2)us granatensis Eosenhauer, Die Thiere Andalusiens, p. 3
(Granada).

1867. [Lcpus] MsiMnicus Fitziuger, Sitzungsber. kais. Akad. Wissensch.
Wien. Math.-Naturwiss. Classe, lvi, p. 161 (Substitute for grana-
tensis. " Natt." cited as authority).

1897. Lepus meridionalis Graells, Mem. Eeal Acad. Sci., Madrid, xvii,

p. 525 (Vicinity of Madrid).

1898. Lepus lilfordi de Winton, Ann. and Mag. Nat. Hist., 7th ser., i,

p. 153, February, 1898 (Seville). Type in British Museum.

1906. L[ep2is] cluro^jxtis] granatensis Hilzheimer, Zool; Anzeiger, xxx,

p. 512, August 14, 1906.

1907. Lepus granatensis granatensis Miller, Ann. and Mag. Nat. Hist.,

7th ser., xx, pp. 399, November, 1907.
1910. Lepus granatensis Trouessart, Faune Mamm. d'Europe, p. 222.

Type locality. — Granada, Spain,

Geographical distribution. — The greater portion of Spain,
extending at least from the Province of Burgos to the south and
east coasts ; Balearic Islands.

Diagnosis. — Ear long, its height from crown in dried
specimens 105-115 mm. ; general aspect pallid, the ground colour
of back nearly the cream-colour of Ridgway and much in excess
of black.

Colour. — Underfur (17 mm.) bluish grey at base, the hairs
becoming buffy terminally, the extreme tips slightly darker but
not definitely black ; longer ^airs (30 mm.) greyish at base, then
black to tip, each with a conspicuous, sharply defined sub-terminal
annulation (4 mm.) of pale cream-buff". Clear area along sides
(ill-defined) between cinnamon-rufous and ochraceous-rufous,
becoming somewhat more dull on inguinal patches and darker
and brighter on outer surface of thighs. Outer side of front leg
a duller shade of the same colour ; inner surface white to wrist,
where the white crosses to upper surface and forms a conspicuous
patch on metacarpals. Collar pale wood-brown strongly washed
with pale dull ochraceous-buff". Chin somewhat dusky.

LEPUS 517

Skull and teeth. — In the typical form the skull is of maximum
size for the species, occipitonasal length of full-grown individuals
ranging from 86 "4 to 90*4 mm.

Measurements. — Adult from Granada: hind foot, 115; ear
from crown, 120. Two males from Seville (the second the type
of lUfordi) : hind foot, 114 and 116 ; ear from crown, 118 and 120.
Two females from the same locality : hind foot. 111 and 114 ; ear
from crown, 110 and 113. Adult male and female from Silos,
Burgos : head and body, 500 and 520 ; tail, 95 and 87 ; hind
foot, 117 and 118 ; ear from crown, 110 and 112. For cranial
measurements see Table, p. 521.

Specimens examined. — Fifteen, from the following localities in Spain :
Silos, Burgos, 2 ; Castrillo de la Reina, Burgos, 2 ; Seville, 4 ; Las Marismas,
near Seville, 1 ; Granada, 1 ; Jerez, Cadiz, 1 ; Goto Dofiana, Huelva, 2 ;
no exact locality, 1 ; Selva, Majorca, Balearic Islands, 1.

2 9, Castrillo, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 120-121.

2. Coto Donana, Huelva. A. Chapman (c & p). 8. 3. 8. 8-9.

2 (5, 2 9. Seville. (Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 11-14.

(95. 3. 3. 12. Type of L. lilfordi de W.)

1. Las Marismas. near H. F. Witherby 0. 2. 17. 1.

Seville. (c & p).

1. Granada. Col. J. H. Irby (c & p). 96. 7. 29. 1.

juv. Jerez, Cadiz. A. Chapman (c & p). 8. 3. 8. 16.

1. Spain. Lord Lilford (p). 75. 2. 11. 3.

1. Selva, Majorca, Balearic 0. Thomas (p). 1. 3. 6. 2.
Islands. (M. Biutort.)

Lepus granatensis gall^cius Miller.

1907. Lepus granatensis gallfecius Miller, Ann. and Mag. Nat. Hist.,
7th ser., xx, p. 400, November, 1907. Type in British Museum.

1910. Lepus granatensis gallaecius Trouessart, Faune Mamm. d'Fjurope,
p. 223.

Type locality. — ^La Coruna, Province of Coruila, Spain.

Geographical distribution. — At present known from extreme
north-west Spain only.

Diagnosis. — Similar to Lepus granatensis (jranatensis, but colour
throughout dark and rich, the ground colour of back nearly the
ochraceous-bufF of Ridgway, and scarcely in excess of black ;
white markings on front leg not so extensive as in the typical
form.

Colour. — ^Underfur with buff distinctly brighter than in
Lepus granatensis granatensis, and extreme tips of the hairs
becoming definitely black. Sub-terminal annulations of longer
hairs light ochraceous-buff. Clear area along sides dull cinnamon-
rufous, the inguinal patches similar, but outer surface of thigh
and of front leg distinctly darker and brighter. Inner surface
of front leg with white extending to wrist or not beyond elbow,
the usual white markings at wrist present in either case, though

518

RODENTIA

slightly less extensive than in L. (jranatensis granatensis. Collar
a dai"k wood-brown washed with yellowish clay-colour. Head and
ears noticeably darker than in the typical form, and eye-ring
thrown into rather strong relief. A conspicuous dull cinnamon-
rufous patch at base of whiskers. Chin conspicuously dusky in
strong contrast with surrounding parts.

Measurements. — Type : hind foot, 107 ; ear from crown, 105.
Adult from Vigo: hind foot, 110; ear from crown, 105. For
ci'anial measurements see Table, p. 521.

Specimens exmnined. — Two, one from La Corufia, the other from Vigo.

1. Vigo, Pontevedra, Spain. Lord Lilford (p). 82. 12. 8. 1.

6. La Corufia, Coruiia, Spain. Dr. F. L. Seoane (c & p). 94. 2. 16. 1.

(Type of subspecies.)

Lepus granatensis iturissius Miller.

1907. Lepus granatensis iturissius Miller, Ann. and Mag. Nat. Hist.,
7th ser., xx, p. 401, November, 1907. Type in British Museum.

1910. Lepus granatensis iturissius Trouessart, Faune Mamm. d'Europe,
p. 223.

Ty2)e locality. — Basses-Pyrenees, near Biarritz, France. The
type was probably bought in the Biarritz market, but a note
published by the collector in the Field, Lxxxix, January 30,
1897, p. 135, refers to the animal as occurring on the Spanish
side of the border.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — Colour essentially as in Lepus granatensis grana-
tensis ; ear short (only 95 mm. instead of 105-115 mm.) ; skull
small and slender, its occipitonasal length 82 mm. instead of
85-90 mm.

Colour. — The colour so exactly resembles that of Lepus
granatensis granatensis as to require no special description.

Shull and teeth. — The skull differs from that of Lepus
granatensis granatensis in its distinctly smaller size (occipitonasal
length only 82 • 4 mm.), in the even less convex dorsal profile,
and the relatively more slender rostrum. The rostral depth at
front of tooth-row in the type specimen is 17 mm., while in
L. granatensis granatensis it ranges from 19 to 21 mm. Teeth
as in the typical form.

Measurements. — Type: hind foot, 110; ear from crown, 95.
For cranial measurements see Table, p. 521.

Specimen examined. — The type.

Biarritz, Basses-Pyrenees, J. E. Harting (p). 97. 6. 15. 1.

France. (Greig.) [Type of subspecies.)

519

LEPUS PARNASSIUS Miller.

1903. Lepus parnassius Miller, Proc. Biol. Soc. Washington, xvi, p. 145,
November 12, 1903. Type in U.S. National Museum.

1908. Lepus e[iLropxus] parnassius Hilzheimer, Jahreshefte des Vereins
fiir vaterl. Naturk. in Wiirttemberg, p. 395.

1910. Lepus europieus parnassius Trouessart, Faune Mamm. d'Europe,
p. 221.

Type locality. — Agoriani (Agokgianne), north side of Lyakuia
(Parnassus) Mountains, Greece.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — General appearance much as in Lepus creticus,
but size larger (hind foot about 135 mm., occipitonasal length of
skull about 95 mm.), and ear much longer (height from crown
about 130 mm. instead of 105 to 117 mm.), the black area on its
outer side noticeably more extensive ; skull with rostral portion
unusually long and slender ; no trace of m^ in the two specimens
known ; m^ with the two enamel cylinders broadly communi-
cating.

Colour. — General colour of upper parts a coarse grizzle of
black and pinkish buff, the latter slightly in excess, much less so
than the cream-buff of the corresponding region in Lepus europxus.
Actual colour bands as follows : (1) whitish smoke-grey, 12 mm.,
(2) black, 7 mm., (3) pinkish buff, 5 mm., (4) black, 5 mm., the
first and second on the underfur, the others on the longer hairs.
Sides and neck not noticeably different from back, but the
grizzle less distinct owing to the replacement of the black of
underfur by hair-brown. Rump essentially like back, but with
a slight, inconspicuous greyish cast due to the light grey bases
of the hairs. The exact shade of this undercolour is about
Ridgway's grey No. 10 at base of hairs, darkening to grey
No. 7 near surface. Cheeks like sides but more finely grizzled,
and with a faint blackish wash below ear. An indistinct greyish-
buft" eye-ring and loral stripe. Crown and face like back but
more finely grizzled, and bases of hairs wood-brown. Nape patch
ill-defined, an obscure russet wood-brown overlaid by the usual
buff. Ears light silvery grey (the exact colour not given by
Ridgway), except for the following markings : a very finely
grizzled stripe essentially concolor with top of head extending
up anterior outer surface almost to tip, and about 20 mm. wide
at middle; a similar area 45 mm. long by 12 mm wide near
middle of posterior inner surface ; a black apical patch 40 mm.
long by 30 mm. wide on posterior outer surface ; a black apical
area on inner surface about 10 mm. wide; a clear ochraceous-
buff area 10 mm. in width between apical black and general grey
of inner surface ; a whitish line 3 mm. in width along inner
anterior margin from base to above middle. Feet, outer surface
of legs, flank patches, and throat, dull ochraceous-buff. Under-

520

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522 RODENTIA

parts and inner surface of le,£?s white, much suffused with
ochraceous-buff on front legs. Collar very wide, essentially like
sides of body, but without black and with a decided tinge of
wood-brown.

Shill. — The skull differs from that of Lepus europseus in
smaller size and much less robust form. This is particularly
noticeable in the rostral region, which is relatively elongated and
narrowed, though not much reduced in depth. In the type the
distance from front of zygoma to gnathion is 41 "4 mm., while in
a Swiss skull of L. europseus with the same occipitonasal length
it is only 38 • 6 mm. The width and depth of rostrum at level of
point of junction between lower border of nasal and upper
border of premaxillary are respectively 16*0 mm. and 19*2 mm.
in the type and 18-2 mm. and 20 mm. in the Swiss specimen.
In other respects the skull shows no tangible characters to
distinguish it from that of L. europseus*

Teeth. — The teeth differ from those of all other European
hares in the absence, in the type, and a very young specimen
(in L. europseus of same age it is plainly visible) of m^, and in the
broad communication of the two cylinders of m^ along their
region of contact, this peculiarity combined with a general
reduction in the relative size of the tooth. In other respects the
teeth show no tangible characters to distinguish them from
those of L. europseus, though the posterior wall of the re-entrant
angle in the second, third and fourth upper cheek-teeth is thrown
more conspicuously into minute folds, like those of anterior wall,
than is usual in European hares.

Measurements. — Type (adult but not old male) : head and
body, 570± ; hind foot, 135 mm. ; ear from crown, 130 mm.
For cranial measurements see Table, p. 521.

Specimens examined. — Two, the type and a very young individual with
crowns of cheek-teeth not worn away, both from Agoriani (U.S.N.M.).

LEPUS TIMIDUS Linnteus.

(Synonymy under subspecies.)

Geographical distribution. — Northern portion of Europe and
Asia ; exact limits of range unknown. In Europe throughout
Norway, Sweden and northern Russia, also in Scotland and the
Alps.f Represented in Ireland by a distinct though nearly
related species.

Diagnosis. — Tail short, its length including terminal hairs
much less than that of hind foot, its upper surface white or
grizzled, never with black median area ; ear only moderately
long, extending when laid forward about to nostril or slightly

* In the type the basicranial region and auditory buUee are missing.
t Often referred to as occurring in the Pyrenees, but this undoubtedly
erroneous (see Trutat, Bull. Soc. Hist. Nat. Toulouse, xii, p. 110, 1878).

LEPus 523

beyond ; general colour of upper parts a finely grizzled greyish
brown (approaching the broccoli-brown and hair-brown of
Ridgway) in summer, white or pale grey in winter, the extreme
lip of ear always black ; root of upper incisor long, extending to
suture between premaxillary and maxillary ; height of posterior
upper premolar measured from crown to upper surface of root
capsule greater than alveolar length of tooth-row.

External characters. — Externally much as in Lejms europaeus
except for the shorter tail and ear, and more woolly fur.
Mammae : p 1-1, a 3-3 = 8.

Colour. — Summer pelage : genei'al colour of upper parts
between hair-brown and sepia, inconspicuously grizzled with
cream-buff, the sides not so dark as back, the rump sooty blackish ;
cheeks, muzzle and outer surface of legs tinged with buify ; nape
and back of neck to shoulders not grizzled, the general colour of
this area between wood-brown and broccoli-brown. The colours
on back are arranged as follows : underfur (10 mm.) light pearl-
grey at base changing to light wood-brown at middle ; longer
hairs (25 to 30 mm.) greyish at base, then dark sepia, each with
a cream-buff sub-terminal annulation about 2 mm. long and a
blackish slightly longer tip ; a few still longer hairs (35 mm.)
entirely blackish except the grey base ; still others in some
specimens jiure white. On sides and rump the grey portion of
underfur extends nearly or quite to tips of hairs. Tail bluish
white, the median region of upper surface an indefinite grizzled
brown. Ear blackish or brownish, the inner surface sprinkled
to a varying extent with whitish or buffy hairs, the outer margin
always bufiy whitish, the tip always clear black, this area either
confined to rim or extending down outer surface for a distance
of from 5 to 20 mm. Inner surface of legs and entire underparts
(except collar) white or whitish usually with a slight buffy tinge
and frequ.ently darkened by a sprinkling of blackish hairs. Chin
and interramia white or greyish, always contrasted with the
grizzled brownish collar, the latter essentially like back or some-
what paler. Feet dull buffy brown varied with white or cream-
buff. General hue of upper parts sometimes more yellowish,
owing chiefly to the wood-brov/n of underfur brightening toward
ochraceous-buff. Winter pelage white or grey. In the former,
more usual phase the animal is white throughout, except the
palms and soles, which are wood-brown, and the tip of ear, which
retains the narrow black area as in summer. Underfur every-
where white, occasionally with a faint greyish tinge at base, or
a slight buffy cast at tips of hairs. Rarely there is a slight
greyish wash along anterior border of outer surface of ear. Grey
phase : upper parts and sides light grey, clear and somewhat
drabby on sides of body, neck and cheeks, noticeably tinged with
wood-brown on face, crown, ears and back, the latter region
further darkened by a faint clouding of black ; region imme-
diately surrounding eye nearly white, this area forming a well

524 RODENTIA

defined band about 5 mm. wide over upper lid ; ear with same
colour pattern as in summer ; underfur of back and sides bluish
grey (about grey No. 8 of Kidgway) at base (10 mm.), the tips
of the hairs (5 mm.) wood-brown on back, light buffy or buffy
grey on sides ; longer hairs of sides M'hitish, those of back dark
brown or blackish, each with a conspicuous annulation (about
4 mm. wide) of whitish cream-buff, the extreme tip black ; outer
surface of legs irregularly washed with light buffy wood-brown ;
underparts white, a little tinged with buffy on throat, where an
ill-defined collar is produced by the buffy wood-brown of underfur ;
tail white, tinged with bluish grey above, the black tip of ear
remaining in all cases as in summer.

Skull. — ^In general form the skull resembles that of Or>/ctolngus
cuniculus. The size is, however, decidedly greater than in either
of the Eui'opean wild rabbits (occipitonasal length about 90 to
100 mm. instead of about 70 to 83 mm.) and the proportions are
more robust. There are also numerous differences in details of
structure. Most important of these are the peculiarities of the
postorbital processes, interparietal, and mesopterygoid region
already alluded to as generic characters. Brain-case relatively
shorter and wider than in Oryctolagus, its greatest width about
equal to distance from lambdoid suture at side of interparietal
to posterior base of postorbital process ; parietal region distinctly
flattened, the lateral convexity much less than in Oryctolagus
and Lepus europseus. Anterior half of frontal noticeably concave,
the orbital rims rising gradually but considerably above general
level of interorbital region. Malar less projecting posteriorly
than in Oryctolagus, its tip extending less than half way from
posterior border of glenoid surface to auditory meatus. Length
of exposed portion of suture between malar and zygomatic
process of squamosal about equal to height of zygomatic process
(in Oryctolagus it is much greater than height of process).
Upper border of malar thickened anteriorly, everted and
obliquely plate-like posteriorly ; elevated region in front of
lateral groove deep and narrow, the least distance from groove
to front of zygoma less than depth through same region.
Rostrum heavy and rather short, its depth to anterior rim of
alveolus of first cheek-tooth about equal to distance from latter
point to front of incisive foramen. Auditory bulla essentially as
in Oryctolagus, the anterior border more abruptly rounded off
than in Lepus europseus.

Teeth. — The teeth do not differ materially from those of
Lepus europseus. Incisors both above and below more squarish
in cross section ; groove on anterior face of i^ in same position
as in the more southern animal ; root extending to suture
between premaxillary and maxillary, the entire course of the
tooth clearly indicated on outer surface of bone. Anterior
upper premolar usually less narrowly elliptical in cross section
than that of L. europseus, its inner re-entrant angle appearing to

525

Fm. 103.
Lejms timidus scoticus. Nat. size.

526

RODENTIA

lie on inner border of crown rather than on inner side of
anterior border. Anterior section of first lower premolar longer
and narrower than in the related species. Other teeth not
certainly distinguishable from those of L. eiiropseus except that

the roots of the upper cheek-teeth
extend further into the orbital cavity ;
distance from crown of third pre-
molar to upper surface of root capsule
usually greater than alv^eolar length
of tooth-row.

Bemarks. — As a group the vary-
ing hares are readily distinguishable
from the other Eviropean members
of the genus by their shorter ears and
tail, on both of which the dark mark-
ings are reduced in extent, and by
their slight but evident cranial and
dental peculiarities. That these more
fundamental characters indicate a
higher degree of specialization than
that attained by Lepus europseus has been shown by Winge
(Danmarks Fauna, Pattedyr, p. 58, 1908). Lepus timidus is
represented in Europe by three local forms, which, though
isolated geographically, seem most conveniently treated as sub-
species. The Irish hare is completely diflferentiated from the
continental and British forms.

Fig. 104.
Lepus timidus. Teeth. X 2.

Lepus timidus timidus Linnseus.

1758. [Lep2is] timidus Linnseus, Syst. Nat., i, 10th ed., p. 57.

1777. [Leims timidus] alpinns Erxleben, Syst. Eegiii Anim., i, p. 328 (the

varying hares in general, from Switzerland to Greenland). Not
of Pallas, 1773, or of Erxleben, p. 337.

1778. Lepus variabilis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 2 (Renaming

of timidus).
1778. [Lepus] algidus Pallas, Nov. Sp. Quadr. Glir. Ord., p. 2 (Alternative

for alpinus Pennant).
1778. [Lepus] borealis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 2 (Alternative

for alpinus Pennant).
1795. L[eptis] septentrionalis Link, Beytrage zur Naturgesch., Bd. i,

Stiick 2, p. 73 (Substitute for variabilis).
182C. Leptis borealis Nilssou, Skand. Fauna, i, p. 211 (Substitute for

variabilis).
1827. [Lepus timidus] j8 alba Billberg, Synopsis Faunae Scandinavise, p. 7

(Northern Scandinavia). Not L. timidus albtis Bechstein, 1801.
1829-32. Lepus borealis collinus Nilsson, Ilium. Fig. Skand. Fauna, i,

pi. 19 (Sodermanland, Sweden).
1829-32. Lepus borealis sylvaticus Nilsson, Ilium. Fig. Skand. Fauna, i,

pi. 22 (Heavily wooded portions of Sweden).
1844. Lepus cancscens Nilsson, Ofversigt af Kongl. Vetensk.-Akad. For-

handl., Stockholm, i, p. 133 (Renaming of sylvaticus).

LEPUS 527

1857. Lepus variabilis b. Forra der Mittelregion uud der Alpen, Blasius,

Saugethiere Deutschlands, p. 424 (part).
1910. Lcpus timidus and L. timidus collinus Trouessart, Faune Mamni.

d'Europe, p. 216.

Type locality. — Upsala, Sweden.

Geographical distrihution. — Scandinavian Peninsula.

Diagnosis. — Largest of the varying hares of western Europe,
the occipitonasal length of adult skulls ranging from 95 to
103 mm.

Measurements. — Adult from Helsingland, Sweden : hind foot,
150 ; ear, 94. Adult male from Stockholm, Sweden : hind
foot, 155 ; ear from crown, 105. Adult male from Hellestad,
Ostergotland, Sweden: hind foot, 151 ; ear from crown, 99. For
cranial measurements see Table, p. 532.

Specimens examined. — Thirty-two, from the following localities : —
Sweden : Helsingland, 1 (U.S.N.M.) ; Kronen, Vesterbotten, 7

(U.S.N.M.) ; northern Sweden, no exact locality, 1 skeleton (U.S.N.M.) ;

near Stockholm, 1 (U.S.N.M.); Furusund, Stockholm, 13 (U.S.N.M.);

Hellestad, Ostergotland, 1 (U.S.N.M.) ; Lund, Skane, 1 (U.S.N.M.) ; no

exact locality, 1.

Norway : Hammerfest, 1 ; Enebek, 1 ; Eidsvold, Christiania, 2 ;

near Christiania, 1 ; Jeederen, Stavanger, 1 ; Holme, Mandal, 2.

Memarlcs. — The typical form of Lepus timidus is readily
distinguishable from the Alpine and Scotch races by its larger
size, particularly as shown by the length of the skull. "Whether
more than one local race should be recognized among the
Scandinavian members of the species must still be regarded as
an open question owing to the absence of adequate material. It
appears to be a fact, however, that the grey winter coat, though
not invariably assumed by the animals of the region where it
occurs, is met with nowhere except in southern Sweden and the
extreme south of Norway west to Jsederen. Correlated with the
occurrence of the grey winter coat appears to be the tendency
for the black at tip of ear to .spread downward from the rim and
form a noticeable patch on outer surface. The names sylvatictis
and canescens of Nilsson were based on the southern animal,
while the horealis and collinus of the same author were applied to
the northern form.*

Stockholm Museum (e). 46. 6. 2. 73.

Christiania Museum (e). 93. 3. 1. 21.

Christiania Museum (e). 0. 2. 7. 1-2.

Christiania Museum (e). 93. 3. 1. 20.

Christiania Museum (e). 0. 5. 2. 2.

n. J. Cuninghame (c & p). 8. 8. 9. 38-39.

1.

Sweden.

6.

Enebek, Norway.

2 6.

Eidsvold, Christiania.

6.

Christiania.

6.

Jpederen, Stavanger.

(J juv, 9.

Holme, Mandal.

* A good account of the two races is given by CoUett, Norges Pattedyr,
pp. 58-59, 1911.

528

Lepus timidus varronis Miller,

1857. [Lepus variabilis'] b. Form der IMittelregiou und der Alpen, Blasius,

Saugethiere Deutschlands, p. 424 (part).
1901. Lejms varronis Miller, Proc. Biol. Soc, Washington, xiv, p. 97,

June 27, 1901. Type in U.S. National Museum.
190G. Llep^is] vi[edi'us'] varronis Hilzheimer, Zool. Anzeiger, xxx, p. 511,

August 14, 1906.
190G. Llcpusl mledius] brcviaiiritus Hilzheimer, Zool. Anzeiger, xxx,

p. 511, August 14, 1906 (Bernese Alps, Switzerland). With regard

to status see Mottaz, Bull. Soc. Zool. de Geneve, 1908, pp. 172-174,

November 15, 1908.
1910. Lepus medius varronis Trouessart, Paune Mamm. d'Europe, p. 218.

Type locality. — Hinzenberg, Grisons, Switzerland.

Geographical distribution. — Alp.s and adjacent mountain
chains, mostly at elevations above 1,300 m.

Diagnosis. — Decidedly smaller than Lepus timidus timidus,
the occipitonasal length of skull rarely if ever reaching 95 mm.
(usually 85 to 93 mm.) ; summer pelage lighter and more greyish ;
winter pelage always white, the middle of back usually with a
faint greyish wa^h ; anterior portion of outer boi'der of ear
nearly always buffy gx'ey ; black area at tip of ear confined to rim.

Measurements. — Type (adult male) : head and body, 582 ;
tail, 53; hind foot, 144-6; ear from crown, 106. Two adult
males from Grisons, Switzerland : head and body, 570 and 585 ;
tail, 65 and 50; hind foot, 148 and 141; ear from crown,
98 and 101. Two adult females from the same region: head
and body, 575 and 610 ; tail, 55 and 56 ; hind foot, 146 and 142 ;
ear from crown, 100-5 and 99. Adult female from Padola,
Cadore, Italy : hind foot, 142 ; ear from crown, 101. Two males
from Col-de-Vars, Hautes-Alpes, France : hind foot, 138 and
139. For cranial measurements see Table, p. 533.

Sp)ecimens examined : — Twenty-four, from the following localities : —

France : Col-de-Vars, Hautes-Alpes, 2 ; above Barcelonnette, Basses-
Alpes, 2 (Mottaz) ; Petit Bornand, Haute-Savoie, 2 (Mottaz).

Switzerland : Canton St. Gallen, 3 ; Hinzenberg, Grisons, 1
(U.S.N.M.) : Lugnetz, Grisons, 2 (U.S.N.M.) ; Obersaxen, Grisons, 1
(U.S.N.M.); Untervatz, Grisons, 1 (U.S.N.M); Grisons, no exact locality,
5 (B.M. and U.S.N.M.) ; Centovalli, Ticino, 1 (U.S.N.M.) ; San Lucio,
Val CoUa, Ticino, 1 (U.S.N.M.).

Italy: Padola, Cadore, 2 (Turin); Dronero, Cuneo, 1.

Bemarhs. — The Alpine varying hare is a strongly characterized
race, which should perhaps be treated as a species distinct from
Lepus timidus. The winter coat appears to be always white or
incompletely white ; a special blue grey winter pelage like that
of " sylvaticus " is unknown.
2 6. Col-de-Vars, Hautes-Alpes, O. Thomas (p). 8.8.10.131-132.

France. (C. Mottaz.)
9. St. Gallen, Switzerland. O. Thomas (p). 2. 8. 4. 52.

[e. H. Zollikofer.)
6,2 9 Grisons. {E. H. Zollikofer.) O. Thomas (p). 2. 8. 4. 49-51.

1. Dronero, Cuneo, Italy. W. Playters Stark 11. 1. 2. 111.

(c & p).

529

Lepus timidus scoticus Hilzheimer.

1816. ■? Lepus albus Leach, Syst. Catal. Spec. Indig. Mamm. and Birds,
Brit. Mus., p. 7 (nonien nudum: "White Hare"). Not Lepus
timidus albus Beclistein, 1801.

1833. Lepus albus Jenyns, Man. Brit. Vert. Anim., p. 35. Not L(pus
timidus albus Bechstein, 1801.

1857. [Lepus variabilis] h. Form der Mittelregion und der Alpen, Blasius,
Siiugethiere Deutschlands, p. 424 (part).

1900. Lepus timidus typicus Barrett-Hamilton, Proc. Zool. Soc, London,
p. 88 (part).

1906. L\_cpus'] m[edius'] scoticus Hilzheimer, Zool. Anzeiger, xxx, p. 511,
August 14, 1906.

1910. Lepus ti7nidus scoticus Trouessart, Faune IMamm, d'Europe, p. 218.

Type locality. — Northern Scotland.

Geographical (listrihution. — Highlands of Scotland ; range now
extended by artificial introduction irregularly into Wales and
northern England, also in Ireland.

Diagnosis. — Size even less than in Lepus timidus varronis, the
occipitonasal length of fully adult skulls ranging from 83 to
89 mm. ; ear shorter than in varronis, its length from crown
80 t(t 90 mm., instead of 90 to 100 mm. ; winter pelage white,
but apparently never without some trace of grey.

Colour. — ^Summer pelage essentially indistinguishable from
that of Lepus timidus timidus, though perhaps not attaining
the same degree of sepia brown as in extreme specimens of the
Scandinavian race, and underparts usually more bufFy. the
interramia less contrasted Avith collar. Winter pelage apparently
never clear white, the middle of back in all the specimens
examined showing at least a faint grey tinge as compared with
the snowy white of thighs and tail, this grey usually suft'using
the entire dorsal surface. It is produced by the presence among
the white hairs of a few that are entirely black, and others that
are black with a light cream-buft" sub-terminal annulation as in
summer pelage. Underfur very pale buft", this colour appearing
at surface when hairs are disarranged. Ear with anterior outer
surface conspicuously grizzled (much more than in varronis),
though usually j^aler than in summer, the basal half often mostly
white. Sides of muzzle usually tinged with buffy.

Skull and teeth. — The skull is immediately distinguishable
from those of the other European forms of Lepus timidus by its
small size. The teeth show no tangible peculiarities.

Measurements. — Adult male and female from Cromlix, Dun-
blane, Perthshire : head and body, 482 and 480 ; tail, 63
and 57; hind foot, 130 and 132; ear from crown, 85 and 83.
Adult male from Altyre, Morayshire : head and body, 501 ;
tail, 57 ; hind foot, 131 ; ear fi-om crown, 90. Adult from New
Galloway, Kirkcudbright : head and body, 535 ; tail, 58 ; hind

2 M

530 RODKNTIA

foot, 133 ; ear from crown, 77. For cianial measurements see
Table, p. 533.

Specimens exumiiied. --Fovty, from the following localities in Scotland :
Boi-gie, Sutherland, 1 ; Nairn, 5 ; Forres, Elgin, 1 ; Altyre, Morayshire, 5 ;
Dallas, Morayshire, 2 ; IMorayshire, 3 ; Glengroullie, 3 ; Gromlix, Dunblane,
Perthshire, 8 ; Caparoch, Selkirkshire, 1 ; Thornhill, Dumfriesshire, 2 ;
New Galloway, Kirkcudbright, 2 ; Clonas, 4 ; no exact locality, 3 (B.M.
andU.S.N.M.').

Reiuarhs.' -While readily distinguishable from true LepUH
timidus by its smaller skull, the Scotch hare rather closely
resembles the Alpine race. Its ears are, however, noticeably
shorter, and the winter pelage rarely if ever becomes so nearly
pure white. As in L. timidus ran-oni)^ no special grey winter coat
is known.

Individuals supposed to be hybrids between this animal and
Lepm eurojjseus occidentalis are sometimes unusually heavy
examples of scoticus, and in other instances occidentalis in grey
winter pelage. After eliminating these, however, a few specimens
remain in which the characters of the two animals appear
to be blended. The British Museum contains three such
skins.* In each of these the ear is longer than in scoticus, and
its colour pattern nearly as in occidentalis, though less distinct ;
the tail is short as in scoticus, but its dorsal surface has a nearly
black median area ; in two the body colour is intermediate
between that of the supposed parent species in summer coat,
while in the other (No. 63. 8. 23. 1) it more nearly approaches
that of the common hare.j

1.

Borgie, Sutherlandshire,

Rev. H. H. Slater

0. 2. 24. 3.

Scotland.

(c & P).

6,2 9.

Nairn.

Earl Cawdor (p).

98. 11. 21. 1-3.

99. 5. 1. 1-2.

1.

Knockie, Inverness-shire.

E. Hargitt (p).

86. 9. 9. 8.

1.

Forres, Elginshire.

Sir W. Gordon Gum-
ming (p).

3. 1. 27. 1.

4 c?, 1 9.

Altyre, Morayshire.

Sir W. Gordon Gum-

93. 12. 4. 1-3.

ming (p).

94. 2. 15. 1-2.

3 St.

Morayshire.

Sir W. Gordon Gum-
ming (p).

97. 12. 27. 1-3.

2 St.

Dallas, Morayshire.

Sir W. Gordon Gum-
ming (p).

1. 1. 15. 1-2.

3.

Glengroullie.

Earl Gawdor (p).

99. 3. 20. 1-3.

G 6, 2 9.

Dunblane, Perthshire.

Gapt. Hon. A. Hay

92. 2. 15. 2-5.

Drummond (p).

97. 5. 13. 1-3.

2.

Selkirkshire.

Rev. H. H. Slater
(c & P).

0. 2. 24. 2-3.

* No. 63. 8. 23. 1. No history.

No. 2. 11. 28. 1. Male, Graigmyle, Aberdeenshire, Scotland.
No. 6. 12. 26. 1. Male, Gapenoch, Thornhill, Dumfriesshire, H. S.
Gladstone.

t Swedish specimens combining the characters of timidus and euro-
pneus have been described and figured by Lonnberg, Proc. Zool. Soc,
London, 1905, ii., pp. 278-287.

LEl'US 531

1. Thornhi!!, Dumfi-iesshire. H. S. Gladstone 7.8.7.1.

(c&p).
;2. New Galloway, Kircud- Col. Gordon Maitland 95. 10. 6. 1-2.
brightshire. (c & p).

6, 3 9. Clonas. W. R. Ogilvie-Grant 99. 2. 17. 1-4.

LEPUS HIBERNICUS Bell.

1833. Irish Hare Yarrell, Proc. Zool. Soc, London, p. 88. No technical

name.
1837. Lepus hibcrnicus Bell, History of Brit. Quadrupeds, p. 341.
1857. [Lepus i;ariabilis'] a. Form der warmeren Klimate, Blasius, Siiuge-

thiere Deutschlauds, p. 424 (part).
1900. Lepus timidus hiberniciis Barrett-Hamilton, Proc. Zool. Soc,

Loudon, p. 89.
1900. Lepus timidus lufesccns Barrett-Hamilton, Proc. Zool. Soc, London,

p. 89 (Donobate, Co. Dublin, Ireland). Type in British IMuseum.
190G. Llepus] t[imidus'] hibernicus Hilzheimer, Zool. Anzeiger, xxx, p. 510,

August 14, 1906.
1910. Lepus timidus hibcrnicus and L, timidus lutescens Trouessart, Paune

Mamm. d'Europe, p. 216.

Tijije locality. — Ireland.

Geograpliical dlstrUmtion. — Ireland, chiefly in the more hilly
districts. Now introduced and established in portions of Wales
and Scotland (Carnarvonshire and the island of Mull).

Diagno^'iH. — Differing from the other European members of
the Lepus timidus group in its strongly russet colour and in the
partial or complete absence of the white winter coat. Size
noticeably greater than in its nearest geographical ally,
L. timidus scoticus.

Colour. — General colour of upper parts a finely and incon-
spicuously grizzled russet brown, aj^proaching the russet and
wood-brown of Ridgv/ay, brighter and more reddish on shoulders
and back, paler, though strongly russet, on crown and face,
duller and fading to ochraceous-bufi' on cheeks, sides, and outer
surface of legs ; nape and back of neck to between shoulders
scarcely grizzled, the nape usually with a suffusion of drab. The
colours on the back are arranged as follows : underfur (15 mm.)
light pearl-grey (about grey No. 10 of Eidgway) at base, the
terminal half of the hairs abruptly light russet ; loiiger hairs
(25 mm.) light pearl-grey at extreme base, becoming black at
level of russet portion of shorter hairs, each with a dull buff sub-
terminal annulation about 4 mm. long and a rather shorter black
tip ; a few still longer haix's (35 mm.) entirely black except the
grey base. Rump usually with an evident greyish cast, this
often rather conspicuous in winter specimens, the black tips to
the hairs occasionally ])roducing a markedly clouded effect. Tail
white, usually with a bluish tinge, the upper surface sprinkled
to a varying degree Avith blackish or brownish hairs, these
occasionally producing a decided grizzled effect, and never

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entirely absent. Ear grizzled ochraceous-buff", riinined with
black at tip, the inner surface becoming clear ochraceous-buff
sub-terruinally, the posterior outer surface washed with grey,
and with a bufFy or whitish border from base to black terminal
area ; antei"ior border occasionally whitish near middle. A
faintly defined pale eye-ring, but no other light markings on
side of head. Inner surface of legs and entire uuderparts
(except collar) white, slightly tinged with bufFy, especially on
inner side of thigh ; inguinal region slightly clouded ^\'ith a
mixture of blackish and bufFy ; collar between ochraceous-
bufF and clay-colour though paler than either, the grey (about
No. 7 of Eidgway) underfur appearing irregularly at surface.
Feet ochrnceous-bulF, duller and somewhat grizzled on meta-
podials, clear and brighter on toes. Soles an indefinite bufFy drab.

Skull and teeth. — The skull difFers from that of Ltpus
thnidus scoticus in the larger sizes attained by fully adult
individuals, the occipitonasal length in such specimens ranging
from 91 to 94 mm., Avhile in the Scotch hare it seldom exceeds
88 mm. Similarly the mandible in Mhernicus is usually more
than 70 mm. in length, while in scoticus it rarely if ever reaches
70 mm. The brain-case in the Irish hare often appears more
elongate than in the Scotch form, but the difFerence is not
constant. Otherwise I can detect no cranial or dental
peculiarities by which the two species may be distinguished.

Measurements. — Adult female from Arthurstown, Waterford,
Ireland: head and body, 517; tail, 40; hind foot, 139.
Average and extremes of six specimens from LisdufF, Co. Cavan ;
ear from crown, 90 "3 (85-95). For cranial measurements see
Table, p. 533.

Specimens examined. — Twenty-nine, from the following localities : — ■

Ireland : Gosford Castle, Armagh, 2 ; Lisduff, Co. Cavan, 12 ; Rath-

robbin, King's County, 2; Arthurstown, ^Yaterford, 1; Kilmanock,

Waterford, 8 ; no exact locality, 2. Douobale, Dublin, 1 (type of lutescens).

Wales: Vaynol Park, near Bangor, Carnarvonshire, 1 (introduced).

HemarJcs. — The Irish hare is strikingly difi'erent from the
other European members of the thnidus group in its peculiar
russet colour. In size it noticeably exceeds the Scotch hare, but is
itself surpassed by the Scandinavia,n true timidus. The yellowish
specimens from Co. Dublin, to which the name lutescens has been
applied, occurring, as they do, together with animals of normal
colour, appear to be examples of some pathological or dichromatic
condition rather than the representatives of a recognizable
geographic race.

2. Gosford Castle, Armagh, Hon. L. Powys (c &r). 76. 1. 25. 1-2.

Ireland.
4. Lisduff, Cavan. Hon. B. E. B. Fitz- 76. 3. 26. 1-2.

Patrick (c & p). 76. 5. 8. 1-2.

4. Lisduff, Cavan. Hon. B. E. B. Fitz- 76. 7. 12. 1-2.

Patrick (c & p). 76. 9. 7. 1-2.

SIMPLICIDENTATA

535

4. Lisduff, Cavan. Hon. B. E. B. Fitz 70. 11. 18. 1-2.

Patrick (c & p). 77. 1. 18. 1-2.

2 6, St. Rathrobbin, King's County. Col. M. W. Biddulpb 98. 2. 10. 1-2.

(c & p).

st. Donabale, Dublin. Charles Cobbe(c&p). 82.2.4.1.

{Type of L. lutesccns Barrett-
Hamilton.)

?,. Kilmanock, Waterford. G. Barrett-Hamilton 99. 2. IG. 1-3.

(c k p).

2(^,9. Kilmanock, Waterford. G.Barrett-Hamilton 9.12.15.1-3.

(c & p).

6,9. Waterford. G.Barrett-Hamilton 9.10.4.1.

(c & p). 8. 2. 27. 1.

1. Bangor, Carnarvonshire, G. W. Assheton 3.1.28.1.
Wales. Smith (c & p).

Sub-Order SIMPLICIDENTATA.

1891. Sim.plicidcntata Flower and Lydekker, Mammals, Living and
Extinct, p. 448.

GeograpliiraJ ilhtrihution. — Same as that of the order
Modentia.

Characters. — Upper incisors 1—1, their enamel covering not
extending to posterior surface ; distance between mandibular and
maxillary tooth-rows approximately equal, both pairs of rows
capable of opposition at the same time, the primary motion of
the jaws in mastication longitudinal or oblique ; premolars never
more than f;f- ; incisive foramina moderate or small, not confluent
posteriorly ; bony palate never reduced to a narrow bridge
between premolars ; facial portion of maxillary entire ; fibula
not articulating with calcaneum.

Bemarks. — This sub-order contains the vast majority of exist-
ing I'odents, among which about twenty families are now recog-
nized. Eight of these occur in Europe (for keys see pp. 481-483).

Family ZAPODID^.

1875. ZapodicLv Cones, Bull. U.S. Geol. and Geogr. Surv. Terr., 2d ser., i,

p. 253.
1901. Zapodichv Lyon, Proc. U.S. Nat. Mus. xxxin, p. 666, May 2, 1901.

Geographical distrihitlon. — Boreal portions of both hemi-
spheres, east in Europe to Norway.

Characters. — Skull and teeth with a genei'al resemblance to
those of the Dlpodidse though less highly modified ; but infra-
orbital foramen similarly lai-ge and with distinct supplemental
canal along inner border, and jugal in contact with lachrymal ;
hind legs much less specialized than in the related family, the
foot with five distinct metatarsals.

Bemarks. — The family Zapodldae, though related to the

536

RODENTIA

Dipodtdse,* is readily distinguished by the more primitive murine
character of the skull and hind foot. It is divisible into two
main groups, the Zapodime with two genera in North America
and one in China, and the Sichtinefi with a single genus peculiai-
to the Old World.

Sub-Family SICISTINiE.

1857. Murina Blasius, Saugethiere Deutsclilauds, p. 299 (part).

1887. Sminthi (section of Dipodini) Winge, Jordfundne og nulevende Gna-

vere (Eodentia) fra Lagoa Santa, Minas Geraes, Brasilien, p. 122.
1901. Smintldmv Lyon, Proc. U.S. Nat. Mus. xxxiii, p. 666, May 2,1901.

Geographical distribution. — Boreal portion of eastern hemi-
sphere, exact limits of range not known ; in Europe west to
Norway.

Characters. — Form strictly murine, the hind legs not elongated,
cheek-teeth tuberculate.

Remarhs. — The sub-family Sicistinse, a strictly Old World
group less sjDecialized than the Zapodinse, contains the single
genus Sicista, two members of which occur in Europe.

Genus SICISTA Gray.

1827. Sicista Gray, Griffith's Cuvier, Animal Kingdom, v, p. 227 {Mjis

subtilis Pallas).
1839. Smintlius Nathusius in Nordmann, Demidoff's Voyage dans la

Russie JMeridionale, iii, p. 49 [S. lorigcr Nordmann).
1857. Smintlius Blasius, Saugethiere Deutschlauds, p. 301.
1901. Sicista Allen, Proc. Biol. Soc. Washington, xiv, p. 185, December 12,

1901.

I'ype species. — Mus suhtilis Pallas.

Geographical distrihution. — From central Asia to Denmark
and southern Norway. Details of distribution imperfectly
known. For the possible occurrence of a member of the genus
in the Orkney Islands see Major, Zool. Garten, XLVi, pp. 129-134,
Moay 1905.

Characters. — ^As in the sub-family Sicistinfe. Dental formula :
*' fii' ^"* o-o' "* -Ti = ^^ J crowns of cheek-teeth with complicated
enamel pattern which does not form transverse ridges ; lai'ger
teeth with four well developed tubercles.

Remarhs. — The genus Sicista is at once recognizable among
European rodents by its murine external characters combined
with the large infraorbital foramen, low-lying zygoma and 5—5
upper cheek-teeth the tubercles of which are arranged in two
longitudinal series.

The species are very imperfectly known. About eight are
currently recognized ; two of these occur in Europe.

* Waterhouse, Ann. and Ma^. Nat. Hist., x., p. 203, November, 1842.

537

KEY TO THE EUROPEAN SPECIES OP SICISTA.

Least depth of rostrum behind iucisors about equal to

breadth of rostrum in same region ; crown area of m^

about four times that of premolar ; sides distinctly

more yellow than back, with rather noticeable

line of demarcation (Roumania) S. lorigcr, p. 537.

Least depth of rostrum behind incisors decidedly greater

than breadth of rostrum in same region ; crown

area of m^ about three times that of premolar ; sides

essentially concolor with back (Hungary, Denmark,

south-eastern Norway) .S'. irt^ona, p. 539.

SICISTA LORIGER Nathusius.

1840. Sminthus loriger Nathusius in Nordmann, Demidoff's Voyage dans

la Russie M6ridionale, iii, p. 49 (Odessa).
1840. Sminthus nordmanni Keyserling and Blasius, Wirbelth. Europas.,

p. 38 (Crimea).

Tyjjc localitij. — Odessa, Russia.

Geoiirapliical distribution. — South-westevn Russia, eastern
Roumania, Bulgaria ; limits of i-ange not known.

Diagnosis. — Least depth of rostrum behind incisors about
equal to width of rostrum in same region ; crown ai^ea of m^
at least four times that of premolar ; sides distinctly more yellow
than back, with rather well-defined line of demarcation.

External characters. — General appearance essentially as in
Mus muscnliis, ])ut size less (about as in 3Ius sjjicilegus), and body
more slender ; tail slightly longer than head and body, obscurely
four-sided, the fine hairs not concealing annulations (of which
there are about twenty to the centimeter at middle of tail) and
not forming any decided pencil at tip ; ear moderately large,
extending a little beyond eye when laid forward, its outline
simple, ovate, its posterior border with large thickened lobe at
base, capable of completely closing the meatus, its anterior
border with minute though evident sub-basal notch and projec-
tion ; both surfaces of ear densely clothed with short hairs ;
basal lobe conspicuously tufted ; anterior border of nostril
thickened, a small wart beneath it ; space between nostrils
narrow, crossed by a well developed vertical ridge ; upper lip not
grooved in front ; feet more slender than in Mus iimsculus and
its allies, the fingers and toes longer relatively to palm and sole ;
front foot with thumb reduced to a mere tubercle to support the
short but evident blunt Jiail ; third digit longest, fourth slightly
shorter, second extending to middle of third, fifth just beyond
middle of fourth, pads as in Mus inusciilas ; hind foot with
relative lengths of digits as in Mus musculus, the second, third
and fourth sub-equal and longest, fifth extending nearly to
middle of fourth, first not quite to base of second ; claws
essentially like those of front feet ; sole naked, the pads
arranged as in Mus musculus ; postero-internal tubercle extended

538 RODENTIA

backward so that its length is slightly more than double its
greatest width. Mammte : p 2-2, i 2-2 = 8.

Colour. — Back a peculiar light brown perhaps best described
as a buffy, greyish isabella-colour, tinged with ecru drab on sides
of neck, and everywhere slightly clouded (not "lined") with
black. Median dorsal region with a sharply detined black stripe
about 2 mm. wide thrown into relief posteriori}! by a distinct
lightening of groutd colour along its edges. tSides ochraceous-
buff, not strongly contrasted with back, but distinctly more
yellow and with rather sharply-drawn line of demarcation ;
below the colour of sides passes insensibly into the lighter, less
yellow ochraceous-buff of belly. Ear black sprinkled with bufty
hairs and with a narrow butty edge. They are rather strongly
contrasted with surrounding parts, this effect heightened by the
presence of an ill-defined though evident patch behind ear.
Feet a light indefinite buffy brown rather lighter than belly, the
toes a little more pallid. Tail obscurely bicolor, sepia above,
dull cream-buff below. Longer hairs of back black to extreme
tip ; basal colour everywhere slaty.

Sl^uJl and teeth. — The skull difiers from that of S. trizona

Fig. 105.
Sicista loriger. Xat. size.

FIG. lOG.
Sicista loriger. Xeetli. x 9.

merely in the somewhat more slender rostrum ; least depth
behind incisors about equal to width in same region. Teeth as
in the related species, but discrej^ancy in size between premolar
and first molar more pronounced, the crown area of the smaller
tooth about one-fourth that of the larger.

Measurements. — Three adult females from Malcoci, Dobrudscha
(in alcohol): head and body, 55, 56 and 58 ; tail, 69, 75 and 76 ;
hind foot, 14, 15 and 15; ear from meatus, 11, 12 "4 and 12.
For cranial measurements see Table, p. 541.

Specimens examined. — Six, five from Dobrudscha (B.IM. and U.S.N.M.),
and one from Bulgaria, no exact locality (Lataste).

BemarJcs. — The different specimens show no individual
variations worthy of note. From the better-known animal of

sicisTA 539

Hungary, Denmark and Norway this species is immediately distin-
guishable by the contrast between the colour of sides and back.

1. Dobrudscha, Eoumania. Purchased (Pruliere). 86. 4. 2. d.

SICISTA TRIZONA Petenyi.

1857. Sminthus vagus Blasius, Siiugethiere Doutschlands, p. 302 (probably
not Mhs vagus Pallas).

1882. Mus trizonus Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Hungary).

1882. Mus interzonus Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Alterna-
tive for trizonus).

1882. Mus intcrstriatus Petenyi, Termeszetrajzi Fiizetek, v, p. 103
(Alternative for trizonus).

1882. M^i.s tripartitus Pet6nyi, Termeszetrajzi Fiizetek, v, p. 103 (Alterna-
tive for trizomis).

1882. ikfws virgulosus Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Alterna-
tive for trizonus).

1882. [Mus] tristriatus -Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Alter-
native for trizonus).

1910. Sicista suhtilis Trouessart, Faune ]\Iamm. d'Europe, p. 205 (probably
not Mus suhtilis Pallas).

Type localiti/. — Hungary.

Geographical distrihnt'wn. — Hungary ; Denmark ; * south-
eastern ISTorway.f Details of distribution very imperfectly
known.

Diagnosis. — Least depth of rostrum behind incisors decidedly
greater than breadth of rostrum in same region ; crown area of
m^ about three times that of premolar ; sides essentially concolor
with back.

Colour. — Back, sides and head uniform raw-umber with a
slight buffy tinge, this more noticeable on cheeks and sides of
neck than elsewhere ; back very inconspicuously " lined " with
black. The individual hairs of the underfur are slate-colour at
base (5 mm.), the tips (1-5 mm.) buffy raw-umber; longer hairs
(12 mm.) black, the tips (2 mm.) colourless and with a silvery
gloss. Median dorsal line with clear black stripe about 3 mm.
wide, extending from crown to base of tail, better defined
posteriorly (except within a few mm. of base of tail) than
anteriorly, but not thrown into relief by lightening of background
along any part of its extent. Underparts light ochraceous-buff
without line of demarcation along sides, much dulled by
appearance at surface of slaty under colour. Ears essentially
concolor with back, but with a stronger admixture of black, the
rim clear light buffy. Feet a light indefinite buffy brown rather
paler than belly, the toes still paler. Tail obscurely bicolor,
sepia above, dull cream-buff below.

SlmU. — In general appearance the skull suggests that of

* See Winge, Danmarks Fauna, Pattedyr, p. 65, 1908.
t Sec Collett, Norges Pattedyr, pp. 68-77, 1911.

540

RODENTIA

Mus miisculus, but brain-case more squarish in outline, rostrum
slightly less elongate, and dorsal profile convex throughout, more
strongly so posteriorly than anteriorly. Interpariental narrow,
the lateral extremities pointed. Auditory bulhti rather large, well
inflated, their greatest transverse diameter along squarely truncate
anterior border. Interorbital region much broader than rostrum,
nearly flat, its edges faintly angular. Zygoma abruptly sjjreading
anteriorly, much deeper in front than behind, its lower edge
horizontal and at level of alveolar line ; anteorbital foramen large,
much wider below than above. Rostrum sloping anteriorly, but
rather narrow, the least depth just behind incisors decidedly
greater than width at same region ; nasals convex anteriorly, flat
posteriorly, the posterior border squarely truncate at level of
middle of lachrymal, the nasal bi'anches of premaxillaries slightly
longer. Incisive foramina very large, extending from about
1 mm. behind alveolus of incisor to level of front of m^, the
width of each foramen slightly greater posteriorly than anteriorly.
Palate rather large, flat throughout, extending at full width
conspicuously behind last molars, rather more than half of its
surface formed by the palatine bones ; a noticeable transverse
ridge at level of middle of h;^, and high narrow longitudinal
ridge extending from this to posterior border of palate ;
mesopterygoid space about one-half longer than broad, nearly
parallel sided, the hamulars slightly bowed outward and coming
in contact with antei"o-internal angle of bullte ; ectopterygoid well
developed, forming a conspicuous, neai-ly horizontal plate, the
rather large pterygoid fossa, therefore, shallow. Mandible slender,
with no special peculiarities, the well developed coronoid process
rising a little above level of condyle ; angular process long,
concave on inner side, the lower border bent abruptly upward
and outward at nearly a right angle.

Teeth. — Upper premolar single-rooted, the crown terete, with
high, narrow antei-o-external cusp, and low ridge-like internal
cusp, the three cusps separated by two evident furrows. Its area
is about one-third that of succeeding tooth. First upper molar
3-rooted, the general outline of crown squarish, slightly longer
than wide and slightly wider externally than internally ; enamel
of crown thrown into four main cusp-like folds, the outer of which
are larger than the inner, and three secondary folds extending
outward between and alternating with outer cusps. Second molar
like first but slightly longer and a little narrower in proportion
to its width. Third molar slightly larger than premolar, and of
about the same shape, the two anterior cusps small but evident,
the two posterior scarcely more than slight elevations on rim
of tooth. Lower molars essentially like the upper teeth, but
ci'owns longer and less squarish in outline, and pattern of folding
reversed, the intermediate folds extending inward instead of out-
ward. Third lower molar lai'ger than the coiresponding upper
tooth.

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542 RODENTIA

Measurements.— Adult male from Vestervig, Denmark : head
and body, 65 ; tail, 87 ; hind foot, 16"4 ; ear from meatus, 10 "6.
For cranial measurements see Table, p. 541.

Specimens examined. — Five, from the following localities : —
Austria-Hungary : Zuberec, northern Hungary, 2 ; Tatra Mts.,
northern Hungary, 2.

Denmark: Vestervig, Thj% 1.

BemarJcs. — Until it is possible to compare the two European
species of Sicista with authentic specimens of the Asiatic forms
named by Pallas it seems preferable to use the names trizona and
loriger, the status of which admits of no doubt.

2. Zuberec, Hungary. Budapest Museum (e). 94. 3. 1. 73-74.

1 al. Tatra Mountains. Zoological Society. 87. 1. 4. 1.

1 al. Tatra Mountains. Dr. R. Collett (p). 91. 1. 21. 3.

1 al. Vestervig, Thy, Den- Copenhagen Museum (e). 88.1.9.1.

mark.

FAMILY HYSTRICID.^.

1821. Hystridw Gray, London Med. Repos., xv, p. 804, April 1, 1821.
1831. Hystricidx. Burnett, Quart. Journ. Sci. Lit. and Art, xxviii, 1829,
p. 350, 1880.

Geographical distribution. — Warmer portions of the Old
World, Madagascar, Australia and the outer portions of the
Malay Archipelago excepted. In Europe confined to the
Mediterranean region.

Characters. — Middle portion of zygoma formed exclusively by
the large jugal bone which does not come in contact with
lachrymal ; anteorbital foramen large, without secondary canal ;
mandible with angular part arising from outer side of alveolus
of incisor ; frontal region of skull inflated (especially in genus
occurring in Europe) ; clavicle incomplete ; ^tibia and fibula
distinct ; teeth hypsodont, incompletely rooted ; body heavy,
covered with long quills.

BemarJcs. — The family Hystricidse contains four or five genera,
one of which reaches the Mediterranean region of Europe.

Genus HYSTRIX Linn«?us.

1758. Hystrix Linnaeus, Syst. Nat., i, 10th ed., p. 56 (cristata by tautonymy).
1798. Histrix Cuvier, Tabl. Element, de I'Hist. Nat. der Anim., p. 170

(Modification of Hystrix).
1866. QSdocephalus Gray, Proc. Zool. Soc. London, p. 308 {Acanthiun

cuvieri Gray).

Type species. — Hystrix cristata Linnaais.

Geographical distribution. — Africa and locally in Mediter-
ranean region of Euroi^e.

Characters. — Inflation of facial region of skull maximum for
the family, the contrast between depth at front of tooth-row and
that behind alveolus of incisor very conspicuous ; nasal bones

HYSTRIX 543

extending back to glenoid level, their median length about three
times that of frontals, the posterior margin of the two together
forming a broad, even curve ; dental formula : i ^ \ niii 1-, m ^"■'
= 20 ; tail short ; neck with conspicuous crest of long coarse
bristles.

Heniarks. — The genus Hystrix as here restricted contains the
African and Italian species only. Several forms have been
described, the status of which is inij^erfectly understood. One
occurs in Europe.

HYSTRIX CRISTATA Linujfus.

1758. [Hystrix'] cristata Linnaeus, Syst. Nat., i, 10th ed., p. 5G (near Rome,

Italy).
1792. H[ystrix] cristata europxa Kerr, Anim. Kiiigd., p. 213 (Renaming of

cristata) .
1839. ? [Hystrix cristata] \B,v. alba De Selys-Longchamps, Etudes de Micro-

mamni., p. 152 (nomeu nudum).
1910. Hystrix cristata Trouessart, Faune Mamm. d'Europe, p. 212.

Type local'dij. — Near Rome, Italy.*

Geographical distribution. — Northern Africa ; central and
southern Italy ; Sicily. (Wrongly attributed to the Iberian
Peninsula.) f

.Diagnosis. — General characters as in the genus Hystrix ;
size medium, the condylobasal length of skull about 130 mm.,
depth of skull at front of tooth row about 50 mm.

External characters. — Form heavy, the body appearing much
larger posteriorly than anteriorly on account of the mass of
specially elongated quills on posterior half of back ; head short,
the eye rather large, the ear low and inconspicuous ; legs short ;
feet broad ; tail shorter than hind foot, though bearing a dense
mass of quills which make it appear much longer. Head rather
short and deep ; ear low, rounded, overlaid by the bristle-like
hairs springing from region above and behind eye, a small but
evident lobe in front of meatus ; muzzle pad and median portion
of upper lip finely pubescent with fine and short but rather stift"
hairs ; nostrils situated in a deep groove crossing muzzle hori-
zontally ; upper lip with narrow median cleft. Front foot short
and broad, with four short, well developed digits and a rudi-
mentary thumb, the two median digits longest and bearing the
largest claws, the second and fifth successively shorter, all four
with strong slightly curved claws, the thumb a mere tubercle
covered on its upper surface by the rudimentary nail ; palm

* The statement "Habitat in Asia" is merely equivalent to saying
that the animal is foreign. The reference to the Systema [4th ed.], p. 9,
No. 1, leads directly to Ray (Syn. Meth. Auim. Quadr., p. 209), whose
account is based on a specimen from the mountainous districts near
Rome (see Thomas, Proc. Zool. Soc. London, 1911, p. 144).

t The peculiar distribution suggests the possibility that the animal's
presence in Italy is due to artificial introduction.

544 KODENTIA

naked, its surface almost entirely occupied by three tubercular
masses, separated by narrow furrows ; anterior pad at bases of
digits obscurely trilobed ; posterior pads roundish, the outer
larger than the inner. Hind foot much like fore foot, but
median claws shorter and hallux better developed than thumb,
its claw differing from those of other digits in its smaller size
only ; transverse groove separating trilobed tubercular mass at
base of digits from posterior mass less distinct than in front
foot ; posterior mass single, its anterior border showing a
tendency to bilobation. Tail very short, completely hidden
in the mass of quills which arise from it. Head, neck, shoulders,
limbs and underparts covered with coarse grooved bristles,
ranging in length from 20 to 45 mm., their width usually about
2 mm., though those on head are more slender and hair-like,
and those along median line from between eyes to shoulders
are greatly elongated, attaining a length of 300 mm. and pro-
ducing a conspicuous erectile crest or mane ; posterior half of
back and sides thickly beset with terete quills, ranging in length
from 35 to 300 mm., the longest on or near median line, the
extremities for the most part stiff and awl-like, but occasionally
drawn out into a hair-like filament ; base of tail with stiff terete
quills like those of body ; tip bearing a cluster of about a dozen
highly modified quills with slender thread-like bases (about
20 mm.) and a hollow terminal portion 20 to 30 mm. long, the
hollow part compressed at its base but becoming terete distally ;
whiskers stiff and long, extending to shoulders when laid back ;
throughout the body there is a sparse growth of coarse hairs
among the spines and quills, these hairs most numerous and
longest on posterior half of back, where they reach a length of
100 to 150 mm.

Colour. — Underparts, sides and legs blackish brown, the
bristles becoming horn-colour at base ; head, neck and shoulders
drab, many of the bristles in mane white or whitish ; whiskers
black ; quills on back, sides and tail blackish horn-colour, with
whitish terminal area 50 to 80 mm. long, and one to three
whitish annulations, each about 10 mm. in length, the general
effect of the quilled region black conspicuously spotted and
streaked with white.

ShtU. — In general appearance the skull differs strikingly
from that of other European rodents in the great depth and
width of the facial portion (post-lachrymal width about equal to
depth through same region, and fully one-half basilar length ;
occipital depth only about two-thirds that at middle of palate),
the unusual area occupied by the nasal bones (posterior margin
of nasals near glenoid level ; greatest combined breadth of
nasals, in lachrymal region, half or two-thirds their length, and
nearly equal to depth at front of tooth-rows), and the very
slender, weak anterior extremity of rostrum. Dorsal profile
evenly convex from front of nasals to about their posterior

HYSTIIIX 545

border, then sloping abruptly downward to the relatively low,
squarely truncate occipital region. Ventral profile not parallel
with dorsal profile, scarcely curved. Brain-case not differen-

FIG. 107.
Hystrix cristata.

tiated as an evident region, its width greater anteriorly than
posteriorly, the entire surface of each parietal occupied by a
deep depression for muscular attachment, the posterior margins

2 N

546

RODENTIA

of the depressions forming a distinct lambdoid crest, their inner
borders uniting to form a short sagittal ridge. Occipital region
almost squarely truncate, though sloping a little forward so that
the condyles are just visible when skull is viewed from above
(unless hidden by unusual development of lambdoid crest), its
outline forming a broad arch about four-fifths as high as wide,
slightly flattened at the sides and above, the lower outer extremities
of the arch formed by the prominent, triangular, slightly in-curved
paroccipital processes, the points of which descend nearly to level
of lower surface of condyles. Foramen magnum wider than
high. Floor of brain-case without special peculiarities ; a
slightly developed median longitudinal ridge ; auditoiy bulla
rather small, globular, the inflated portion longer than wide,
noticeably broader anteriorly than posteriorly, the meatus with
a short though evident tube ; anterior border in contact with
hamular ; posterior border free from paroccipital process.
Interorbital I'egion, the broadest portion of skull, highly arched
laterally, without evident lidges or depressions. Zygoma short
and heavy, not widely spreading, its anterior portion deep, the
posterior portion shallow, the lower border nowhere above
alveolar level thoughsloping evidently upward behind ; anteorbital
foramen considerably more than half as large as the rather con-
tracted orbit, its lower wall appearing as a conspicuous anterior
prolongation of zygoma. Nasals very large, their widest region
slightly behind level of posterior termination of nasal branches
of premaxillaries, their extension behind this region at least
equal to their anterior extension in front of anteorbital foramina,
the posterior outline of the two bones together evenly rounded,
their outer borders gradually converging anteriorly to a width a
little more than half greatest width ; anterior border of each
nasal strongly oblique with a slight but evident emargination at
outer edge. Nares somewhat naiTowly cordate in outline^ the
apex projecting as a flat shelf or shallow trough over space
between shafts of incisors. Facial portion of maxillary
terminating posteriorly in line with posterior termination of
premaxillary and with middle of lachrymal. Palate narrow,
without special features, its width scarcely exceeding alveolar
width of cheek-teeth, the portion formed by palatine bones
relatively small ; incisive foramina small (length about 6 mm.),
narrow, slightly divergent posteriorly, nearer to incisors than to
cheek-teeth, their outer margins continuous with ridges which
pass backward to anterior borders of alveoli of premolars and
unite anteriorly to form a noticeable crest between foramina and
incisors ; septum dividing foramina continued backward as a
high median ridge flattening out rather abruptly at level of front
of premolar. Mesopterygoid space considerably more than half
as wide as long, narrowing rapidly forward from a little behind
m^, the evenly concave anterior border at level of front of *»•' ;
hamular short, obliquely expanded, the posterior, longer limb of

547

the expansion bi'oadly in contact with inner portion of anterior
border of bulla. INIandible relatively slender, the posterior
portion not conspicuously deeper than ramus, including cheek-
tooth, the anterior portion a mere shell enclosing the robust
incisor ; articular process broad and low ; coronoid process small,
not rising conspicuously above level of posterior alveolar border,
widely separated from articular process ; angular y:)ortion broad,
lying cons23icuously on outer surface of alveolus of incisor, its
posterior border slightly concave. Dental foramen at alveolar
level.

Teeth. — Incisors robust, without special peculiarities of form,
the shafts scarcely compressed, but with antero-posterior diameter
perceptibly greater than lateral diameter ; enamel surface yellow.
Upjjer incisor with root extending to immediately in front of
alveolus of premolar, where its position is indicated ))y a sliglit
elevation on the surface of the bone ;
anterior face of tooth slightly oblicjue,
nearly flat though a little rounded oft'
at edges ; posterior surface narrow^
rounded, so that the outline of cross-
section is nearly an isosceles triangle
with all the angles rounded. Lower
incisor much longer and less curved
than upper, its root extending almost
to base of articular surface but not
prodvicing any very noticeable swell-
ing on outer side of mandible, and
scarcely rising above level of molar
crowns ; anterior face of tooth very
oblique, scarcely marked off from pos-
terior face ; inner face flat. Cheek-
teeth flat crowned, rather large, not
showing any very noticeable contrasts
of size or structure among themselves. Roots very imperfectly
developed, smallest in the upper teeth where they appear as
mere irregular tubercles, three of which can usually be distin-
guished, and sometimes a fourth ; shafts higher than wide,
slightly tapering downward. Crowns of upper cheek-teeth
broadly elliptical in outline, distinctly longer than wide. Their
enamel pattern shows individual irregularities, but its scheme
is as follows : outer side of crown with three re-entrant folds,
the anterior and posterior of which soon become isolated as
narrow islands, the middle fold more persistent and often con-
fluent with re-entrant fold of inner side ; first and second folds
siuiple ; third curving backward and outward, its extremity
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2 N 2

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HYSTRIX 549

molars but showing more tendency to irregularities. Lower
teeth with the same pattern reversed, the outer side with one
simple, oblique, backward sloping fold, the inner side with three
folds, the posterior and middle of which are usually simple, the
anterior complex and tending eventually to become divided into
two or three irregularly shaped islands arranged somewhat in a
spiral. The premolar shows the same tendency as the corres-
ponding upper tooth to become irregular in details of enamel
pattern, though the fundamental plan of its folds appears to be
always the same as in the molars.

Measurements. — Young adult (probably female) from Vizziri.
Sicily (approximate) : head and body, 600 ; tail, 40 ; hind foot,
75. Immature individual from Province of Rome, Italy (approxi-
mate) : head and body, 600 ; tail, 50 ; hind foot, 85. For cranial
measurements see Table opposite.

Siiecimens examined. — Seven, from the following localities: —
Italy : Near Rome, 4 (B.M. and U.S.N.M.) ; Vizzini, Sicily, 2 ; uo
exact locality, 1 (Stockholm).

3. Rome, Italy. Genoa Museum (e). 97. 2. 22. 1-3.

2,juv. Vizzini, Sicily. J. I. S. Whitaker (p). 3.4.21.1-2.

Family MUSCARDINID^.

1821. MyoxidiB (misprinted Mrjosidx) Gray, London Med. Repos., xv,

p. 308. April 1, 1821.
1857. Myoxina Blasius, Saugethiere Deutschlands, p. 285.

1898. Gliridx Lydekker, Zool. Rec. xxxiv. (1897), Mamm. p. 27, 1898.
Not of Ugilby, 1837.

1899. Muscardinidx, Palmer, Science, N.S., x, p. 413, September 22, 1899.

Geographical distrihution. — Warmer portions of the Old World
from Japan to England and from central Sweden south through
Africa. Absent from Madagascar, the Malay Archipelego and
Australia.

Characters. — In general as in the Muridse (p. 591) but
c;ecum absent, jugal bone larger and forming a more important
mechanical part of zygoma (in some genera approaching the
condition characteriatic of the Sciuridse), angular process of
mandible bent outward at middle so that its lower border
bears a noticeable secondary angle, and cheek-teeth always
rooted, brachyodont, their crowns with transverse cross ridges
variously arranged and sometimes obsolete, but referable to a
primitive 3-ridged pattern ; one premolar typically j^resent in
each jaw.

Bemarks. — Six genera of Muscardinidse are currently recog-
nized, four of which occur in Europe. They are mostly arboreal
animals with habits and aspect somewhat intermediate between
mice and squirrels, but readily distinguishable from the Muridne

550 ~ RODENTIA

by the presence of premolars, the cross-ridged pattern of the
enamel folding of the cheek-teeth and the peculiar secondary
angulation of the angular process of the lower jaw.

KEY TO THE EUROPEAN GENERA OF MUSCARDINID.i:.

Outer margin of first and second upper molar with
two high cusps (occasionally one or two others
low and obsolete).
Cro\vns of cheek-teeth deeply concave ; premolar
both above and below noticeably cuspidate ;
tail not uniformly haired, the basal half
terete, the terminal portion slightly flattened Eliomys, p. 550.
Crowns of cheek-teeth slightly concave ; premolar
both above and below not noticeably cuspidate ;
tail uniformly haired, moderately distichous... Dyromys, p. 56G.
Outer margin of crown of first and second upper
molar with five or six low cusps.
Crowns of cheek-teeth very slightly concave ; first
and second upper molars essentially alike in
form and in arrangement of cross-ridges ; tail

conspicuously distichous Glis, p. 572.

Crowns of cheek-teeth flat ; first and second upper
molars conspicuously unlike in form and in
arrangement of cross-ridges ; tail not evidently
distichous Muscarclinus, p. 583.

Genus ELIOMYS Wagner.

1843. Eliomys Wagner, Abhandl. k. Bayer, Akad. Wissensch. Miinchen,
Math.-Phys. CI., iii, p. 176 (Sub-genus of Myoxus).

1857. Eliomys Blasius, Saugethiere Deutschlands, p. 289 (Sub-genus of
Myoxus) part.

1890. Eliomys Reuvens, Die Myoxidae oder Schlaefer, p. 22 (Sub-genus of
Myoxus) part.

Tjipe species. — -Eliomys melanurus Wagner.

Geographical disfrihution. — Europe from the Atlantic coast
eastward into Asia Minor, and from northern Germany (Mecklen-
burg) south into northern Africa.

Characters. — Skull deep, smooth and rounded, without evident
ridges or depressions in interorbital region ; brain-case deep, the
auditory bullte large : ectopterygoid well developed ; jugal short
anteriorl3% not approaching lachrymal ; mandible with angular
portion fenestrate ; dental formula : / J]J, pm ^ J, m -j^-j = 20 ;
crowns of cheek-teeth deeply concave laterally, the inner and
outer margins noticeably elevated, the outer margin of each
maxillary tooth with two high cusps (one or two others low and
inconspicuous sometimes present) ; crown of rii^ and m- with live
transverse ridges, of which the third is incomplete ; tail not
uniformly haired throughout, the basal half terete, the terminal
portion slightly flattened.

Remarks. — The genus Eliomijs as now understood contains

ELIOMYS 551

about a dozen forms peculiar to the Mediterranean and neigh-
bouring regions. Five of them occur in Europe.

KEY TO THE EUROPEAN FORMS OF ELIOMYS.

Size large, hiud foot often more than 30 mm.,
condj'lobasal length of skull iu adults 36 to

39 mm. (Southern Spain and Portugal) E. luaitanicus, p. 560.

Size medium, hind foot never more than 30 mm.,

condj'lobasal length of skull in adults less than

36 mm.

Underside of tail normally * white from base

to tip.

Ear from meatus iu adults 23 mm. or more

(Central and south-western Europe) E. qncrci)ius, p. 551.

Ear from meatus in adults 20 to 22 mm.

(Balearic Islands) E. gymnesicus, p. 558.

Underside of tail normally f with white inter-
rupted by a black sub-terminal ring.
Condylobasal length of adult skulls 33 to
35 '6 mm.; brain-case slightly flattened,
the ratio of depth through bulla to
condylobasal length about 37 '5 (Sicily and

southern Italy) E. '})allidas, p. 559.

Condylobasal length of adult skulls 30 to
32 mm. ; brain-case deep, the ratio of depth
through bulla to condylobasal length
about 42 (Corsica and Sardinia) E. sarclus, p. 560.

ELIOMYS QUERCINUS Linnieus.

1766. [Mas'] quercinus Linnaeus, Syst. Nat., i, 12th ed., p. 84. Germany.
1778. Mus nitedida Pallas, Nov. Sp. Quadr. Glir. Ord., p. 88 (part ;

synonymy, not description).
1782. Myoxus nitela Schreber, Saugthiere, pi. ccxxvi (text, iv, p. 833,

1787). Germany.
1857. Myoxus qiiercinus Blasius, Siiugethiere Deutschlands, p. 289.
1890. Elioniys quercimis Reuvens, Die Myoxidae oder Schlaefer, p. 26.
1904. Eliomys liortualis Cabrera, Bol. Real Soc. Espafi. Hist. Nat., iv,

p. 183, April, 1904 (Valencia, Spain). Type in ^Madrid Museum.

1907. Eliomys Jiamiltonii Cabrera, Bol. Real Soc. Espan. Hist. Nat. vii,

p. 225 (June-July), issued October, 1907. (El Pardo, near Madrid,
Sx:)ain.) Type in Cabrera collection.

1908. Eliomys quercinus Cabrera, Ann. and i\Iag. Nat. Hist. 8th ser., i,

p. 192, February, 1908 {horttialis and hamiltonii united with
qtiercinus).
1910. Eliomys qtiercimcs Trouessart, Faune Manim. d'Europe, p. 136.

Type locality. — Germany.

Geographical distribution. — Central Europe from northern
Germany (Mecklenburg) to central Spain (Valencia) and northern
Italy ; west to the Atlantic coast.

Diagnosis. — Size medium (condylobasal length of largest
skulls i-arely attaining 33 mm. ; hind foot usually 25 to 27 mm.) ;

* Three partial exceptions among 75 specimens.
t Two exceptions among 34 specimens.

552 RODENTIA

ear large, its height from meatus usually about 23 to 25 mm. ;
under side of tail white throughout.

External characters. — General form slender, the head rather
large, not much elongated, the tail about as long as body without
head. Muzzle bluntly conical ; the nostrils near together,
separated by a deep groove which continues downward to lip ;
eye rather large and prominent ; ear large, extending nearly
half-way from eye to muzzle when laid forward, its general
outline a simple rather narrow oval, its inner and outer surfaces
covered with a fine velvety pubescence ; a low inconspicuous ridge
extending inward above meatus, its outer extremity not forming
angle on margin of conch ; front foot with four digits, the two
median equal and longest, the inner and outer sub-equal and
extending about to second phalanx of median ; palm naked, the
palmar tubercles live, three of which are sub-equal and closely
crowded at bases of phalanges, and two larger at internal and
postero-external edge of palm, the internal tubercle with pro-
jecting semi-detached extremity representing last vestige of
thumb ; hind foot with five digits, the thumb abruptly shorter
than the others and extending barely to end of first phalanx of
second digit ; fourth longest, third and second successively a little
shorter, fifth extending to middle of second phalanx of fourth ;
sole rather narrow, naked except at heel, the six tubercles well
developed but less crowded than those on palm, the postero-inner
much elongated and conspicuously difterent from others in form,
their surface smooth, the region between them noticeably
papillose ; claws well developed, small, those of fore foot slightly
exceeded in size by those of hind foot, except by that of thumb,
which is decidedly smaller than the others ; tail rather thick, the
skin readily breaking and slipping off" from the vertebra?, after
which the stump partly regenerates and puts forth a dense brush
of long hairs ; * hairs of tail completely concealing the annulations,
their length about 5 mm. at median terete portion of tail,
increasing to about 20 mm. at slightly flattened terminal portion.
Mammae : pi -I ; al — 1; i2 — 2 = 8.

Colour. — Upper parts light grey, the back, middle of neck and
whole of crown strongly tinged with russet, the sides clearer
grey. In some specimens the grey predominates throughout the
dorsal region, while in others the russet is in excess ; sides
ranging from ecru-drab to smoke-grey, usually forming a slight
but evident contrast with back. Underparts and inner surface
of legs dull creamy or buffy white, irregularly darkened by the
slate-colour bases of the hairs. The white covers cheek almost
to lower eyelid. Face with conspicuous black area beginning at
roots of whiskers, surrounding eye and extending to beneath and
slightly behind ear, the posterior portion becoming decidedly
broader ; a small but noticeable white spot immediately in front

* Bee Thomas, Proc. Zool. Soc. London, 1905, ii, p. 491.

ELIOMYS 553

of meatus, this succeeded by a largei- black spot just above base
of upper margin of conch. Ear dusky, sometimes incHning to
blackish along antei'ior border. Muzzle and sometimes face and
crown tinged with bufty clay-colour. Feet white, this colour
extending over entire wrist and most of forearm, the outer side
of upper arm with a longitudinal dusky, sometimes almost
blackish streak extending upward to fade into general body colour.
Tail sharply bicolor, the under surface white throughout, the
upper surface like back through basal half, then darkening
abruptly to clear black, this giving way in the pencil to white.
Young with back and sides an obscure slaty drab, the russet
gradually appearing on back, though often not until after the
general colour has assumed the light smoky grey of the adult
pelage.

Skull. — The general form of the skull is deep and rather

t^^^

Fig. 109.

Eliomys qiiercinus. Nat. size.

(The incisive foramina are too short.)

broad posteriorly, tapering noticeably to the slender rostrum.
Dorsal profile nearly flat from front of nasals to posterior
extremity of frontal or a little beyond, then uniformly and
rather strongly con\ex to interparietal across which it is again
flat ; ventral profile parallel with dorsal to front of tooth-row
then bent down to lowest region of bulla (which lies decidedly
below line of pterygoids), behind which it rises abruptly ; brain-
case squarish in outline when viewed from above, somewhat
wider and deeper anteriorly than posteriorly, its surface smooth
and without ridges other than a slight angularity along anterior

554

RODKNTIA

portion of suture between squamosal and parietal in old
individuals, and occasionally a low, backward-projecting lambdal
ridge ; parietals abruptly bent downward at sides, giving the
upper surface of brain-case a peculiar narrowed appearance ;
interparietal somewhat variable in form, but usually ligulate, with
pointed, rounded or squarish extremities and slightly projecting
median anterior angle, the anteru-posterior diameter from one-
third to one-half transverse diameter ; auditory bulhe t'ormiiig a
very conspicuous feature of brain-case when viewed from Ijehind,
their area decidedly greater than that of occipital, the depth
through bulla about equal to distance between outer borders of
paroccipital processes ; foramen magnum rather large, broader
than high, its inferior lip about on level with tips of paroccipital
processes ; floor of brain-case without special features, the suture
between basioccipital and basisphenoid remaining open until late
in life ; auditory V)ulla? large, smoothly inflated, the area of each
bulla somewhat greater than that of space enclosed between
zygoma and side of skull, the anterior border extending forward
to level of middle of glenoid fossa, the posterior border solidly
moulded against paroccipital process and extending upward along
side of exoccipital considerably above level of zygoma, so that
this portion of bulla is plainly visible when skull is viewed from
above ; meatus large, not tubular, but with conspicuous, forward-
curved lip in region nearest zygoma ; interorbital region slightly
hour-glass shaped, the upper surface flat or a little convex
laterally, the edges behind middle becoming somewhat angular in
old individuals ; zygomata rather weak, compressed, a little
expanded at middle, gradually sj^reading anteriorly, the gi'eatest
breadth nearly at glenoid level, the lower border essentially
straight though sometimes a little concave ; anteorbital foramen
rather large, considerably higher than wide, its greatest width
above middle ; rostrum almost parallel-sided when viewed from the
side, slightly tapering when viewed from above ; nasals a little
narrower posteriorly than anteriorly, the posterior border sti'aight
or emarginate, rarely pointed, usually about on level with
posterior termination of nasal branches of premaxillaries and
with middle of lachrymal ; incisive foramina large, narrow and
parallel-sided anteriorly, becoming abruptly much wider through
posterior half or two-thirds, the greatest width of both together
slightly more than half length, the length fully equal to one-half
diastema ; * a noticeable curved ridge extending from posterior
border of each incisive foi'amen to front of alveolus of corre-
sponding premolar ; palate wider anteriorly than posteriorly,
the small portion formed by the palatine bones conspicuously
fenestrate ; mesopterygoid space nearly parallel-sided, slightly
wider posteriorly than anteriorly, the least width about one-third
length, the anterior border rounded, with or without a small

* Not long euough iu fig. 109.

ELIOMYS 555

median spine, the slightly curved hamular in contact with audi-
tory bulla at tip. External pterygoid plate evident though small,
foruiing a distinct pterygoid fossa. Mandible slender, the ramus
curved in front of tooth-row, coronoid process long, sloping
gradually backward, its point considerably above level of
condyle ; angular process well developed, short and deep, the
middle of lower border with abrupt angle, above which the bone
is perforated by a conspicuous round vacuity about 1 • 5 mm. in
diameter.

Teeth. — The teeth are moderately large relatively to size of
skull, the length of tooth-row contained about 2i times in palatal
length ; tooth-rows wider apart anteriorly than posteriorly, the
alveolar length equal to distance between anterior roots of
prejnolars. Incisors with no special peculiarities, the anterior

Fig. 110.
Eliumijs quercinus. Cheek-teeth. X lu.

sui'face smooth, light yellow, its diameter slightly less than
antero-posterior diameter, the discrepancy slightly greater in the
lower than in the upper teeth. Root of lower incisor extending
to that of m.^ ; dental foramen at level of alveolar line. Upper
cheek-teeth with crowns deeply concave, the inner and outer
borders noticeably elevated. Molars sub-equal, the second
slightly the lai'gest, the crowns squarish, slightly wider than
long, the inner border a little rounded, the posterior border of
m^ shortened ; inner border of crown with a robust, moderately
high cusp, the most elevated portion of which is in front of
middle, its base occupying nearly entire inner border, but in
some specimens succeeded by a minute posterointernal cusp
early disappearing with wear ; outer border with two moderately
large subterete cusps, the second not so large as first (the
discrepancy greater in «t^ than in the other teeth) and two or

556

RODENTIA

three minute secondary cusps, one in front of tirst main cusp, the
others behind second ; surface of crown crossed by four complete
transverse ridges, the first, second and third extending from large
inner cusp respectively to small anterior cusp, to large anterior
cusp and to large posterior cusp, the fourth from small postei'o-
internal to small postero-external ; on outer half of crown
between the second and third complete cross-ridges occur two
incomplete ridges extending inward for a varying distance
from posterior base of anterior large cusp and anterior base of
posterior large cusp, the outer extremity of one or the other
sometimes forming a minute supplemental cusp ; in m^ the third
complete ridge is relatively smaller and second incomplete ridge
relatively larger than in the other teeth, the first incomplete
ridge obsolete or absent. Upper premolar essentially like
molars in arrangement of main cusps and ridges, but crown
slightly smaller than that of m^, and somewhat distorted by the
more central, anterior position of antero-external root, and ridges
tending to be incomplete, especially the two anterior ; secondary
ridges and cusps essentially absent ; milk tooth much smaller
than permanent premolar, its nearly triangular crown with area
barely half that of m^, its ridges less regular. The milk tooth
remains in place until m^ is fully grown. Lower cheek-teeth
much like upper, but with crowns slightly less concave than
those of maxillary teeth, and tending, especially that of r)/3, to
become basin-shaped, their cusps and ridges less sharply defined,
those of 7H3 often very irregular ; outer side of crown with three
low but well defined cusps, inner side irregularly crenulate in m.^
and m^, with two rather low but well defined cusps in m^, the
posterior of which is decidedly the larger ; main cross ridges
four, the first and second tending to be distorted and incomplete ;
an incomplete intermediate ridge on inner side of crown between
second and third main ridges. Lower premolar triangular in
outline, its apex in front, its crown area about four-fifths that of
m^ ; a well developed cusp at each corner and an indistinct
transverse ridge across middle ; milk tooth smaller than
permanent tooth, but discrepancy in size less than in the
corresponding upper teeth ; crown of milk premolar essentially
like that of succeeding tooth, but with a small cusp at middle of
antero-external border, and transverse ridge absent.

Measurements — Two adult females from Maredsous, Namur,
Belgium : head and body, 136 and 137 ; tail, 116 and 121 ; hind
foot, 28 and 29 ; ear from meatus, 24 and 24. Adult male from
I'Hospitalet, Ariege, France: head and body, 116; tail, 97"6;
hind foot, 25*4 ; ear from meatus, 25. Two adult females from
the same locality: head and body, 118 and 127; tail, 108
and 111 ; hind foot, 26 and 26'4; ear from meatus, 24 and 25.
Five adult males from Valescure, Var, France : head and body,
125(115-131); tail, 107 (100-111); hind foot, 26-3 (25-27);
ear from meatus, 23 "6 (22-25). Five adult females from Silos,

Burgos, Spain: head and body, 120 (115-130); tai], 107
(100-112); hind foot, 26 (25 -4-26 -6); ear from meatus, 23-4
(22 "6-24). Two males from Dehesa de Valencia, Valencia,
Spain: head and body, 120 and 125; tail, 100 and 107 ; hind
foot, 26 and 26 ; ear from meatus, 24 and 24. For cranial
measurements see Table, p. 562.

Specimens examined. — One hundred and twenty, from the following
localities : —

Belgium : Maredsous, Namur, 2.

France : Pas-de-Calais, 2 ; I'Hospitalet, Ariege, 7 ; Port^, Pyrenees-
Orientales, 2 ; Nimes, Gard, 14 (B.M. and Mottaz) ; Valescure, Var, 8 ;
Chamonix, Haute-Savoie 3 (U.S.N.M.) ; Mounetier, Haute-Savoie, 1
(Mottaz) ; no exact locality, 2.

Spain : Cabanas, Corufia, 2 ; Jaca, Huesca, 4 ; Panticosa, Huesca, 1 ;
Lerida, 1 ; Pajares, Leon, 7 ; Silos, Burgos, 11 ; Castrillo de la Reina,
Burgos, 1 ; La Granja, Segovia, 4 ; Villalba, Madrid, 2 ; Rascafria, Madrid, 1 ;
Barracas, Castellon, 5; Dehesa de Valencia, 3.

Germany : Schmilka, Saxony, 1 ; Rudolstadt, Thiiringen, 2 ; Marxheim,
Bavaria, 1 ; South Germany, 2.

Austria-Hungary : Spalato, Dahnatia, 1 ; Mittelberg, Vorarlberg, 2.

Switzerland : Near the Dole, Vaud, 3 (Mottaz) ; St. Cergues, Vaud, 2
(B.M. and U.S.N.M.) ; Les Plans, Vaud, 3 (U.S.N.M.) ; Miirren, Bern, 1 ;
St. Gahen, 1 (Mottaz); Untervatz, Grisons, 9 (U.S.N.M.); Vulpera-
Tarasp, Grisons, 2 (Rothschild) ; Fusio, Ticino, 1 (U.S.N.M.) ; no exact
locality, 1.

Italy: Padola, Cadore. 1 (Turin) ; Mondovi, Cuneo, 4 (Genoa).

2 9. Maredsous, Namur,

Belgium.
2 9. Boulogne, Pas-de-Calais,

Franco.
4 ?. L'Hospitalet, Ariege.
2 c?, 9. L'Hospitalet, Ariege.
(A. Robert.)
9. Port6, Pyr6n6es-Orien-

tales. {A. Robert.)
9. Port6, Pyr6n6es-Orien-
tales.
2(5,4 9. Nimes, Gard. {C. Mot-
taz.)
5 (5, 2 9. Valescure, Var.
9. Valescure, Var.

1. Prance.

6. France.
9 al. Cabanas, Coruiia, Spain.

1. Cabanas, Coruiia.

2 6. Jaca, Huesca. [N. Gon-
zalez.)
6. Panticosa, Huesca.

{N. Gonzalez.)
6. L6rida. (N. Gonzalez.)
2 9. Pajares, Leon. (N. Gon-
zcitcz \
2 c5, 3 9. Pajdres, Leon. [N. Gon-
zalez.)

Rev. G. Pournier

(c & p).
C. G.Danford(c&p).

G. S. Miller (c).
0. Thomas (p).

0. Thomas (p).

G. S. Miller (c).

0. Thomas (p).

G. S. Miller (c).

Hon. N. C. Roths-
child (c & p).

Dr. J. E. Gray (p).

Tomes Collection.

Prof. V. L. Seoane
(c & p).

Prof. V. L. Seoane
(c & p).

0. Thomas (p).

O. Thomas (p).

O. Thomas (p).
0. Thomas (p).

0. Thomas (p).

3. 3. 30. 1-2.

89. 10. 6. 2.
10. 7. 17. 1.
8. 8. 4. 172-175.
8. 9. 1. 72-74.

8. 9. 1. 75.

8. 8. 4. 171.

8. 8. 10. 57-62.

8. 8. 4. 176-182.
10. 0. 13. 1.

43. 12. 29. 17.

7. 1. 1. 76.
94. 1. 1. 9.

94. 3. 12. 2.

8. 2. 9. 75-76.

8. 2. 9. 77.

8. 2. 9. 80.
8. 2. 9. 78-79.

8. 2. 9. 81-85.

558

8 9.

Silos, Burgos.

G. S. Miller (c).

8. 8. 4. 57-64.

9.

Castrillo, Burgos.

G. S. Miller (c).

8. 8. 4. 65.

2 9.

Villalba, Madrid.
{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 70-71.

S, 3 al.

La Granja, Segovia.

M. de la Escalera (c).

6. 11. 4. 5-8.

3 6.

Barracas, Castellon.

(N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 72-74.

3 6.

Dehesa de Valencia.
{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. G7-G9.

S,9.

Rudolstadt, Thiiriiigen,
Germany. (W. ScJiiiltei

Lord Lilford (p).
■•)

0. 5. 5. 1-2.

6.

Marxheim, Bavaria.

(Wolter.^toif.)

Lord Lilford (p).

0. 5. 5. 3.

2.

S. Germany.

Dr. A. Giinther (c).

59. 9. 6. 25-26.

1.

Spalato, Dalmatia.

Lord Lilford (p).

95. 4. 6. 1.

2 9.

Mittelberg, Austria.
(Dr. F. Major.)

0. Thomas (p).

8. 11. 30. 5-6.

9.

St. Gergues, Vaud, Swit-
zerland.

G. Mottaz (p).

6. 2. 6. 5.

1.

Miirren, Bern.

W. Giirtner (p).

92. 10. 5. G.

1.

Switzerland.

E. E. Alston (p).

79. 9. 12. 3.

ELIOMYS GYMNESICUS Thomas.

1901. Eliomys qucrcinus Thomas, Proc. Zool. Soc, London, p. 41.

1903. Eliomys gymnesicus Thomas, Ann. and Mag. Nat. Hist., 7th ser., xi,

p. 494, May, 1903. Type in British Museum.
1910. Eliomys quercimis gymnesicus Trouessart, Faune Mamm. d'Europe,

p. 137.

Type locality. — San Cristobal, Minorca, Balearic Islands.

Geographical distribution. — Balearic Islands.

Diagnosis. — Similar to Eliomys quercinus but with smaller
ear (height from meatus 20 to 22 mm. instead of 23 to 25 mm.).

Colour. — The colour is in all respects like that of Eliomys
quercinus, though the upper parts appear to be a little more
suffused with reddish brown than usual in the mainland animal.
Tail markings exactly as in E. quercinus.

Measurements. — Type (adult male) : head and body, 131 ;
tail, 107 ; hind foot, 26 ; ear from meatus, 21. Two other adult
males from the type locality: head and body, 116 and 120;
tail, 100 and 102; hind foot, 27 and 25*5; ear from meatus,
22 and 20. For cranial measurements see Table, p. 564.

Sxieclmens examined. — Six, all from the Island of Minorca.

Remarks. — The Balearic Eliomys appears to be well differen-
tiated from the mainland animal by its smaller ear.

5 6, 6 San Gristobal, Minorca- 0. Thomas and R. I. 0. 7. 1. 44-49.
skull. Pocock (c & p).

(0. 7. 1. 47. Type of species.)

559

ELIOMYS PALLIDUS Barrett-Hamilton.

1899. Eliomys pallidus Barrett-Hamilton, Arm. and Mag. Nat. Hist.,
7th ser., in, p. 226, March, 1899. Type in British Museum.

1901. Eliomys cincticauda Miller, Proc. Biol. Soc, Washington, xiv, p. 39,
April 25, 1901. (Sorrento Italy). Type in U.S. National Museum.

1910. Eliomys cincticauda and E. pallidus Trouessart, iFaune Mamm.
d'Europe, pp. 137, 138.

Type locality. — Palermo, Sicily.

Geograpliical distrlhittion. — Sicily and southern Italy.

Diagnosis. — Size greater than in Eliomys qucrcinus (condylo-
basal length of largest skulls usually 33 to 35 mm. ; hind foot
usually 27 to 29 mm.) ; general colour not so dark as in the
more northern animal ; tail with sub-terminal black area forming
a complete ring ; skull with posterior termination of nasals
usually pointed.

Measurememts. — Average and extremes of ten adult males
from Sicily : head and body, 143 (130-150) ; tail, 118 (110-131) ;
hind foot, 28-8 (27-30) ; ear from meatus, 23-6 (22-25). Pom-
adult females from the same region: head and body, 142 (136-
150); tail, 111-4 (105-120); hind foot, 27-7 (27-29). Three
adult females from Calabria, Italy : head and body, 143 (140-
145); tail, 130-6 (125-135) ; hind foot, 28-6 (27-30) ; ear from
meatus, 22 - 6 (22-24). Por cranial measurements see Table,
p. 564.

Specimens examined. — Thirty-four, from the following localities : —
Italy: Tuscany (no exact locality), 1 (U.S.N.M.) ; Sorrento, 5; type

and paratypes of cincticauda (U.S.N.M.); Santa Eufemia d'Aspromonte,

Calabria, 9.

Sicily : Palermo, 8 ; Ficuzza, 7 ; San Giuglielmo, Castelbuono, 3 ;

Mondello, 2 ; Madonna del Alto, 4.

ItemarJi-s. — In general the South Italian Eliomys so closely
agrees with E. quereiims as to require no detailed description.
The larger size, readily appreciable in skull, teeth and hind foot,
is alone sufficient to distinguish it ; but in addition to this there
is the peculiar black-ringed tail, a character nearly always
diagnostic as compared with E. qiierciniis, though shared by the
much larger Iberian E. lusitanicus. In details of form the skull
shows no peculiarities other than the tendency of the nasals to
terminate posteriorly in a distinct median point. The type
specimen is an immature individual in the light transitional
pelage between the plumbeous first coat and the russet-tinged
livery of the adult.

3 6, 6 9. Aspromoute, Calabria, O. Thomas (r). 6. 8. 4. 2-5.

Italy. {A. liobcii.) 8. 9. 1. 33-35.

6, 9. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 6-7.

(98. 10. 6. 0. Tyix of species.)

9 juv. Palermo, Sicily. J. I. S. Whitaker (p). 11. 1. 1. 110.

5 6,19. Ficuzza, Sicily. O. Thomas (p). 6. 8. 4. 28-32.

(A. Robert.) 8. 9. 1. 32.

560 KODEXTIA

2 6. Mondello, 1000m. Sicily. 0. Thomas (p). 6. 8. i. 33-34,

(A. Robert.)
2 c5, 2 9. Madonna del Alto, Sicily. O. Thomas (p). 8. 9. 1. 28-31.

[A. Bobcrt.)

ELIOMYS SARDUS Barrett-Hamilton.

1901. Eliomys sardus Barrett-Hamilton, Ann. and ]\Iag. Nat. Hist.,

7th ser., vii, p. 340, April, 1901.
1910. Eliomys sardus Trouessart, Fauna Mamm. d'Europe, p. 13S.

Tyjie locality. — Tricoli, Cagliostra, Sardinia.

Geographical disirihntion. — Sardinia and Corsica.

Diagnosis. — Similar to Eliomys pallidus but with smaller skull
(condylobasal length 30 to 32 mm.) and relatively deeper brain-case.

Colour. — The colour does not differ appreciably from that of
Eliomys pallidus.

Skull and teeth. — The skull does not attain the large size of
that of E. jKdlidus. In form it is peculiar in the relatively
greater depth of brain-case, the ratio of depth through bulla to
condylobasal length I'anging in four specimens from 41-f- to
43 + , while in seven of the larger animals it ranges from 37 to
38 + . Teeth as in the related form.

Measurements. — Type (old female): hind foot, 26 '6. Three
other Sardinian specimens: hind foot, 25*4, 26 and 28. For
cranial measurements see Table, p. 565.

Specimens examined. — Nine, from the following localities : —
Sardinia : Tricoli, 2 ; Lanusei, 2.
CoESiCA : Asco, 5.

6 juv., 9. Tricoli, Sardinia. O. Thomas (p). 1. 3. 8. 1-2.

[G. Meloni.) (1. 3. 8. 1. Ty2ye of species.)

S, 9. Lanusei, Sardinia. O. Tliomas (p). 1. 3. 8. 3-4.
(G. Meloni.)

2 6, 2 9. Asco, Corsica. {Dr. 0. Thomas (p). 8. 11. .30. 1-4.
F. Major.)

ELIOMYS LUSITANICUS Reuvens.

1890. El[io7nys'] nitela var. lusitanica Reuvens, Die Myoxidae oder

Schlaefer, p. 28, footnote (Lisbon, Portugal).
1897. Mioxus nitela var. amori Graels, Mem. Real Acad. Sci., Madrid,

XVII, p. 481 (Cordova, Spain).
1901. E[liomys] amori Thomas, Proc. Zool. Soc, London, p. 41.
1904. Eliomys amori Cabrera, Bol. Real Soc. Espafi. Hist. Nat., xvii,

p. 187, April, 1904.
1908. Eliomys lusitanicus Cabrera, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 189, February, 1908.
1910. Eliomys qucrcinus lusitanicus Trouessart, Fauue Mamm. d'Europe,

p. 137.

Tyjie locality. — Lisbon, Portugal.

Geographical distribution. — Southern portion of the Iberian
Peninsula ; exact limits of range not known.

Diagnosis. — Largest European member of the genus Eliomys
(hind foot in adults usually more than 30 mm.) ; skull con-

ELIOMYS 561

spicuously larger and heavier than iji E. querciniis, the conclyio-
basal length of skull, 36-39 mm. ; colour slightly darker and
more brighty russet than in E. quercinus, the tail usually with
black sub-terminal area forming a complete ring.

Colour. — Paratype (No. 46. 11. 21. 1): hairs of back with
three distinct colour bands: (1) slate-grey (8 mm.); (2) light
buffy grey (2 mm.) ; (3) russet (2 mm.), the extreme tips of hairs
whitish. Longer hairs black except for the slate-grey bases.
The general eftect is a peculiar dull reddish brown, faintly
" lined " by the longer black hairs and a ^ittle "powdered "by
the whitish tips to the ordinary hairs. On sides the bufFy grey
band becomes longer (4 mm.) and paler (a clearer, more whitish
grey), the russet at the same time gradually disappearing. Dark
markings of head as in E. quercinus. Tail as in E. quercinus,
except that the black ring usually present in E. lusitanicus is
indicated on lower surface by dark bases to the hairs in the
region where it should occur.

Specimens from Seville, Spain, resemble the paratype except
that the russet of the back is slightly more intense, a difference
that might readily be accounted for by their fresher condition,
the paratype having been in the Museum collection more than
fifty years longer than the others. In live of the Seville skins
the black ring on the tail is complete, the black area on under
side ranging from 40 mm. to 60 mm. in width. In one, however,
as well as in a specimen from Goto DoSana, Province of Huelva,
the tail is like that of the paratype, the ring being merely
indicated by dark bases to the hairs of the white under surface.

Skull and teeth. — Except for its noticeably larger size the
skull resembles that of Eliomys quercinus. As in the smaller
animal, the nasals are truncate or emarginate posteriorly, not
pointed as they usually are in the other ring-tailed species,
E. iiaUidus and E. sardus. Teeth as in E. quercinus, but appre-
ciably larger, the difference more noticeable to the eye than
might be supposed from the actual measurements.

Measurements. — Paratype (sex not known) from Lisbon,
Portugal : hind foot, 30. Adult male from Goto Doiaana, Huelva,
Spain : head and body, 166 ; tail, 120 ; hind foot, 31 ; ear from
meatus, 27. Three adult males fi'om Seville, Spain : hind foot,
31, 31 and 32. For cranial measurements see Table, p. 565.

Specimens examined. — Twelve, from the following localities :^

Portugal : Lisbon, 1.

Spain : Seville, 6 ; Goto Donana, Huelva, 3 ; Jerez, Cadiz, 2.

RemarJcs. — The South Spanish Eliomys is so readily distin-
guishable from the other ring-tailed species by its large size that
no special comparisons are required.

1. Lisbon, Portugal. C. Friend (c & p). 46. 11. 21. 1.

{Paratype of species.)
2 6, ?. Seville, Spain. {Dr. A. Lord Lilford (p). 95. 3. 3. 19-21.

Ruiz.)

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Genus DYROMYS Tlionias.

1780. Myoxus Zimmermann, Geogr. Gesch., ii, p. 351 (part).

1857. Eliomys Blasius, Siiugethiere Deutschlands, p. 288 (Sub-genus of

Myox7is) part.
1890. Myoxus Reuvens, Die Myoxidae oder Schlaefer, p. 24 (Sub-genus of

Myoxus) part.
190G. Dryomys Thomas, Proc. Zool. Soe., London, 1905. ii, p. 8-15,

April 5, 1906 (Sub-genus of Eliomys). Not of Philippi, An. Mus.

Nac. de Chile, xiv, p. 20, 1900.
1907. Dyromys Thomas, Ann. and j\Iag. Nat. Hist., 7th ser., xx, p. 406,

November, 1907 (Substitute for Dryomys).

Type. — Mus nitedula Pallas.

Geographical distribxtion. — From central Asia through Asia
Minor to Hungary and eastern Switzej-land.

Characters. — Teeth resembling those of Eliomys but relatively
smaller than in any of the other European members of the
family, the crowns with concavity less pronounced than in
Eliomys and cross-ridges better developed, the first intermediate
ridge in the upper molars nearly complete and scarcely lower
than the others ; three rudimentary ridges intercalated between
the main ridges of jh^ m^ and m.^ ; outer border of crowns of
upper molars with two high main cusps nearly as well developed
as in Eliomys, these usually supplemented by a minute anterior
or posterior cusp ; lower molars with outer cusps relatively lower
than in Eliomys and sub-equal in size, inner cusps essentially
as in the related genus ; premolars both above and below with
crowns sub-quadrate in outline, nearly flat, the cusps obsolete,
the maxillary tooth crossed by four or five ridges, the anterior
of which are better developed than in Eliomys. Skull essentially
as in Eliomys, but parietals as broad as in Muscardinus ; the
angular process of mandible fenesti^ate. Externally diff"ering
from Eliomys in the uniformly haired, distichous tail.

lie marks. — Though nearly related to Eliomys this group
seems worthy of recognition as a genus. Seven forms are known,
four of which (three of them apparently local races of one species)
occur in Europe.

KEY TO THE EUROPEAN FORMS OF DYROMYS.

Skull broad and robust, the zygomatic
breadth about 17 mm. ; auditory bullae
decidedly enlarged (Rustschuk, Bul-
garia) D. robustus, p. 572.

Skull slender, the zygomatic breadth about
15 mm. ; auditory bullae not specially
enlarged D. nitedula, p. 567.

567

Teeth weak ; ccowii area of upper pre-
molar decidedly more than half that

of first molar (Greece) D. nitedula iciivjei, p. 570.

Teeth robust ; crown area of upper pre-
molar scarcely half that of first
molar.
General colour of adults yellowish
brown above (Hungary and east-
ward) D. nitedula nitedula, p. 568.

General colour of adults greyish brown

above (Tirol and eastern Alps) D. niiedida intermedins,^. 569.

DYROMYS NITEDULA Pallas.
(Synonymy under subspecies.)

Geographical disirihiition. — South-eastern Europe from eastern
Switzerland and north-eastern Italy into Asia Minor ; north to
northern Hungarj-. Limits of range not known.

Diagnosis. — General characters as in the genus : tail about as
long as head and body ; face with dark line from muzzle to ear.
Skull slender, the rostrum rather long ; auditory bulla^ not
specially enlarged.

External characters. — Externally as in Elionu/s quercinus,
except that tail is uniformly rather long-haired throughout, its
form moderately distichous ; ear smaller, extending barely to
middle of eye when laid forward, the ridge on its inner surface
above meatus high and valve-like, terminating externally in a
conspicuous angular projection on mai'gin of conch ; relative
lengths of digits as in E. quercinus ; fore foot longer and
narrower, the terminal lobe of postero-internal tubercle larger
and more distinct ; hind foot with surface of sole betw^een
tubercles less papillose, scarcely more than reticulate, and
postero-internal tubei'cle not much longer than that at base of
thumb, though narrower. Mammse : p 2 — 2, i 2 — 2 = 8.

Colour. — Upper parts ranging from a light, clear greyish
brown to a distinctly yellowish brown with a russet tinge, the
face usually paler than back, the tail like rest of dorsal surface,
but lighter or more greyish, and sprinkled with wdiitish hairs at
edges. Underparts and feet whitish or very pale buff, this
colour usually suflusing under side of tail. A well defined
blackish line extends from base of whiskers to ear, including eye
(width on upper lid about 1*5 mm., on lower lid about '5 mm.),
and is often rendered more conspicuous by a light edging along
upper border.

Skull. — In general apj^earance the skull resembles that of
Eliomys quercimis, though in size and certain peculiarities of form
it more nearly approaches that of Muscardinus avellanarius. It
suggests that of Eliomys quercinus in form of brain-case (though
the parietals are relatively broader), of interorbital region, of
rostrum as viewed from above, and in the gradually spreading

568

RODENTIA

;^yg<)mata. The foi'm of the auditory bulla is also, as in Eliomys,
much more inflated anteriorly tlian posteriorly, and readily dis-
tinguishable from the rounder, more evenly inflated bulla of 6r//.s'
and MuscardlnuH.

Teeth. — The detailed structure of the teeth has already been
sufliciently described. As compared with those of Eliomi/s
quercinuH the molar rows are more nearly parallel, and there is

Fig. 111.
Dijromys nitedula. Nat. .size.

FIG. 112.
Dijromys nitedula. Cheek-teeth.

X 10.

usually less contrast in size both above and below between the
premolar and last molar with the two middle teeth.

Memarlxs. — Among the European Muscardinidx this species is
immediately recognizable by the combination of a dark face
streak and uniformly bushy tail. In size it is intermediate
between 3Inscardinus avellanarius and Eliomys quercinus, but
nearer the former. In different parts of its range Dyromys
nitedula has become modified into four local races, three of
which occur in Europe.

Dyromys nitedula nitedula Pallas.

1778. Mils nitedula Pallas, Nov. Sp. Quadr. Glir. Ord., p. 88, part, des-
cription, not synonymy (Region of the lower Volga, Russia).

1782. Mijoxus dnjas Schreber, Siiugthiere, pi. ccxxv b. ; description, iv,
p. 831, 1787 (Region of the lower Volga, Russia).

1857. Myoxus dryas Blasius, Siiugethiere Deutschlands, p. 295.

1906. Eliomys nitcdulus Thomas, Proc. Zool. Soc, London, 1905, ii, p. 348,
April 5, 1906.

1910. Dyromys nitedula Trouessart, Faune Mamm. d'Europe, p. 133.

Type locality. — Region of the lower Volga, Russia.

Geograplucal distribution. — South-eastern Europe north of
the Balkan Peninsula, west to Hungary. Exact limits of distri-
bution not known.

DYROMYS 569

Diagnosis. — Skull with rostrum railier long, the distance from
anterior root of zygoma to tip of nasal }nore than 8 mm. ; teeth
robust ; crown area of upper premolar scarcely half that of first
molar ; general colour of adults yellowish brown above.

Colour. — Adult : upper parts a yellowish brown intermediate
between the russet of Ridgway and yellowish wood-brown, the
back inconspicuously "lined" with black, the sides lightening
almost to ochraceous-buft" ; underparts pale cream-buff; tail hair-
brown above in noticeable contrast with back, the sprinkling of
whitish hairs along edge indistinct, underside of tail mixed
whitish and mouse-grey, much darker than belly ; feet whitish ;
muzzle and entire region between black face stripes as far as
eyes light grey, this colour continuing to ear as a border to black
stripe about 2 mm. wide. An obscurely defined buffy white spot
at anterior base of ear. Young similar to adult but duller, the
upper parts light wood-brown, sometimes with a greyish tinge.

Measurements. — Hind foot in an adult male from Belgrade,
Servia, 21 mm. Hind foot in three specimens from Zuberee,
northern Hungary, 20 "5^ 21 and 21 ram. For cranial measure-
ments see Table, jj. 571.

Specimens examined. — Twelve, from the following localities : —
Austria-Hungary : Zuberec, northern Hungary, 6 ; Herkulesbad, 3 ;
Hungary, no exact locality, 1.
Servia : Belgrade, 2.

4, 1 juv. Zuberec, Hungary. Budapest Museum (e). 94. 3. 1. 37-41.

1. Zuberec. G. Barrett-Hamilton 11. 1. 2. 59.

2 6,19. Herkulesbad. {F. Cox.) Hon. N. C. Rothschild 7. 9. IG. 12-14.

2 ?. Belgrade, Servia. Dr. R. B. Sharpe (p). 75. 8. 24. 3-4.

Dyromys nitedula intermedius Nehring.

1S02. Myoxus intermedius Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 155.
1910. Dyromys nitedula intermedius Trouessart, Faune jMamm. d'Europe,

p. 134.

Type locality. — Near Lienz, Tirol, Austria-Hungary.

Geographical distrUmtion. — Tirol and eastern portion of the
Alps (Engadine and Cadore).

Diagnosis. — Similar to Dyromys nitedula nitedula, but general
colour of upper parts greyish brown without tinge of yellow.

Colour. — Upper parts between the broccoli- brown and mouse-
grey of Ridgway, a little suffused with wood-brown across thighs
and rump ; tail slightly darker than back but not different in
colour, its edges ci'eamy white in strong contrast ; underparts
and feet buffy white, this colour suffiusing whole underside of
tail. Face markings as in true nitedula, but grey edging along
upper side of black lines less evident.

Measurements. — Two adult males from Vulpera-Tarasp,

570 RODENTIA

Grisons, Switzerland : head and body, 92 and 93 ; tail, 90 and
70 (imperfect); hind foot, 19 '5 and 20'5; ear from meatus,
14 '5. Adult from Padola, Cadore, Italy: head and body, 8 G ;
tail, 86; hind foot, 19.

Specimens examined. — Four, from the following localities : —
Switzerland : Vulpera-Tarasp, Grisons, 2 (Rothschild).
Italy : Padola, Cadore, 2 (B.M. and U.S.N.M.).

Bemarks. — At first sight this animal might be supposed to be
the young of Dyromys nitedula nitedula, but none of the three
specimens examined shows any indication of immaturity.

1 al. Cadore, Venetian Alps, Dr. E. Festa (r). 9. 1. 18. 1.
Italy.

Dyromys nitedula wingei Nehring.

1881. Eliomys dryas Winge, Vidensk. Meddel. fra den naturh. Foren. i
Kjobenhavn, 1881, p. 50 (near Athens). Not Mus dryas Pallas.

1902. Myoxus loingei Nehring, Sitz.-Ber. Gesellsch. Naturforsch. Freunde,
Berlin, p. 5 (Parnassus region, Greece).

1910. Dyromys nitedula ivingei Trouessart, Faune Mamm. d'Europe, p. 134.

Type locality. — Parnassus region, Greece.

Geographical distribution. — Known only from the type locality
and from the vicinity of Athens.

Diagnosis. — Similar to Dyromys nitedula nitedula but skull with
rostral portion less elongate, the distance from anterior root of
zygoma to tip of nasal less than 8 mm., and teeth weak ; upper pre-
molar with crown area decidedly more than half that of first molar.

Colour. — The colour appears to be indistinguishable from that
of D. nitedula nitedula.

Slcull. — The only skulls examined are broken. While the
bram-case does not appear to differ from that of true D. nitedula,
the rostrum is shorter and weaker than in the allied .species,
though not different in form. Auditory bullse apparently larger
than in D. n. nitedula, but the material is not sufficient to show
whether this character is constant.

Teeth. — While resembling in all essential characters those of
D. n. nitedula the molars are distinctly less robust, particularly
the first and second. As a result the premolar is larger relatively
to the first molar, and the tooth-row as a whole is more nearly
parallel-sided, and with less broadening at middle.

3Ieasurements. — Two adult females from Tatoi, near Athens,
Greece : head and body, 80 and 93 ; tail, 80 and 86 ; hind foot,
20 and 18 '8 ; ear from meatus, — and 12*4.

Specimens examined. — Two, both from Tatoi, north of Athens, Greece.

MemarTcs. — Though at present very impei'fectly known the
Grecian Dyromys appears to be a well-characterized local form.

2. Tatoi, Greece. (C. Mottaz.) Hon. N. C. Eoths- 8. 10. 2. 24-25.

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572 RODENTIA

DYROMYS ROBUSTUS Miller.

1910. Dyromys lubiistus Miller, Ann. and Mag. Nat. Hist., 8th ser., vi,
p. 459, November, 1910. Type in British Museum.

Type locality. — Ru.stschuk, Bulgaria.

Geographical dixtrihution. — Known from the tyj^e locality only.

Diagnosis. — Like Dyromys nitediila but skull broader and
more robust, with relatively shorter rostrum and mvich enlarged
auditory bulke.

External cliaracters and colour. — Not appreciably different
from those of D. nitedida (no skins examined).

Sliull and teeth. — Skull of essentially the same length as in
the largest specimens of Dyromys nitedida, but conspicuously
broader and more robust ; brain-case relatively shorter and
broader^ its width above zygomatic roots nearly equal to distance
from posterior border of iiaterparietal to narrowest portion of
interorbital constriction, its posterior outline appearing more
squarely truncate when viewed from above, and wider and lower
when viewed from behind ; in lateral aspect, owing to the
enlarged bulla?, the brain-case appears deeper than in the related
species ; auditory bullae of the same form as in D. nitedida, but
so enlarged that their length from paroccipital process is nearly
equal to distance from front of bulla to infraorbital foramen ;
mandible heavy but with no special peculiarities of form. Teeth
with enamel pattern as in D. nitedida ; relative size of upper
premolar about as in D. n. wingei.

Measurements. — Type (adult female in alcohol) : head and
body, 95 ; tail, 87 ; hind foot, 22 ; ear from meatus, 15. For
cranial measurements see Table, p. 571.

Specimen examined. — The type.

9 al. Rustschuk, Bulgaria. K. Andersen (c). 11. 10. 8. 1.

{Type of species.)

Genus GLIS Brisson.

1762. Glis Brisson, Eegn. Anim. in Classis ix distrib., 2nd ed., p. 13(G7i6).
1780. Myoxus Zimmermann, Geogr. Gesch., ii, p. 351 (Type by tautonymy

Sciurus glis Linnasus).
1843. Myoxiis Wagner, Abhandl. kais Bayer. Ak. Wissensch., Miinchen,

Math.-Phys. Classe iii, p. 185 (Sub-genus of Myoxus).
1857. Glis Blasius, Siiugethiere Deutschlands, p. 288 (Sub-genus of

Myoxus). '
1890. Myoxus Reuvens, Die ]\Iyoxidae oder Schlaefer, p. 56 (Sub-genus of

Myoxus).
1895. Glis Merriam, Science, N.S., i, p. 376, April 5, 1895 (Genus).
lEOO. Elius Schulze, Zeitschr. fiir Naturwiss., Stuttgart, lxxiii, p. 200

(Sub-genus of Myoxus) part, included glis and nitedula.

Type species. — Glis Brisson = Sciurus glis Linnaeus.
Geographical distribution. — Central and southern Europe from

GLIS 573

the Atlantic coast of the mainland east to Asia Minor, north to
northern Germany, south to Sicily, Sardinia and northern
Spain.

Characters. — Skull flattened, slightly angular, the interorbital
region with lateral ridges which unite to form a median crest in
old age ; brain-case rather broad and low, the auditory bulla?
mode)-ate ; ectopterygoid reduced to a low ridge ; jugal very long
in front, extending almost or quite to lachrymal ; mandible with
angular portion entire ; dental formula as in Eliomys ; crowns of
cheek-teeth very slightly concave, the inner and outer margins
scarcely elevated, the outer side of m^ and m'^ with five low
cusps ; crowns of m} and ?»'^ alike in form, each crossed by four
complete transverse ridges, in the spaces between which lie three
incomplete ridges ; external appearance squirrel-like, the tail
conspicuously distichous.

Remarls. — The single known member of this genus is one of
the most characteristic mammals of central and southern Europe.
In the mechanics of the zygomatic arch Glis is more sciurine than
any of the other European members of the family. The structure
of the teeth and the peculiarities of external form are less
specialised than in Muscardinus.

GLIS GLIS Linnfeus.
(Synonymy under subspecies.)

Geographical distribution. — From the northern portion of the
Iberian Peninsula to Asia Minor ; north to the Baltic coast of
(lermany ; south to Sardinia and Sicily ; west to the Atlantic.

Diagnosis. — General characters as in the genus ; size largest
of the European Mmcardinidse (head and body about 160 to
190; hind foot about 27 to 34; condylobasal length of skull
about 36 to 44) ; colour greyish above, whitish below, the tail
usually darker or more slaty than body ; no blackish markings
anywhere.

External characters. — General form and appearance squirrel-
like. Details c>f structure much as in Eliomys quercinus except
in the following particulars. Ear much smaller-, extending barely
to eye when laid forward ; scarcely a trace of ridge on inner
surface of conch, but meatus with a small though evident anti-
tragus-like lobe. Feet more robust than in the other European
members of the family, but relative lengths of digits as in
Eliomys ; tubercles larger relatively to the area in which they
occur ; posterointernal palmar tubercle divided into two nearly
equal parts, on the outer surface of the anterior of which an
exceedingly small remnant of the thumb nail may usually be
detected. Sole rather broad and robust, hairy on posterior third,
its tubercles not diftering noticeably among themselves in form,
the postero-internal tubercle not elongated, but the three posterior

574

RODENTIA

larger than the three anterior. Claw of inner toe relatively
more reduced, closely appressed to pad. Tail easily broken as in
Ellomys and showing the same tendency to produce an abnormally
thickened pencil when injured, its hairs long throughout, its
form strictly distichous in adult, but terete through basal half
in young. Mammae : p2 — 2, a '1 — 1, i2 — 'l = 12.

Colour. — Upper parts ranging from a yellowish broccoli-brown
to bluish smoke-grey, a little darkened on back by a sprinkling
of long blackish hairs ; underfur slate-grey at base, the general
body colour appearing at exti'eme tips only ; hairs unusually
glossy throughout, producing a noticeable metallic silvery lustre,
especially on posterior half of back ; head faintly lighter than
body ; a narrow dusky eye-ring ; ears like surrounding parts or
somewhat dusky ; underparts and inner surface of legs pale buff,
the line of demarcation rather well defined and extending on cheek
to lower edge of eye-ring ; feet dull pale buff', irregularly clouded
with dusky or plumbeous ; tail with upper surface essentially
like back but usually darker or more slaty, its under surface
lighter, with ill-defined pale median stripe sometimes extending
nearly to pencil but more often indistinct beyond middle. Young
dull plumbeous grey above, bufty white below, the light area on
under side of tail usually better defined than in adult.

Fig. 113.
Glis glis. Nat. size.

SJcull. — In general the skull differs from that of Eliomyx
qnercinus chiefly in its broader, lower form, a peculiarity that is
especially noticeable in posterior view. Profiles essentially as in

GLis 575

E. querchms except that nasals are abruptly bent downward at
tip and venti'al border of auditory bulla does not descend evidently
below level of pterygoids. Surface of brain-case smooth, its
general outline between squarish and ovate when viewed from
above, the parietals bent downward along outer edge as in
Eliomys ; interparietal lozenge-shaped, its antero-posterior diameter
about half transverse diameter, rarely approaching ligulate.
Auditory bulla much less inHated than in Eliomys, particularly
in its posterior portion, which is not visible when skull is viewed
from above, and is not closely appi'essed to paroccipital process,
the general outline of bulla from side sub-circular, the anterior
border extending about to posterior border of glenoid surface ;
width of basioccipital at basal suture considerably more than
half distance from suture to foramen magnum instead of barely
one-third this distance as in Eliomys. Interorbital region with
well developed lateral ridges, low in the young, becoming higher
in adults and finally uniting to form a median crest. Zygomata
more abruptly spreading than in Eliomys, their median portion
often parallel, the middle of each jugal angularly expanded
upward, the zygomatic process of squamosal with abrupt posterior
concavity when viewed from above. Posterior termination of
nasals pointed or truncate, about at level of nasal branch of
premaxilla and front of lachrymal. Incisive foramina as in
Eliomys but shorter, the greatest length about one-third that of
diastema. Palate and mesopterygoid space essentially as in
Eliomys, but hamular not in contact with bulla, and ectopterygoid
reduced to a low ridge. Mandible much more robust than that
of Eliomys, especially in its anterior portion ; coronoid process
nK)re robust, strongly curved, its point high above level of
condyle ; angular portion entire.

Teeth. — Incisors as in Eliomys but more robust. Upper
cheek-teeth relatively larger than in Eliomys, their outlines
s(][uarish, the first and second molars sub-equal, the third smaller,
narrowed posteriorly ; premolar slightly less than half as large
as first molar. Crowns low and nearly flat, but inner and outer
border slightly raised. The crown of each tooth is crossed by
four complete ridges, the postei'ior of which is least developed ;
between these three incomplete ridges extend inward to or a
little beyond middle, the second reaching outer side. These ridges
are best developed in m^ and ?h-, least developed in the premolar, in
which the first is often absent. Inner border of crown with four
low tubercles formed by the inner extremities of the four main
ridges, the fourth smaller than any of the others. Outer margin
of m^ and m- with five similar but smaller tubercles formed by
the outer extremities of the four main ridges and the middle
incomplete ridge. In the premolar the incomplete ridge does
not extend to outer margin of crown, while in m^ the structure
of all the ridges becomes indefinite near periphery of crown.
Lower molars essentially like the maxillary teeth, but with

576 llODENTIA

arrangement reversed, the thicker more tuberculate termination
of ridges at outer side of crown, and none of the intermediate,
ridges extending to border ; lower premolar usually with oidy

FlG. 114.
Glis ijlis. Cheek-teetli. X 10.

two incomplete ridges (the second and third) ; m., with crown
longer and all the ridges better developed than in m^.

BcmarJcs. — Glis glis is immediately recognizable among
European members of the family by its large size, squirrel-like
aspect, and the absence of dark markings on face. Three local
races are known to occur in Europe and a fourth in Asia Minor.
A fifth, which I have not seen, has been described from northern
Spain.

KEY TO THE EUROPEAN RACES OF GLIS GLIS.

Size medium, hind foot usually less than 30 mm. ; cou-
dylobasal length of skull usually 35 to 39 mm.

(Central Europe, south to northern Italy) G. glis <jlis, p. 577.

Size large, hind foot usually more than 30 mm. ; condy-
lobasal length of skull usually 39 '6 to 44 mm.
Skull frequently more than 42 mm. in condylobasal
length ; dark terminal area of tail often involving

distal half or more (Italy and Sicily) G. g. italicus, p. 578.

Skull rarely more than 41 mm. in condylobasal
length ; dark terminal area of tail usually con-
fined to distal third or less (Sardinia) G. g. melonii, p. 579.

577

Glis glis glis Liunteus.

1766. [Sciiirus] glis Linnaeus, Syst. Nat., i, 12th ed., p. 87 (Central

Europe).
1779. [Gils'] esciilentusBlumenhsich, Handb. der l^aturgescb., p. 79 (Central

Europe).
1816. Glis vulgaris Oken, Lehrbuch der Naturgesch., iii, pt. 2, p. 868

(Germany).
1832. ? Mlyoxii.s] giglis P. Cuvier, N. Ann. Mus. d'H. N. Paris, i, p. 444.

Nomen nudum, " le loir proprement dit."
1840-45. Myoxus avcllanus Owen, Odontography, ii, p. 25, pi. 105.
1857. Myoxus glis Blasius, Siiugethiere Deutschlands, p. 292.
1890. Myoxus glis Reuvens, Die IMyoxidae oder Schlaefer, p. 61.
1895. Gtis (sic) glis Merriam, Science, N.S., i, p. 376, April 5, 1895.
1910. Glis glis Trouessart, Faune jNIamm. d'Europe, p. 181.

Type locality. — Germany.

GeograjMcal distribution. — Central Europe from northern
Germany (Mecklenburg) to the Pyrenees and northern Italy,
and from the Atlantic coast eastward.

Diagnosis. — Size medium, the hind foot about 30 mm. or less,
the condylobasal length of skull about 35 to 39 mm. ; molars
not enlai'ged ; general colour of uj^per parts a yellowish broccoli-
brown scai'cely darkened by the longer blackish hairs ; tail
essentially concolor with body, the extremity usually tinged
with drab, but hardly ever noticeably darker than basal portion.

Measurements. — Adult male from Ziirich, Switzerland : head
and body, 165; tail, 150; hind foot, 29. Adult male from
Bruggen, St. Gallen, and adult male from Mels, Rheinthal,
St. Gallen: head and body, 180 and 170; tail, 120 and 130;
hind foot, 28 and 29. Adult female from Hatszeg, Hunyad,
Hungary: head and body, 147; tail, 135; hind foot, 29 "4.
Adult male and female from Ceresole d'Alba, Turin, Italy : head
and body, 170 and 176 ; tail, 146 and 149 ; hind foot, 27*4 and
27. Sixteen adults from various localities in northern Italy
(Genoa) : hind foot, 27 to 28 '5. For cranial measurements see
Table, p. 580.

Specimens examined. — Ninety-two, from the following localities : —

France : No exact locality, 5 ; near Nimes, Gard, 4.

Germany: Saxony, 1 (U.S.N.M.) ; Bavaria, 1 (U.S.N.M.) ; Marxheim,
Bavaria, 2 ; Ekernkrug, 1.

Austria-Hungary : Steiermark, 1 ; HoUos, Eisenburg, 1 (U.S.N.M.) ;
Trieste, 1 ; Herkulesbad, 1 ; Hatszeg, Hunyad, 6.

Switzerland ; Les Plans, Vaud, 3 (U.S.N.M.) ; Interlaken, Bern, 1
(U.S.N.M.) ; Miirren, Bern, 1 ; Lucerne, 1 ; St. Gothard, Uri, 1 ; Ziirich, 1
(U.S.N.M.) ; Thayngen, Schaffhausen, 2 (B.M. and U.S.N.M.) ; Bruggen,
St. Gallen, 1 (U.S.N.M.) ; Degersheim, St. Gallen, 2 (U.S.N.M.) ; Toggen-
burg, St. Gallen, 1 (U.S.N.M.). ; Mels, St. Gallen, 2 (B.M. and U.S.N.M.) ;
Wolfhalden, Appenzell, 1 (U.S.N.M.) ; Biasca, Ticino, 1 (U.S.N.M.) ; Bug-
giolo, Ticino, 1 (U.S.N.M.); Carmago, Ticino, 2 (U.S.N.M.); Lonvico,
Ticino, 1 (U.S.N.M.) ; Vezia, Ticino, 1 (U.S.N.M.) ; no exact locality, 1.

Italy : Padola, Cadore, 1 (Turin) ; Porlezza, Como, 23 (Ghidini) ;

2 P

578

RODENTIA

Gozzano, Novara, 3 (Genoa) ; Ceresole d'Alba, Turiu, 10 (Turin) ; Monca-
lieri, Turin, 7 (Turin).

2 (5, 2 9. Nimes, Gard, France. 0. Thomas (p). 8. 8. 10. 53-56.

(C. Mottaz.)

2, 2 juv. France. Purchased 46. 1. 2. 1-4.

(Lefebvre).

1. France. A. Baillon (c). 40. a.

2 ?. Marxheim, Bavaria. Lord Lilford (p). 11. 1. 1. 102-103.

{Sdiucliardt.)

?. Ekernkrug, Germany. Lord Lilford (p). 11. 1. 1. 104.

1. Trieste, Austria-Hungary. Lord Lilford (p). 11. 1. 1. 152.

{J. G. Haggard.)

6. Herkulesbad, Austria- Hon. N. 0. Roths- 7. 9. 16. 11.
Hungary. {F. J. Cox.) child (p).

4 cJ, 2 ?. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 21-26.
sylvania.

1. Miirren, Bern, Switzer- W. Giirtner (c & p). 92.10.5.4.

land.

1. Lucerne. E. Cavendish Taylor 5.5.6.15.

(c & p).

1. St. Gothard, Uri, Swit- Tomes Collection. 7. 1. 1. 75.
zerland. (Verreaux.)

9 juv. Thayngen, Schaffhausen. O. Thomas (p). 2. 8. 4. 35.

(E. H. Zollikofer.)

9. Mels, St. Gallen". {E. H. 0. Thomas (p). 2. 8. 4. 34.

Zollikofer.)

i St. Switzerland. E. R. Alston (p). 79. 9. 12. 1.

Glis glis italicus Barrett-Hamilton.

1898. Glis italicus Barrett-Hamilton, Ann. and Mag. Nat. Hist., 7th ser.,

II, p. 424, November, 1898 (Siena, Italy). Type in British
Museum.

1899. ? Glis insularis Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., iii, p. 228, March, 1899, Monte Aspro, near Palermo,
Sicily. Type in British Museum.
1910. Glis italicus and ? Glis insularis Trouessart, Faune Mamm.
d'Europe, p. 132.

Type locality. — Siena, Italy.

Geographical distribution. — Italy, from region of Genoa,
southward ; Sicily 1

Diagnosis, — Larger than Glis glis, the hind foot more than
30 mm., the condylobasal length of skull about 40 to 44 mm. ;
cheek-teeth distinctly enlarged ; the incomplete cross ridges
tending to be better developed ; general colour of upper parts a
clear broccoli-brown noticeably darkened by the longer blackish
hairs ; tail usually much darker than body, its terminal half
drab, slaty, or occasionally blackish.

Measurements. — Adult male from the type locality : head and
body, 190 ; tail, 152 ; hind foot, 33 ; ear from meatus, 24. Type
and a second adult female from the type locality : head and
body, 180 and 187; tail, 153 and 130 (probably injured); hind
foot, 34 and 33*4; ear from meatus, 23 and 23. Adult female
from Monte Sirino, Lagonegro : head and body, 175; tail, 150;

uLis 579

hiud foot, 34; ear from meatus, 21. Young female fi'om near
Palermo, Sicily (tyj^e of insularly) : head and body, 160; tail,
130; hind foot, 31 ; ear from meatus, 21. For cranial measure-
ments see Table, p. 581.

Specimens examined. —Forty, from the following localities in Italy :
near Genoa, 5 ; Tana del Mosto, Finalborgo, Genoa, 1 (Genoa) ; Perti,
Finalborgo, Genoa, 6 (Genoa) ; near Vittoria, western Liguria, 2 (Genoa) ;
Caluzzano, 1 (Genoa) ; Malasana, 2 Genoa) ; near Montariolo, 1 (Genoa) ;
Verona, 1 ; Florence, 1 ; Siena, 8 ; Viterbo, near Rome, 2 ; Monte Cimino,
Rome, 7 (Genoa) ; Monte Sirino, Lagonegro, 1 ; near Palermo, Sicily, 2.

RemarJis. — Throughout southern and central Italy, north
to about the region of Genoa, the large dormouse appears to be
very constant in its characters, diftering from that of central
Europe in its greater size and heavier teeth, as well as in its
somewhat darker coloration. Further north, notably in the
region of Turin and at Porlezza, the colour becomes essentially
as in true glis, but the size remains a little above that of the
typical form. These northern specimens are probably best
treated as intermediates between glis and italicus, lying nearer
to the former than to the southern race. The two immature
specimens from Sicily which formed the basis of Glis insularis
are in very bad condition. Allowance being made foi" this, there
appears to be nothing to distinguish them from the ordinary
south-Italian form.

2 9. Genoa, Italy. Genoa Museum (e). 7. 2. 28. 1-2.

6, 2 ?. Genoa. Marquis G. Doria(p). 89. 12. 11. 1-3.

9. Verona. {Conie delle 0. Thomas (p). 99. 11. 10. 4.

Oddi.)
9. Florence. A. Savage Landor 97. 3. 7. 2.

(c & P).
6,5 9. Siena. (Brogi.) Dr. E. Hamilton (p). 98.10.2.11-16.

(98. 10. 2. 14. Type of subspecies.)
2. Siena. (Brogi.) G.Barrett-Hamilton 11. 1. 2. 60-61.

9. Monte Sirino, Lagonegro. 0. Thomas (p). 6.8.4.1.

(A. Robert.)
2 9. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 4-5.

(98. 10.6.4. Ty2}e of G. insularis Barrett-Hamilton.)

Glis glis melonii Thomas.

1907. Glis melonii Thomas, Ann. and Mag. Nat. Hist., 7th ser., xix,

p. 445, ]\Iay, 1907. Type in British Museum.
1910. Glis melonii Trouessart, Faune Mamm. d'Europe, p. 182.

Type locality. — Marcurighe, Urzulei, Ogliastra, Sardinia.

Geograpliical distribution.- — Sardinia and jjrobably Corsica also.

Diagnosis. — Similar to Glis glis italicus, but not attaining so
large a size (skull rarely more than -11 mm. in condylobasal
length), and dark terminal area on tail not so extensive, seldom
involving more than distal third.

2 p 2

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Measurements. — Type (adult male) : hind foot, 32 • 6. Two
other adult males from the type locality : head and body, 154
and 160; tail, 134 and 142; hind foot, 30 and 32; ear from
meatus, 20 and 20. Adult female from the same locality : head
and body, 165; tail, 134; hind foot, 32; ear from meatus, 20.
For cranial measurements see Table, p. 581.

Specimens examined.— Twentj-iour, all from Sardinia (B.M. and
Genoa) .

JRemarks. — Although not a strongly marked race the Sardinian
Glis differs appreciably in average characters from that of the
Italian mainland. Its most obvious features are the slightly
smaller size and less darkened tail.

6 6,5 9. Sardinia. G. Meloni (c). 8. 4. 6. 1-11.

(8. 4. 6. 1. Type of subspecies.)

Note. — The Spanish form of Glis has been distinguished sub-
specitically from the other European races. I have not seen the
animal, and the following account is taken entirely from the
original description.

Glis glis pyrenaicus Cabrera.

1908. Glis glis pyrenaicus Cabrera, Ann. and Mag. Nat. Hist., 8th ser., i,
p. 193, February, 1900.

Type locality. — Alio, Navarra, Spain.

Geographical distribution. — Northern Spain ; exact limits of
range unknown.

Diagnosis. — Similar to the typical form in all essential
respects, but readily distinguishable by larger skull, and strong
buffy tinge of back.

Colour. — Upper parts buffy grey, the hairs being iron-grey
with yellowish buff ends. In the middle of the back there are
numerous black hairs, showing a bright metallic gloss. Under
surface creamy white, separated from the upper colour by a
narrow, ill-defined zone of pure yellowish buff extending from
the cheek to the hip. Tail glossy brownish grey, with the usual
whitish line along under side. Ears and orbital rings brown.
Hands and feet white ; a broad brown metatarsal patch.

Skull. — Like that of typical G. glis, but larger, approaching
that of Glis glis italicus.

Measurements. — Type (in flesh): head and body, 169; tail-
vertebrje, 137; hind foot, 28; ear, 16. Skull: greatest length,
41*5; basilar length, 33; zygomatic breadth, 24; breadth of
brain-case, 18; interorbital breadth, 5' 5; length of nasals, 14;
palatilar length, 16 ; diastema, 10; upper tooth-row, 7*5.

MUSCARDINUS 583

Genus MUSCARDINUS Kaup.

1829. Muscardiniis Kaup, Entw.-Gescb. u. Natiirl. Syst. Europ. Thierwelt,

I, p. 134.
1843. Muscardinus Wagner, Abhandl. k. Bayer. Akad. Wissensch., Miin-

cheu, Math.-Phys. CI., iii, p. 185 (Sub-genus of Myoxus).

1857. Muscardinus Blasius, Saugetbiere Deutscblands, p. 289 (Sub-genus

of Myoxus).
1890. Muscardinus Reuvens, Die Myoxidae oder Scblaefer, p. 25 (Sub-genus
, of Myoxus).

Type species. — Mus avellanarius Linnpeus.

Geographical distribution. — From England east into Asia
Minor and from central and southern Sweden to the Mediter-
ranean coast and Sicily, though not at present known from the
Iberian and Balkan Peninsulas.

Characters. — Skull deep, as in Eliomys and Dyromys, but inter-
orbital region flattened, with slightly raised edges, its median
portion usually with one or two small irregular perforations at
narrowest part ; ectopterygoid absent ; jugal moderately long in
front, extending about half-way up anterior border of orbit ;
mandible with angular portion fenestrate ; dental formula as in
the other European members of the family ; crowns of cheek-
teeth essentially flat, the margins not elevated, the outer border
of m} and m^ with five or six low cusps ; crowns of m^ and m^
conspicuously iinlike in size and form, the five cross ridges of the
larger first molar mostly oblique, the seven ridges of the smaller
second molar strictly transverse ; external form not peculiar ;
tail long, moderately bushy ; feet more specialized than in the
other European members of the group.

Remarks. — This genus with its flat-crowned molars and
unusually prehensile feet represents the most highly specialized
stage attained by the European Muscardinidse. As might be
anticipated the habits of the species are more strictly arborial
than in the other genera. Three species are now known, two
of which occur in Europe, the other in Asia Minor.

KEY TO THE EUROPEAN SPECIES OP MUSCARDINUS.

Upper parts dull yellowisb brown ; no sbarp line
of demarcation along sides ; no wbite stripe
between ear and eye (Distribution general)... M. avellanarius, p. 583.

Upper parts brigbt buff ; a sbarp line of demarca-
tion along sides ; a vertical wbite stripe
between ear and eye (Central Italy) M. pulcher, p. 590.

MUSCARDINUS AVELLANARIUS Linnseus.

1758. [_Mus'] avellanarius Linnasus, Syst. Nat., i, lOtb ed., p. 62 (Sweden).
1782. Myoxus muscardinus Scbreber, Saugtbiere, jil. ccxxvii ; text, iv,
p. 835, 1788 (Germany).

1858. Myoxus avellanarius Blasius, Saugetbiere Deutscblands, p. 297.

584

RODBNTIA

1869. Mas corilinum Fatio, Faune Vert. Suisse, i, p. 183 (name attributed
to Schreber).

1890. Muscardinus avellanarius Reuvens, Die Myoxidae oder Schlaefcr,
p. 69.

1900. M{^iiscardimis'] avellanarius anglictis Barrett-Hamilton, Proc. Zool.
Soc, London, p. 86 (Bedford Purlieus, Thornhaugh, Northampton-
shire, England). Type in British Museum.

1910. Mnscardinus avellanarius and M. avellanarius anglicus Trouessart,
Faune Mamm. d'Europe, p. 135.

Type locality. — Central Sweden.

Geographical distribution. — Throughout central Europe from
England eastward ; north to central Sweden, south to the
Pyrenees and to Rome, Italy.

Diagnosis. — Smallest European member of the family, the
general size not much greater than that of a large house mouse ;
colour dull yellowish brown above, buffy beneath, the chin and
throat usually whitish, the flanks between the clay-colour and
ochraceous-bufF of Ridgway ; no whitish strijDe in front of ear.

External characters. — General form essentially as in Eliomys
quercinus, but eye relatively lai'ger and more prominent, ear
relatively shorter (extending to outer canthus of eye when laid
forward), its inner surface with conspicuous ridge above meatus
as in Dyromys, and tail uniformly haired throughout. Fore
foot with digits relatively longer than in the other European
genera, and closing obliquely inward so as to come in opposition
with the much enlarged inner tubercle, the unusual size of which,
about equal to that of all the other tubercles together, enables
it to function as a low, broad thumb ; terminal lobe of this
tubercle representing true thumb small though evident ; propor-
tional lengths of fingers as in Eliomys. Hind foot rather short
and broad, as in Glis, the sole naked to heel, the two inner
tubercles somewhat enlarged, but not specially elongated ; inner
digit a mere rudiment scarcely larger than its corresponding
tubercle, its claw obsolete ; other digits relatively longer than
in the related genera. Tail partly prehensile, slightly
flattened, extending about to ears when laid forward, uniformly
haired and loosely short-bushy throughout. Mammre :
pl_l, al-l i2-2 = 8.

Colour. — Entire upper parts and outer surface of legs
yellowish brown, the exact shade between the wood-brown or
clay-colour and bufi" of Ridgway, sometimes with a slight greyish
or ecru-drab cast across shoulders, the middle of back incon-
spicuously sprinkled with blackish hairs ; sides, flanks, cheeks and
ears tinged with ochraceous-buflf ; tail concolor with back above,
faintly lighter below, the tip with a dusky tinge ; underparts
and inner surface of legs light ochraceous-buff, slightly contrasted
with sides and without sharp line of demarcation, the hairs,
like those of back, slate-grey at base ; chin and throat with an
irregular area of pure white (to base of hairs), this occasionally

MUSCARDINUS

585

extending back along median line to middle of belly, but
anteriorly seldom covering lower lip and never extending upward
between ear and eye ; a family indicated, very narrow blackish
eye-ring ; feet ochraceous-buff, the toes sprinkled with silvery
hair.

Skull. — In general form the skull differs from that of the
other European Muscardinidse in greater depth of brain-case,
particularly at its anterior portion, the posterior half sloping
away much more noticeably than in the allied genera, the
occipital region low. Interparietal usually narrowed to a point
at each lateral extremity. Auditory bulla essentially as in Gils,
but a little more inflated posteriorly, so as to be just visible
when skull is viewed from above. Interorbital region with
longitudinal median depression and slightly though distinctly
angular-elevated edges, the elevated regions never coming together
in old age. Zygomata very abruptly
spreading anteriorly so that the arches
of the two sides are essentially parallel
through the greater part of their extent ;
infraoi'bital foramen relatively smaller
and more nearly circular in outline than
in any of the other European members
of the family. Nasals and nasal branches
of preniaxillaries terminating at level
of front of lachrymal, the posterior
border of nasals usually angulate-emar-
ginate. Incisive foramina not widened
behind middle, their aspect more Murine
than Muscardinine. Posterior border of
palate emarginate to level of middle or
front of m", the palatine bones reduced
to a mere rim at front of mesopterygoid
space. Pterygoid fossa and ectopterygoid

absent. Mesopterygoid fossa parallel-sided, about three times
as long as wide ; hamulars long and very slender, a little curved
upward. Mandible relatively more robust than in any of the
other European members of the family ; dental foramen noticeably
above alveolar level ; hamular process short, its point scarcely
rising above level of condyle ; angular region robust, very sharply
angled below ; a distinct vacuity above angle.

Teeth. — In general aspect the cheek-teeth differ from those
of the other European Muscardinidse in their relatively greater
size, the flatness of their crowns, and in the more marked
contrasts of size and form shown by the premolar and first two
molars. Upper incisor with no special peculiarities. Lower
incisor with root extending distinctly beyond that of m^ and
forcing the dental foramen upward to a level slightly above that
of crushing surface of molar crowns. Premolar both above and
below small, single rooted, its crown variable in outline, but

FIG. 115.

Muscardinus avellanarius.
Nat. size.

586

RODENTIA

usually subterete or a little flattened against succeeding tooth,
its area about one-fifth that of first molar ; cross ridges two or
sometimes three, the third smaller than the others and lying at
back of upper tooth and front of lower tooth, the two main
ridges of upper premolar usually joined at inner side, and
occasionally at outer side also, producing a circular, raised rim.
Milk premolar minute, not functional, its crown bluntly spicular,
smooth, scarcely one-quarter as large as that of permanent tooth.
First upper molar about one-half greater than that of second, its
length decidedly greater than its width, the anterior border
oblique, not so long as posterior border, the inner border with
a slight though abrupt emai'gination anteriorly ; surface crossed
by five high, very distinct ridges, the first three wider apart than
the last two and sloping obliquely backward toward inner
margin, the first independent of the others, the second, third,

fourth, and fifth continuous with
a ridge which extends along inner
border as far forward as space
separating first ridge from the
others ; when viewed from the
side the outer terminations of the
five ridges have the appearance of
five low cusps, decreasing regularly
in height from before backward ;
on inner margin there is a single
anterior cusp, the rest of the crown
appearing flat. In some specimens
there is a rudimentary ridge at
extreme anterior border of tooth
and traces of another in the space
between third and fourth main
ridges. Second upper molar
squarish in outline, its length
slightly greater than its width ;
surface of crown crossed by seven low but complete, squarely
transverse, equally spaced ridges, all* extending from border to
border, the outer extremities appearing as five or six low cusps.
Third upper molar like second, but smaller and with posterior
border rounded, the ridges behind the first three or four
usually incomplete and confused. Lower molars essentially
alike in structure, the crown of each tooth crossed by six
ridges rather more distinct than those of iv? (except in m^),
their terminations producing the usual effect of low cusps along
outer border ; crown area diminishing regularly from m-^ to m^
(which is only a little more than half that of anterior
tooth) ; iHo and m^ squarish in outline, slightly longer than wide ;
TOj decidedly longer than wide, narrower anteriorly than
posteriorly.

* Except occasionally the third.

Fig. 116.

Muscardinus aveUanariiis.
Cheek-teeth, x 10.

MUSCARDINUS

587

Measurements. — Adult male from Thornhaugh, Northampton-
shire, England:* head and body, 86; tail, 57; hind foot, 16.
Adult female from Colchester, Essex : head and body, 77 ;
tail, 55; hind foot, 16; ear from meatus, 11. Adult male and
female from Guines, Pas-de-Calais, France ; head and body,
72 and 77 ; tail, 71 and 74; hind foot, 16-2 and 16-4. Adult
male and female from Lucinges, Haute-Savoie, France : head
and body, 78 and 76 ; tail, 61 and 66 ; hind foot, 15 and 15 ;
ear from meatus, 11 and 10. Adult male from Lausanne, Vaud,
Switzerland: head and body, 75; tail, 77; hind foot, 16" 5.
Adult female from Padola, Cadore, Italy : head and body, 80 ;
tail, 70; hind foot, 16; ear from meatus, 12. Adult female
from Arsoli, Rome, Italy : head and body, 75 ; tail, 65 ; hind
foot, 15 '6. For cranial measurements see Table, p. 588.

Specimens examined. — Ninety, from the following localities : —

England : Thornhaugh, Northamptonshire, 1 ; Halton, Buckingham-
shire, 1 ; Chalfont, Buckingham, 1 ; Bury St. Edmunds, Suffolk, 2 ;
Colchester, Essex, 3; London (purchased alive in), 2; Norwood, Headley,
Surrey, 1 ; Ratham, Sussex, 1 ; Sussex, no exact locality, 1 ; Eversley,
Hampshire, 2 ; Honiton, Devonshire, 1 ; no exact locality, 1.

France : Guines, Pas-de-Calais, 2 ; Manonville, Meurthe-et-Moselle, 4 ;
Cranves-Sales, Haute-Savoie, 2 ; Lucinges, Haute-Savoie, 2 ; Montauban,
Haute-Savoie, 1 ; Scientriers, Haute-Savoie, 1 (IMottaz) ; no exact locality, 1.

Germany : Oberwald, near Gross, Silesia, 1 ; Silesia, no exact locality,
1 ; Wolfshau, Riesengebirge, Silesia, 1 (U.S.N.M.) ; Nuremberg, Bavaria,
2 (U.S.N.M.).

Austria-Hungary: Csallokoz-Somorja, Pressburg, Hungary, 3; Zuberet^,
Hungary, 1 ; Tirol, 2 (U.S.N.M.) ; Brasso, 1 ; Hatszeg, Hunyad, 5.

Switzerland : Geneva, 4 (Genoa and Mottaz) ; Lausanne, Vaud, 2
(U.S.N.M.); Neuchatel, 3 (U.S.N.M.); St. Gallen, 2; Ziiberwangen,
St. Gallen, 1 (U.S.N.M.) ; Vulpera-Tarasp, Grisons, 5 (Rothschild) ;
Ticino, 1 (U.S.N.M.) ; Lugano, Ticino, 1 (U.S.N.M.).

Italy : Padola, Cadore, 2 (Turin) ; Porlezza, Como, 3 ; Gozzano,
Novara, 1 (Genoa) ; Moncalieri, Turin, 1 (U.S.N.M.) ; Timone, Borzoli,
Genoa, 1 (Genoa) ; N.S. della Vittoria, Appennino Ligure, 9 (Genoa) ;
Vaccarezza, 2 (Genoa) ; Siena, 4 ; Rome, 1.

Bemarks. — Skins from different parts of the range of Mus-
cardinus avellanarius show practically no individual variation.
Adults are darker and more richly coloured than the young,
though the difference is slight. British specimens are not
distinguishable from those taken in Switzerland and Italy. f

i. Halton, Buckingham- Hon. N. C. Roths- 11. 1. 3. 490.

shire, England. child (p).

?. Chalfont, Buckingham- Hon. N. C. Roths- 11. 1. 3. 401.

shire. child (p).

6. Thornhaugh, Northamp- Rev. H. H. Slater 99. 11. 27. 6.

tonshire. (c & p).

{Type of M. a. angliciis Barrett-Hamilton.)

S, 9 al. Bury St. Edmunds, Suf- Dr. Giinther (c & p). 90. 5. 27. 1.
folk.

* Type of anglicus Barrett-Hamilton.

f The type of anglicus is an adult in full, rich winter pelage.

588

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RODENTIA

6.

Purchased in London.

G. Barrett-Hamilton
E. R. Alston (p).

11. 1. 2. 113.

6.

Ratham, Sussex.

79. 9. 12. 2.

(W. Jeffrey.)

1.

Sussex.

E. Cavendish Taylor
G. C. Shortridge

11. 1. 3. 403.

6.

Honitou, Devonshire.

11. 1. 3. 402.

(c & P).

1.

England.

Yarrell Collection.

56. 12. 10. 682.

6,9.

Guiues, Pas - de - Calais,
France.

0. Thomas (c & p).

94. 6. 6. 12-13.

3 9, juv.

Manonville, Meurtbe-et-
Moselle. {Lomont.)

Lord Lilford (p).

11.1.1.105-107

2 6.

Cranves - Sales, Haute-
Savoie, 1200 m.

{A. Bobert.)

0. Thomas (p).

5. 11. 18, 9-10.

6,9.

Lucinges, Haute-Savoie.
(.4. Robert.)

0. Thomas (p).

6. 4. 2. 4-5.

6.

Llontauban, Haute-Savoie,
900 m. (A. Bobert.)

, 0. Thomas (p).

6. 4. 7. 6.

1 St.

Prance. (F. Petit.)

E. Hargett (p).

97. 11. 25. 4.

6.

Oberwald, Silesia, 400 m.
Germany. (T7. Baer.)

Lord Lilford (p).

99. 1. 9. 16.

?al.

Silesia.

Dr. Kaup (p).

3.

Csallokoz-Somorja, Hun-
gary.

Budapest Mus. (e).

94. 3. 1. 43-45.

1.

Zubereg, Hungary.

Budapest Mus. (e.)

94. 3. 1. 42.

9al.

Brasso, Hungary.

Prof. L. von Mehely
C. G. Danford (c).

95. 8. 12. 3.

1,4 juv.

Hatszeg, Hunyad, Tran-

3. 11. 8. 27-31.

sylvania.

9, 9 juv.

St. Gallen, Switzerland.
{E. H. Zollikofer.)

0. Thomas (p).

2. 8. 4. 36-37.

9.

Siena, Italy. (Brogi.)

G. Barrett-Hamilton

11. 1. 2. 112.

'

(P).

MUSCARDINUS PULCHER Barrett-Hamilton,

1855. ? Myoxus speciosus Dehne, Allgem. deutsche Naturhist. Zeitung,

neue Folge, i, p. 180, Tursi, Basilicata, Italy.
1898. Mnscardinus pzolcher Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., ii, p. 423, November, 1898, Perugia, Italy.
1900. M[usca7-dinus'] avellanarius speciosus Barrett-Hamilton, Proc. Zool.

Soc, London, p. 87.

1910. Muscardinus avellanarius speciosus Trouessart, Faune ]Mamm.
d'Europe, p. 136.

Type locality. — Perugia, province of Perugia, Italy.

Geoijrapliical disiribiition. — At present known from Perugia,
Rome, and the vicinity of Naples, Italy.

Diagnosis.- — Like Muscardinus avellanarius but general colour
brighter, the flanks nearly raw-sienna; white of throat extending
upward as a conspicuous whitish stripe between eye and ear.
Teeth somewhat larger than in M. avellanarius.

External clmraciers. — In external form I can detect no diiFer-
ence between this species and Muscardinus avellanarius ; mammae 8,
arranged as in the more northern animal.

MUSCARDINUS 591

Colour. — Upper parts including ears and entire tail very
uniform yellowish buff, the exact shade perhaps best described
as intermediate between the buff-yellow and raw-sienna of
Ridgway, somewhat brighter on sides and flanks, faintly darker
along middle of back ; feet concolor with back, the toes
suffused with whitish ; underparts strongly contrasted pale
cream-buff, the line of demarcation well-defined and conspicuous,
the hairs slaty grey at base ; chin and throat usually though not
always with pure white area as in 31. avellanarius, this area when
present not forming noticeable contrast with rest of underparts.

Skull and teeth. — The skull and teeth do not differ appre-
ciably from those of Muscardinus avellanarius, though the teeth
are usually a trifle larger.

Measurements. — Type (adult male) : head and body, 90 ;
tail, 68 ; hind foot, 16 ; ear from meatus, 12. Adult male from
Monte Cimino, Rome : head and body, 74 ; tail, 70 ; hind
foot, 18*5. Average and extremes of ten males from Sorrento :
head and body, 79 • 4 (71-86) ; tail, 65 • 3 (63-73) ; hind foot, 15 ■ 7
(15-17). Average and extremes of ten females from Sorrento:
head and body, 79-9 (75-85); tail, 69-6 (62-73); hind foot,
15*9 (15-17). For cranial measurements see Table, p. 589.

Specimens examined. — Fifty-seven, from the following localities in Italy :
Perugia, 1 (type) ; Monte Cimino, Rome, 1 (Genoa) ; Sorrento, 54
(U.S.N. M.) ; Palermo, Sicily, 1 (too young to be positively identified).

ItemarJcs. — This species is well differentiated from Muscar-
dinus avellanarius by its bi'ight, light colour and by the white
streak between ear and eye. It may eventually prove to be
identical with the Myoxus speciosus of Dehne from the extreme
south of Italy, but the description of the latter indicates a much
darker more reddish animal. A single immature specimen
(No. 8. 9. 28. 1) from Palermo, Sicily, too young and in too
bad condition to be satisfactorily identified, is noticeably different
from any of the numerous young individuals from Sorrento. In
its " foxy " red colour it agrees with the description of Dehne's
animal.

6. Perugia, Italy. (Brogi.) Dr. E. Hamilton (p). 98. 10. 2. 17.

[Type of species.)
?. Palermo, Sicily. J. I. S. Whitaker (p). 8. 9. 28. 1.

Faaiily MURID^.

1821; Muridse Gray, London Med. Eepos., xv, p. 303, April 1, 1823.

Geographical distribution. — As in the order Rodentia.

Characters. — Anterior j^ortion of zygomatic arch not formed
chiefly by jugal ; infraorl^ital foramen large, wider above than
below, without accessory canal; jugal bone splint-like, supported
by long zygomatic processes of maxillary and squamosal, not
in contact with lachrymal anteriorly ; mandible with angular

592

RODENTIA

portion arising from under side of alveolus of incisor, its lower
border not angulated ; tibia and fibula joined ; cheek-teeth never
more than i;"?, rooted or rootless, the crowns tuberculate or
prismatic.

BcmarJiS. — The cosmopolitan family Muridse is more abun-
dantly represented in genera, species and individuals than any
other similar group of mammals. About a dozen sub-families
are currently recognized, three of which occur in Europe.

KEY TO THE EUROPEAN SUB-FAMILIES OF MURIDM.

Molars prismatic, hypsodont or rootless, their crowns

liat (Voles and Lemmings) Microtinx, p. 610.

Molars tuberculate, brachyodont, rooted.

Tubercles of maxillary molars arranged in two primary

longitudinal series Cricetiniv, p, 592.

Tubercles of maxillary molars arranged in three

primary series Murinie,'p. 191.

Sub-Family CRICETIN^.
1866. Cricetinge Murray, Geogr. Distrib. Mamm., p. 358.

Geographical distribution. — Entire American Continent ; cen-
tral region of Asia from eastern China west into south-eastern
and centi-al Europe ; southern Africa.

Characters. — As in the sub-family Murinse, but tubercles of
maxillary teeth arranged in two primary longitudinal rows.

Bemarks. — The sub-family Cricetinse, though characteristically
American, is represented in the Old World by about six genera,
three of which occur in Europe.

KEY TO THE EUROPEAN GENERA OF CBICETINM.

General external and cranial characters essentially
murine ; fur with no special colour pattern (a
narrow dark dorsal stripe in some oriental
species) ; form of rostrum and brain-case about

as in Apodemus (Dwarf Hamsters) Cricetulus, p. 593.

General external and cranial characters not murine,
the body heavy and thick- set ; fur with specialized
colour-pattern of contrasted dark and light areas ;
rostrum broadened and brain-case narrowed, the
parietal bones reduced in size (True Hamsters).
Anteorbital foramen with well-developed forward-
projecting external plate ; flattened area on

dorsal surface of brain-case approximately

parallel-sided ; tail evident, longer than hind

loot ; mammse 8 (Central Europe, west into

Belgium and France) Cricekis, p. 596.

Anteorbital foramen without forward-projecting

external plate ; flattened area on dorsal surface

of brain-case truncate diamond- shaped ; tail

nearly concealed in the fur, shorter than hind

foot ; mammse 16 (Eastern Europe, west into

Roumania and Bulgaria) Mesocricetus, p. 605.

CRICETULUS 593

Genus CRICETULUS Milne-Edwards.

1867. Cricctulus Milne-Edwards, Ann. des Sci. Nat., Paris, 5th ser., Zool.,
VII, p. 375.

Type species. — Gricetulus (jriseus Milne-Edwards.

Geographical distribution. — G-entral region of Asia from
eastern China west through Asia Minor to the eastern portion
of the Balkan Peninsula. *

Characters. — Like Cricetus, but less highly modified: form
murine ; feet with normally developed tubercles ; tail in
European specie?, well developed, slightly longer than hind foot ;
skull murine in appearance, with large (normal) interparietal
and smooth brain-case and interorbital region, its general aspect
much as in Apodemiis ; ectopterygoid fossa long and shallow;
anteorbital foramen with evident external plate, some part of
which is visible when skull is viewed from above ; teeth as in
Cricetus.

Remarhs. — The genus Cricctulus contains about a dozen
described species, whose distribution extends across the entii-e
central portion of Asia. One occurs in the Balkan Peninsula,
but the group is not otherwise represented in Europe west of
Russia.

CRICETULUS ATTICUS Nehring.

1882. Cricetus arenarius Winge, Vidensk. Middel. fra den naturh. Poren.

i Kj0benhavn, 4th ser., iii (1881), p. 31.
1902. Gricetulus atticus Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 3, January, 1902.
1910. Gricetulus atticiis Trouessart, Fauna Mamm. d'Europe, p. 165.

Type locality. — Pentelikon, Attica, Greece.

Geographical distribution. — Greece ; limits of range not
known, but apparently confined to the eastern portion of the
peninsula.

Diagnosis. — Similar to Cricetulus phseus of Asia Minor but
smaller, the condylobasal length of skull about 25 mm. nistead
of about 27 mm., hind foot about 13 '6 mm. instead of 15 mm.
Skull witli brain-case somewhat narrower than in C. phseus. In
the only adult examined the underparts are pale buff instead of
white. Recognizable among European murines by the short tail,
pale greyish colour and large ears. (In the pallid members of
the Pitymys ibericus group, which are of similar size and general
appearance, the ears are nearly concealed in the fur.)

Colour. — Upper parts a light grey approaching the grey
No. 10 of Ridgway but with a slight smoky cast, faintly tinged
with ecru-drab on head and suffused with light buff on rump
and buttocks, the hairs everywhere with dark tips, these incon-
spicuous except on crown and along middle of posterior half of
back, where they produce a slight though evident clouding of

•2 o

594

t:^

blackish. The individual hairs are slate-grey at base, those of
the underfur with a sub-terminal band of grey No. 10 about
2 mm. in width and a minute blackish tip, the longer hairs
blackish throughout. On rump and outer surface of hind legs
the pale gi'ey is replaced by light bull". Sides of body and outer
surface of fore legs white, the line of demarcation well defined
though not remarkably so, and extending along lower portion of
cheeks an(4 over base of whiskers to sides of muzzle, the lips thus
entirely white and sharply defined against the narrow grey
muzzle. A veiy narrow black eye-ring. Ears blackish, frosted
with greyish white, not much contrasted with surrounding parts.
Underparts in one specimen white, in another pale cream-bufF
rather sharply defined from the narrow white area along sides.
Feet butly white, including hairy portion of sole. Tail light
cream-buff', the upper surface sprinkled with blackish hairs along
median region.

Shill. — Apart from its much smaller size (approximately as
in Apodemus flavicoUis) the skull
differs from that of Cricetus cricetus
in its generally less modified, more
murine appearance. Dorsal profile
slightly and evenly convex through-
out. Brain-case relatively narrower
and rostrum relatively broader than
in Apodemus flavicoUis and A. sylva-
ticus, but these peculiarities not
carried to the same extreme as in
Cricetus cricetus. Brain-case dis-
tinctly longer than broad, slightly
wider than deep, smoothly rounded
off^ at sides, the occiput scarcely
oblique, so that condyles are not
visible when skull is viewed from
above ; the general form of brain-
case essentially as in Apodemus sylva-
ticus but narrower and deeper, and
with condyles more projecting. Interparietal about as in
A. sylvaticus, but more convex in front and more nearly straight
behind. Interorbital region about as wide as rostrum, its sur-
face with faintly indicated median longitudinal groove, its edges
slightly ridged. Rostrum a little less than half as wide as brain-
case, distinctly more robust than in Apodemus sylvaticus, but
without any special peculiarities of form. Nasals tapering
gradually backward, squarely truncate posteriorly, much exceeded
by nasal branches of premaxillaries. Zygomata slender, not very
widely spreading, their middle portions approximately parallel.
Auditory bullae moderately large, essentially as in Apodemus
sylvaticus. Mesopterygoid space parallel-sided, about three
times as long as wide, its anterior termination nearly in line

Fig. 117.
Cricetulus atticus. Nat. size.

CRICETULUS

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596

RODENTIA

with posterior border of vi\ the hamulars not diverging pos-
teriorly. Incisive foramina moderately long and narrow, much
as in Cricetus cricetus, the width of both together about one-third
length, their posterior border extending to within about 0 • 5 mm.
of level of tirst molar. Mandible resembling that of Cricetus
cricetus in the strongly bowed ramus and in the length and dis-
tinctness of the three main processes, the coronoid process larger
and more approximating to size and form of angular process than
in Ajpodemus. Outer surface of angular process distinctly convex
laterally, not concave as in Cricetus cricetus.

Teeth. — Except for their much smaller size the teeth are
essentially like those of Cricetus cricetus both in structure and in
length of tooth-row relatively to diastema and to width of palate.
Neither incisors nor molars show any tendency toward the
peculiarities assumed in Mesocricetus.

Measurements. — External measurements of an adult female
from Kephissia, near Athens : head and body, 87 ; tail, 22 ;
hind foot, 16. Eor cranial measurements see Table, p. 595.

Specimens examined. — Three, all from the vicinity of Athens.

Semarks. — Though nearly related to the Cricetulus pJiseus of
Asia Minor, the Grecian animal appears to be sufficiently distin-
guished by its smaller size.

2?. Athens, Greece. (C. Moita^.) Hon. N. C. Roths- 8.10.2.47-48.

child (p).

Genus CRICETUS Leske.

1779. Cricetus Leske, Anfangsgruude der Naturgesch., i, p. 168.

1799. Hamster Lac^pede, Tabl. des divisions ordres et genres des Mamm.,

p. 10 (Hamster nigricans Lac^pede = Miis cricetus Liunseus).
1859. Cricetus Blasius, Saugethiere Deutschlands, p. 305.
1873. Heliomys Gray, Ann. and Mag. Nat. Hist., 4th ser., xii, p. 417,

November, 1873 (H. jeudii = Cricetus cricetus).

Type species. — Mus cricetus Linnaeus (by tautonymy).

Geographical distribution. — Central Europe, from Belgium
and northern France eastward into Russia and Asia Minor.
Eastern limit of range not known.

Characters. — General form heavy and thick set, not murine ;
tail short but evident, slightly longer than hind foot ; mammse,
8 ; skull not murine in appearance, the interparietal much
reduced, the brain-case with two conspicuous, nearly parallel
ridges continued forward through interorbital region ; occiput
obliquely truncate ; rostrum much more than half as broad as
brain-case ; ectopterygoid fossa short, deep, moderately pit-like ;
infraorbital foramen with well developed external plate ; enamel
pattern characterized by presence of noticeable supplemental
median loops or islands in maxillary teeth.

CEICETUS 597

BemarJcs. — The genus Cricctus is too easily recognizable
among the European groups of rodents to require any special
comparisons. As at present understood it is represented by a
single species.

CRICETUS CRICETUS Linnseus.

(Synonymy under subspecies.)!

Geographical distribution. — Central Europe from Belgium and
northern France eastward into Russia and Asia Minor. Eastern
limit of range unknown.

Diagnosis. — Essential characters as in the genus ; size about
as in a large house rat, length of hind foot in adult 30 to 35 mm.,
condylobasal length of skull 43 to 51 mm. ; underparts entirely
black in strong contrast with sides.

External characters. — Form robust and heavy, suggesting a
house rat with short legs and rudimentary tail. Ear rather
large, orbicular, extending about to eye when laid forward ; a
slightly developed ridge behind meatus ; surface of ear well
clothed both externally and internally with fine hairs. Muzzle-
pad ill defined ; inner border of nostril swollen and projecting,
upper border notched ; a narrow median groove between nostrils
continuous below with broad naked and wrinkled median area
on upper lip. Cheek-pouches large, extending backward to
about level of ear, their openings entirely within the loosely
spreading lips. Feet moderately large, the claws stout, slightly
curved, those on front feet slightly exceeding those on hind feet
in size. Front foot with inner digit rudimentary, tubercular,
with minute flattened nail ; second digit slightly longer than
fifth and a little exceeded by fourth, this in turn by third, the
contrasts in length not very noticeable ; palm with five well
developed tubercles, of which the two posterior are largest, and
that at base of second toe smallest ; relatively to size of palm
the tubercles are large, leaving little space between them.
Hind foot with all five toes well developed, the innermost
extending to base of second, the outermost slightly longer, the
three middle digits sub-equal and longest ; tubercles six, all well
developed and of about equal size though rather small, not
arranged in pairs, the postero-internal lying distinctly behind
postero-external ; surface of sole hairy from heel to posterior
tubercles ; median area nearly to anterior tubercles distinctly
pubescent. Tail short but well developed, extending about to
tip of outstretched hind feet ; annulations ill-defined and
irregular, concealed by the thick covering of short hairs.
Mammse : p 2 - 2, z 2 - 2 = 8.

Colour. — Upper parts a uniform light yellowish brown or
brownish buff", the exact shade usually something between the
wood-brown and ochraceous-buff' of Ridgway though occasionally
with a russet or yellowish tinge, particularly on flanks, rump

598 RODENTIA

and anal region ; median dorsal region faintly clouded by a
sprinkling of black-tipped hairs ; side of muzzle, side of face
around and below eye, and a similar area around and below ear
tinged with tawny or dull ochraceous in rather noticeable
contrast with crown and back ; ear tawny edged with white, a
conspicuous clear buff tuft just below and behind meatus ;
underparts (except chin and anal region), inner surface of hind
leg and entire fore leg black, somewhat dulled by the slaty
under colour which appears at surface when hairs are worn or
disarranged ; between the black and brown areas lie the
following markings in clear, light buff: (1) an area including
front of muzzle, upper lip and chin and extending backward to
level of ear in front of which it rises as a narrow band nearly
separating anterior and posterior tawny areas, (2) a large
roundish area on side of neck just in front of fore leg, (3) a
similar area extending backward from axilla and separated from
that on neck by a dusky upward prolongation of black of leg,
and (4) a small spot (about equal to that at base of ear) at front
of thigh, this last sometimes obsolete or absent ; feet buffy
white ; tail dull yellowish brown, sometimes whitening at tip.

Sl-ull. — The skull is robust and heavy, about the size of that
of a large house rat. Dorsal profile nearly flat, but sloping
gradually and inconspicuously forward from lachrpnal region
and abruptly though not very conspicuously convex over anterior
third of nasals ; ventral profile essentially straight from behind
incisors to lower surface of auditory bulla, nowhere parallel with
dorsal profile, but nowhere conspicuously contrasted with it.
Brain-case roundish or somewhat diamond-shaped in genei-al
outline when viewed from above (when of latter form with
ends truncate at lambdal and interorbital regions) ; most
of the surface of bram-case is formed by the broad temporal
areas of muscle attachment, between which lies the narrow flat
median region continvied backward from interorbital trough,
narrower anteriorly than posteriorly, and bounded posteriorly by
that portion of lambdoid crest which lies along posterior border
of small, triangular or crescentic interparietal. No true sagittal
crest, though flat median area of brain-case becomes narrow
and crest-like in old individuals. Lambdal crest well developed,
with strong median concavity and a lateral convexity at each
side. Occipital regior. so obliquely truncate that the condyles
and entire upper surface of supraoccipital, from foramen magnum
to lambdal region, are visible when skull is viewed from above.
Occiput broadly arched when viewed from behind, a little
flattened above and at the sides, its height slightly more than
half mastoid width ; mastoid processes thick and short, their
lower border slightly above level of lower lip of foramen
magnum ; paroccipital processes short but well developed,
extending a little below lip of foramen magnum. Base of
brain-case with no special features, the basioccipital sloping

CRICETUS 599

noticeably upward posteriorly, its anterior half with median
groove, the backward extension of that occupying most of
basisphenoid. Auditory bulla rather large, evenly inflated, not
in contact with paroccipital process, its greatest diameter
oblique ; meatus large, without tube or lip ; a distinct depression
near middle of bulla parallel with meatus ; greatest diameter of
bulla about three times anterior width of basioccipital. Inter-
orbital region narrow, with distinct median furrow and raised
edges, the margin slightly but evidently angled posteriorly ;

Fig. 118.
Crwetus cricetus. Nat. size.

behind angles the ridges and furrow pass gradually into the
narrowest anterior portion of flat median area of brain-case.
Zygoma heavy but without special features, very gradually
spreading, so that greatest zygomatic breadth is at front of
glenoid region ; anteorbital foramen wide above, narrow and
slit-like below, the well developed external plate of canal plainly
visible when skull is viewed from above, its anterior border
nearly perpendicular. Rostrum very heavy, its greatest width
at about middle, decidedly wider than interorbital region and
more than half as wide as brain-case ; width at middle decidedly

600 RODENTIA

greater than depth at same region, extreme anterior portion
much narrower ; nasals gradually narrowing from before back-
ward, their posterior extremity bluntly pointed, slightly exceeded
by nasal ioranches of premaxillaries. Incisive foramina long and
narrow, parallel-sided, the width of the two together a little
more than one-half length ; anteriorly they extend to within
about 4 mm. of alveoli of incisors, ^josteriorly to within about
3 ram. of aveoli of molars. Bony palate with no special features ;
a faintly indicated median ridge continuous with septum between
incisive foramina ; mesopterygoid fossa about three times as long
as wide, its broadly rounded anterior extremity scarcely extending
to level of alveolus of vP, the plate-like, ill-defined hamulars in
contact with auditory bullfe ; ectopterygoids well developed,
lying, except at their anterior extremity, at a higher (more
clorsal) level than the pterygoids. Mandible slender, the ramus
strongly curved ; coronoid process rising decidedly above condyle,
conspicuously curved backward ; angular process long, curved
upward and outward, its upper margin with a well developed
thickening or supplemental outer plate.

Teeth. — Upper incisor with root extending to infraorbital
foramen, at front of which it produces a noticable swelling, the

Fig. 119.
Cricetus cricetus. Teeth X 0. a, tinworn ; h, slightly worn.

shaft slightly and somewhat obliquely compressed, the antero-
posterior diameter a little exceeding transverse diameter, the
anterior face broadly rounded, decidedly wider than posterior
face, that of the two teeth essentially in line with each other ;
cutting edge sloping upward toward the middle, the two edges
together forming a broad but evident angle ; lower incisor with
root extending nearly to base of coronoid process and forming an
evident tubercle on outer side of mandible slightly above level of
grinding surface of molars ; shaft oval in cross section, the inner
side somewhat flattened, the antero-posterior diameter about
double transverse diameter, the anterior and posterior faces

CRICETUS 601

narrowly and equally rounded. Molar rows about half as long as

diastema in fully adult skulls, relatively longer in immature

individuals ; anteriorly the rows diverge noticeably, owing to

somewhat oblique position of wi\ so that length of tooth-row is

only equal to greatest width of palate and alveolus of one side.

Anterior upper molar with four or sometimes five roots and six

tubercles, its crown about as large as m"^ and first pair of

tubercles of m^ ; the two anterior slightly smaller than the

others ; outer tubercles in line with each other and parallel to

main axis of crown, antero-internal tubercle somewhat displaced

outward so that the line of the three inner tubercles curves

outward in front, adding to the oblique appearance of the tooth ;

a deep pit in median line between each pair of tubercles, the pits

at a certain stage of wear assuming the form of enamel islands,

or of re-entrant loops extending obliquely backward from base of

outer tubercles and later disappearing ; at the stage in which the

pits have become loops the crown of the tooth has an enamel

pattern consisting of six triangles arranged in opposite pairs but

nearly closed by the backward extension of the outer re-entrant

angles ; at the later stage when the last trace of the pits has

disappeared, the bases of the triangles open broadly into each

other, forming three transverse loops nearly divided by the equal

external and internal re-entrant angles. Second upper molar

4-rooted, the crown with four tubercles resembling the posterior

four tubercles of first ; anterior border of crown with narrow

transverse ridge, best developed on outer side though scarcely

rising above level of extreme base of anterior tubercles. Third

upper molar 3-rooted, slightly more than half as large as second,

the two anterior tubercles normal in form though slightly reduced

in size, the two posterior tubercles small and confluent, the outer

more reduced than the inner, but traces of all the elements of

the tooth present ; anterior ridge well developed on outer side of

middle but obsolete or absent on inner side. Mandibular molars

resembling the corresponding upper teeth in general form and

structure, but m, i^elatively smaller and iiig relatively larger,

the tubercles slightly but evidently alternating, the inner more

anterior in position than the outer, and deepest re-entrant angles

on inner side instead of on outer side, the inner extremities of

the angles at no stage so distinctly isolated as pits. First lower

molar at all ages with six nearly closed triangles, distorted in

unworn teeth by the cusp-like elevation of their outer walls ;

anterior pair of cusps noticeably smaller than the others but

essentially in line. Second and third lower molars like posterior

four cusps of first, the crown of m^ nearly as large as that of nh,,

its posterior pair of cusps very little reduced ; anterior border of

each tooth with low but evident ridge on outer side.

RemarJcs.—Cricetus cricetus difi:ers conspicuously from all
other European rodents in its unusual colour pattern with
under parts entirely black. Four local forms have been described.

602 KODENTIA

Three of these occur west of Russia. Their status is at present
by no means clearly understood.

KEY TO THE EUROPEAN RACES OF CBICETUS CRICETUS.

Skull large, the condylobasal length about 50 mm.

(Central Germany eastward into Hungary) C. c. cricetus, p. 602.

Skull moderate, the condylobasal length about 45 mm.

Occipital portion of brain-case high, its truncation

moderately oblique as in typical race (Belgium

and western Germany) C. c. canesceru, p. 603.

Occipital portion of brain-case low, its truncation

unusually oblique (Roumania) C. c. nehringi, p. 605.

Cricetus cricetus cricetus Linntt-us.

1758. [Mus] cricetus Linnseus, Syst. Nat., i, 10th ed., p. 60, Germany.
1792. M[tis] Cricetus germanicus Kerr, Anim. King., p. 243 {Cricetus

germanicns in Syst. Catal., inserted between pp. 32 and 33)

Germany.
1799. Hamster nigricans Lacepede, Tabl. des division, ordres et genres des

Mamm., p. 10 (Renaming of Mus cricetus).
1801. J/[7(s] ciricetus'] fulvns Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 1010 (Thiiringen, Germany).
1808. Cricetus vulgaris Geoflroy, Catal. Mammif. du Mus. Nat. d'Hist.

Nat., p. 196, northern and eastern Europe (Renaming of

Mus cricetus).
1811. Cricetus frutnentariics Pallas, Zoogr. Rosso. -Asiat., p. 161 (Renaming

of Mas cricetus).
1857. Cricetus frumentarius Blasius, Saugethiere Deutschlands, p. 306.
1867. {Cricetus vulgaris'] varius Pitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 98

(Europe).
1867. [^Cricetus vulgarus] albus Pitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 98

(Germany).
1867. [Cricetus vulgaris^ niger Pitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 98

(Austria, Hungary, Germany).
1873. Heliomys jeudii Gray, Ann. and Mag. Nat. Hist., 4th ser., xii, p. 417,

November, 1873 (locality unknown).
1894. Cricetus cricetus Dahl, Die Heimat, iv, p. 130, June, 1894.
1906. Crlicetus] vulgaris niger Simroth, Biol. Centralblatt, xxvi, p. 337,

June 1, 1906. Valley of the Saale, Germany.
1910. Cricetus cricetus and C. cricetus niger Trouessart, Fauue Mamm.

d'Europe, p. 159.

Tijpe locality. — Germany.

Geographical distribution. — Locally distributed through central
Germany. Exact limits of distribution not known, and apparently
undergoing rapid change.*

Diagnosis. — Skull large and heavy, the condylobasal length

* For detailed account of local distribution in Germany see Nehring,
Wiegmann's Archiv fiir Naturgeschichte, 1904, pp. 15-32, pi. iii, and
Sitz.-Ber. Gesellsch. Naturforsch. Freunde, Berlin, 1899, pp. 3-4.

CEICETOS

603

about 50 mm. ; brain-case deep posteriorly, the truncation of
occipital region moderately oblique, the angle of the slope as
referred to plain of base of brain-case about 70^.

Measurements. — Adult male from the neighbourhood of
Magdeburg, Germany : head and body, 260 ; tail, 60 ; hind foot
(dry), 36 ; ear from meatus about 27. A second adult male
from the same locality : hind foot (dry), 36. For cranial
measurements see Table, p. 604.

Specimens examined.- — ^Nineteen, from the following localities : —

History unknown, 1 skull (type of Heliomys jeudii Gray).

Geemany: No exact locality, 1; Ingelheim, Rheinhesseu, 6; Ober-

lausitz, Saxony, 1 ; Lotzen, Saxony, 1 (U.S.N.M.) ; near Magdeburg,

Saxony, 6 ; Saxony, no exact locality, 1 (U.S.N.]\I.) ; Strassburg, 1

(Lataste).

Austria-Hungary: Molna Szecsod, Hungary, 1 (U.S.N.M.), an

immature individual, perhaps not true cricetiis.

1. Germany. {2foschler.) E. R. Alston (p). 79. 9. 25. 35.

6 6. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 38-43.

6. Oberlausitz, Saxony. Dr. E. Hamilton (p). 97. 12. 4. 33.

3, 1 juv. Magdeburg. Dr. W. Wolterstorff 92. 12. 1. 24-27.

skull. Lidth de Jeude Col- 67. 4. 12. 314.

lection.
(Type of Heliomys jciklii Gray.)

Cricetus cricetus canescens Nehring.

1899. Cricetus vulgaris var. canescens Nehring, Sitz.-Ber., Gesellsch.

Naturforsch. Freunde, Berlin, p. 1.
1910. Cricetus cricetus canescens Trouessart, Faune Mamm. d'Europe,

p. 160.

Type locality. — Near Fexhe-Slins, banks of the Maas,
Belgium.

Geographical distribution. — Eastern Belgium and north-
western Germany ; probably north-eastern France also.*

Diagnosis. — Similar to G. cricetus cricetus, but skull noticeably
smaller, its condylobasal length about 45 mm. ; form of brain-
case as in the typical race.

Measurements. — External measurements of two adult males
from Tirlemont, Liege, Belgium : head and body, 212 and 218 ;
tail, 50 and 47; hind foot, 35 and 35 (dry, 33 and 33 -4);
ear, 32 and 28. For cranial measurements see Table, p. 604.

Specimens examined. — Four, from the following localities : —
Belgium : Tirlemont, Liege, 3.
Germany : Brunswick, 1 (Field).

Remarks. — The status of this race is not satisfactory. The
Belgian specimens which I have seen show none of the

* I have seen no French specimens, but the form will probably prove
to be canescens. For account of recent changes in distribution in France
see Trouessart, Faune Mamm. d'Europe, p. 159.

604

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MESOCRICETUS 605

peculiarities in colour described by Nehring, but agree perfectly
in this resjDect with those from central Germany. The small
size of their skulls, however, is too pronounced a character to
permit the name canescens to be placed in the synonymy of true
C7'iccius.

2 6,1 al. Tii-lemont, Belgium. Rev. L. Warnau (p). 8. 11. 23. 1-3.

Cricetus cricetus nehringi Matschie.

1901. Cricetus neJiringi Matschie, Sitz.-Ber., Gesellsch. Naturforsch.

Freunde, Berlin, p. 232.
1910. Cricetus cricetus nehringi Trouessart, Faune Mamm. d'Europe,

p. 160.

Type locality. — Described from specimens from Slobosia,
Cernavoda, and Barza, Roumania. The first may be assumed to
be the type locality.

GeograpJdcal distribution. — Known only from a few localities'
in Roumania.

Diagnosis. — Externally as in C. cricetus cricetus, but tail
apparently shorter ; skull small, as in C. c. canescens, the brain-
case noticeably depressed jDosteriorly and truncation of occipital
region more oblique than in the other races, the angle of the
slope as referred to plain of base of brain-case about 60°.

Colour. — The colour does not appear to differ appreciably
from that of true C. cricetus, though in the one adult examined
the light thigh spot is barely indicated by a few scattered
whitish hairs.

Measurements. — ^External measurements of adult (from well-
made skin) : head and body, 230 ; tail, 28 ; hind foot, 30 • 6 ; ear
from meatus, about 22. For cranial measurements see Table
opposite.

Specimen examined. — An adult from Bucharest, Roumauia (U.S.N.M.).

Genus MESOCRICETUS Nehring.

1898. Mesocricetus Nehring, Zool. Anzeiger, xxi, p. 494, September 5,
1898 (Sub-genus of Cricetus).

1898. Semicricetus Nehring, Zool. Anzeiger, xxi, p. 494, footnote (Alterna-
tive for Mesocricetus).

1898. Mediocricetus Nehring, Zool. Anzeiger, xxi, p. 494, footnote (Alter-
native for Mesocricetus).

1900. Mesocricetus Satunin, Zool. Anzeiger, xxiii, p. 301, May 28, 1900
(genus).

Type species. — Cricetus nigricans Brandt.

Geographical distribution. — Caspian region and Asia Minor,
west into eastern Roumania and Bulgaria. Limits of range not
known.

Diagnosis. — Externally as in Cricetus, but tail shorter than

606 RODKNTIA

hind foot, nearly concealed in fur, and mamruEe 14 to 16 * instead
of 8 ; skull resembling that of Cricetus in general features, but
anteorbital foramen without external plate, and flattened area on
dorsal surface of brain-case broad, its outline somewhat truncate-
diamond shaped ; ectopterygoid fossa short, deep, and con-
spicuously pit-like ; teeth not essentially different from those of
Cricdus, though showing a higher degree of specialization.

Remarks. — This well characterized genus contains six described
forms, one of which occurs in eastern Europe. It is the most
aberrant of tlae Old World group of Cricetinse.

MESOCRICETUS NEWTONI Nehring.

1898. Cricetus nciotoni Nehring, Zool. Anzeiger, xxi, p. 329, May 16, 1898.
1898. Cricetus (Mcsocricetus) neivtoni Nehring, Wiegmann's Archiv fiir
Naturgeschichte, 1898, i, p. 386, December, 1898.

1900. Mlesocricetus'] newtoni Satunin, Zool. Anzeiger, xxiii, p. 301, IMay 28,

1900.
1910. Mcsocricetus newtoni Trouessart, Faune Mamin. d'Europe, p. 162.

Type locality. — Schumla, eastern Bulgaria.

Geographical distribution. — Eastern portions of Bulgaria and
Roumania. Exact limits of range not known.

Diagnosis.— ^ize smaller than in Cricetus cricetus (hind foot
less than 20 mm., condylobasal length of skull less than 40 mm.) ;
black of underparts not extending behind fore legs ; a blackish
area on crown continued backwai'd as median line to between
shoulders ; an oblique black stripe from side of shoulder to
lower margin of cheek.

External characters. — General form as in Cricetus cricetus, but
tail nearly concealed in fur, its surface without trace of scales or
annulations, and noticeably more hairy below than above. Ear
as in Cricetus, extending to eye when laid forward. Feet as in
Cricetus, but pads on sole more crowded, the postero-internal
and postero-external at same level, and the others showing a
distinct tendency to become arranged in pairs. Sole almost
naked throughout, the portion behind tubercles slightly pubescent.
Mammse : 7-7 or 8-8 (in skin No. 122097, U.S.N.M.,
apparently p 4 — 4, -i 4 — 4).

Colour. — Upper parts a light drab brown, perhaps most
nearly the broccoli-brown of Ridgway, but paler and with a
slight wood-brown cast, especially along middle of back, every-
where finely and inconspicuously sprinkled with blackish hairs.
Underparts cream-buff, clearer and more yellowish along sides of
body, neck and cheeks, duller and somewhat obscured by the
smoke-grey under colour on belly ; region fi'om interramia to
between fore legs and covering antero-internal surface of leg
nearly to wrist blackish brown, not so dark as the corresponding

* See Nehring, Sitz.-Ber. Gesellsch. Naturforsch, Frennde, Berlin,

1901, p. 155.

MESOCRICETUS

607

region in Cricetus cricetus ; chin dull whitish ; upper lip cream-
buff. Face, muzzle and most oE region between ear and eye like
back but paler ; roundish area covering most of cheek below eye
and extending forward to base of whiskers strongly tinged with
dull ochraceous ; minute hairs on eyelids black ; behind and
somewhat below cheek spot is a clear cream-buff area continuous
with that of side of neck ; ear silvery grey in noticeable contrast
with surrounding parts ; region behind ear strongly tinged with
buff; crown between ears and slightly in front of them blackish,*
this area continued backward as a fairly well defined median
stripe about 3 mm. in width to a little behind shoulders ; a
better defined black stripe about 6 mm. wide extends obliquely
downward and forward from front of shoulder to about angle of
jaw ; this stripe thrown into strong relief by clear cream-buff
area lying between it and the downward extension of somewhat
darkened body colour toward upper portion of leg ; feet and tail
buffy white. Immature individuals are lighter and less tinged
with wood-brown above, and less buffy on sides ; the dark
mai'kings essentially as in the adults.

Skull. — Apart from its smaller size the skull bears a general
likeness to that of Cricetus cricetus ; profiles essentially as in the

FIG. 120.
Mesocricetus newtoni.

Nat. size

larger animal. Brain-case shorter and deeper than in Cricetus
cricetus, its upper surface with much larger flattened area, the
border of which bends abruptly outward at front of parietal and

* This blackish marking ahnost exactly corresponds in size, form and
position to the flattened area on dorsal surface of brain-case.

608 RODENTIA

continues nearly parallel with outer margin of the bone to lambclal
suture, the two borders together producing a diamond-shaped
figure. Interparietal with antero-posterior diameter relatively
greater and tranverse diameter lelatively less than in Cricetus
cricetus. Interorbital region much narrower than rostrum, with
high lateral ridges and well marked median groove. Zygoma not
peculiar in general form, gradually spreading so that greatest
zygomatic breadth is at glenoid level. Anterior base of zygoma
peculiar in the complete absence of the forwardly extending
plate forming outer wall of infraorbital foramen in Cricetus, so
that when viewed from above the anterior margin forms an
unbroken curve to maxillo-premaxillary suture. The absence of
the outer plate causes the foramen to assume an oval outline
somewhat flattened on inner side. Rostrum not tapering
anteriorly, the width of the region immediately in front of
infraorbital foramina not increased by any evident swellings
over roots of incisors. Incisive foramina relatively shorter and
broader than in Cricetus cricetus, the outer margins slightly but
evidently spreading in front of suture, the posterior margins
extending to level of posterior margins of infraorbital foramina.
Palate relatively narrower than in Cricetus cricetus, but with no
special peculiarities of structure. Mandible with coronoid process
relatively longer than in Cricetus cricetus, its apex almost over-
hanging articular surface.

Teeth. — ^^Upper incisor not producing any noticeable swelling
at front of infraorbital foramen, the shaft narrower internally
than externally, its transverse diameter and greatest antero-
posterior diameter equal, its anterior face nearly flat, sloping
conspicuously inward, so that those of the two teeth come
together at a noticeable angle ; cutting edge very oblique.
Lower incisor with shaft flattened in front and on inner side, the
outer and posterior borders evenly rounded, the antero-posterior ^
diameter scarcely exceeding greatest transverse diameter. Molar
rows about two-thirds as long as diastema ; anteriorly the rows
diverge so slightly and the palate is so narrow that length of
tooth-row is distinctly greater than width of palate, including
both alveoli. Form, relative size, and general structure of molars
both above and below as in Cricetus cricetus, but backward pro-
longation of outer re-entrant angles in upper teeth, apparently
at all stages isolated as pits or islands, and width of re-entrant
angles in lower teeth greater relatively to area of tubercles.

Measurements. — Head and body about 180 ; tail about 12
(well-made skins) ; hind foot in three adults, 19 mm. , ear from
meatus about 15. For cranial measurements see Table opposite.

Speciviens examined. — Nine, from the following localities : —
Roumania: Malcoci, Dobrudscha, 7 (B.M. and U.S.N.M.).
Bulgaria : Melschka, near Bustchuk, 2 (Andersen).

BemarTis. — Apart frgm the generic characters, its peculiar

MESOCRICETUS

609

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610 RODENTIA

coloui' pattern will at once distinguish this animal from all other
European rodents.

1. Malcoci, Dobrudscha, Purchased (Pruliere) (p). 86. 4. 2. 5.
Roumania.

1. Rustchuk, Bulgaria. Dr. Knud Andersen (p). G. 5. 17. 1.

Sub-Family MICROTIN^.
1906. Microtinx Miller, Nortli Amer. Fauna, No. 12, p. S.

Geographical distrihution. — Northern portion of both hemi-
spheres, south to Mexico, northern India, and the Mediterranean
coast of Europe

Characters. — Murine rodents with definitely prismatic,
hypsodont or rootless, flat-crowned molars.

BemarJcs. — The sub-family Microtinse contains about thirty
groups, genera or sub-genera, the status of which is still
imperfectly understood. These fall naturally into three main
sub-divisions, the Lemmi, characterized by the shortness of the
lower incisor, the root of which remains on the lingual side of
the molar alveoli, the Microti, containing the true voles, and the
ElJohii, the last highly modified for underground life. The
Lemmi and Microti are represented in western Europe, the first
by two living genera, the second by four. Fossil and sub-fossil
remains of membei-s of the sub- family are found throughout
this region, some of them representing genera now extinct.

KEY TO THE EUROPEAN GENERA OF MICBOTINJE.
Root of lower incisor lying entirely on lingual side of
molar teeth (Lemmi).
General form vole-like ; feet slender (normal) ; palms
and soles with well developed functional (normal)

tubercles Mijopus, p. 611.

Ger eral form short and heavy ; feet broad, the
metapodials shortened and ungual phalanges
lengthened ; palms and soles with rudimentary,
non-functional tubercles concealed by a dense

growth of stiffened hairs Lcmmus, p. 614.

Root of lower incisor extending to outer side of molar
teeth {Microti).
Bony palate terminating posteriorly in a thin-edged

shelf; molars rooted in adult Evoioimjs, ]). 623.

Bony palate terminating posteriorly in a sloping
median ridge and two lateral pits.
Sole with six well developed tubercles ; mammse, 8. Microtus, p. 658.
Third upper molar with three re-entrant angles

on inner side Microtus, p- 659.

Third upper molar with two re-entrant angles on

inner side Chionomys, p. 712.

Sole with five well developed tubercles ; mammse,
8 or 4.
First lower molar with first outer and first
inner triangle communicating ; mammse, 4 ;
general form modified for underground life... Pitymys, p. 752.
First lower molar with first outer and first inner
triangle not communicating ; mammas, 8 ;
general form modified for aquatic (rarely
subterranean) life Arvicola, p. 723.

611

Genus MYOPUS Miller.

1910. Myopus Miller, Smithsonian IMiscell. Coll., Lii, p. 497, January 12,
1910.

Type species. — Myodes schisticolor Lilljeborg.

Geographical distribution. — Southern Norway and central
Sweden ; eastward into Finland and perhaps much further.
Limits of range not known.

Characters. — Skull and teeth as in Lnumus ; general form
voledike, though tail rather short ; feet
slender, the palm and sole with fully
developed, functional tubercles and no
unusual growth of hair ; metacarpals of
third and fourth fingers slightly longer
than phalanges ; ungual phalanges of
raanus normal, much shorter than first
and second phalanges combined, the
claws not enlarged ; ear well developed
though small, with distinct meatal valve.

RemarJcs. — The genus Myopus is
characterized by the combination of the o, „i 4. ,f ? + p

y .11 Skeleton of fore foot of

skull and teeth or Lemmus with the Lemmus (a) a,nd Myopus (b).

general body-form and foot structure of

the voles. It is, therefore, less specialized than the other Old

World lemmings, representing a stage of development equivalent

to that of the American Synaptomys. Only one species is

known.

MYOPUS SCHISTICOLOR Lilljeborg.

1844. Myodes scliisticolor Lilljeborg, Ofversigfc af Kongl. Vetenskaps-Akad,
Forhandl., Stockholm, i, p. 33, March 20, 1844.

1910. Lemmus schisticolor Trouessart, Paune Mamm. d'Europe, p. 199.

1911. Myopus schisticolor Collett, Norges Pattedyr, Hefte 3, p. 130

March, 1911.

Type locality. — ^Near Lillehammer at north end of Mjosen,
Gudbrandsdal, Norway.

Geographical distribution. — Fir forests of southern Norway
and central Sweden ; eastward into Finland and perhaps much
further.* Exact limits of range not known.

D/ar/nos/s.— General characters as in the genus ; size
essentially as in Microtus agrestis (head and body about 100 mm. ;
condylobasal length of skull, 25 mm.) ; tail short, extending
barely as far as outstretched hind foot ; colour slaty grey with
reddish brown dorsal patch.

* Recorded by Middendorff from Ajan, west coast of the Okhotsk Sea
(Sibirische Reise, ii, Siiugethiere, Vogel und Amphibien, p. 108, 1853).
The validity of this record questioned by Allen (Bull. Amer. Mus. Nat.
Hist., XIX, pp. 153, 154, March 31, 1903).

2 K 2

612 RODENTIA

External characters. — Form vole-like, slightly more robust
than in Microtus agrestls, the legs not shortened and feet not
enlarged as in Lemmiis lemmus. Head rather large, the anterior
portion blunt ; ear rounded, well developed, but nearly con-
cealed in the fur, reaching to within about 2 mm. of eye when
laid forward, the meatus with a distinct though not very high,
broadly triangular valve, both surfaces of ear thickly covered
with rather short hairs, a noticeable tuft of longer hairs at outer
base of meatal valve ; muzzle pad slightly developed, the upper
and inner margins of nostrils strongly projecting ; upper lip with
deep median cleft. Feet slender, strictly vole-like in form and
relative size, the claws on hind feet slightly larger and less
compressed than those on fore feet. Fore foot with the inner
digit essentially as in Lemmus lemmus, its strongly compressed,
strap-shaped nail extending to base of second digit ; outer digit
as long as second and a little less than half as long as the sub-
equal second and third ; pads four, well developed, essentially as
in the smaller voles, except that that at base of thumb is absent,
and postero-external is relatively larger and more elongated, its
extremity reaching nearly to level of tip of thumb-nail when
the two are approximated ; surface of palm between pads
granular-tuberculate, naked with a few scattered minute hairs.
Hind foot at least as slender as that of Evotomys glareolus, its
five tubercles relatively larger and more crowded, occupying
much more, than half the surface of the region in which they
occur, its digits with no special peculiarities of size or form ; sole
densely hairy behind pads, naked between them except for
a sprinkling of very fine hairs, the naked surface conspicuously
granular-tuberculate ; main tubercles sub-equal in size and
essentially alike in form except that the two hinderinost are
somewhat smaller and more tei'ete than the others, these two
nearly in contact, the outer slightly in front of the inner. Tail
short, but not unusually robust, densely covered with hairs that
nearly conceal the narrow, ill-defined annulations ; pencil slender,
about one-half as long as vertebra. Mammae : p 2 — 2 ;
i 2 - 2 = 8.

Colour. — Except for the dorsal patch the entire animal is a
uniform dark grey approaching the slate-colour of Ridgway, the
upper parts with a slight steely effect due to the metallic, almost
silvery, appearance of the tips of the shorter hairs, and a less
evident dark "lining" produced by the longer wholly black
hairs ; a broad, rather ill-defined area on middle of back,
extending from shoulders to within about 15 mm. of base of tail,
a reddish brown intermediate between russet and rufous. Feet
and tail .slaty black, the hairs on under side of latter with a
strong silvery gloss.

Skull and teeth. — Apart from their noticeably smaller size the
skull and teeth do not differ essentially from those of Lemmus
lemmus. The general outline of the skull when viewed from

613

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614 RODENTIA

above is somewhat less broadened than in the larger animal, and
the emargination of posterior border of palate extends relatively
further forward : to level of last prism of iir, instead of to level
of first prism of m^, anteoi'bital foramen with slit-like lower
portion so shortened as to be practical^ absent.

In the few specimens examined the inner border of cutting
edge of upper incisor is less folded backward than in Lemmus
lemmuii, and the outer re-entrant angle of yn.^ is occasionally so
little developed that it fails to cut off a distinct triangle from
outer extremity of second transverse loop, a condition, rarely, if
ever, found in L. lemmus.

Measurements. — Adult (old) female from Eidsvold, Norway
(in alcohol) : head and body, 95 ; tail, 19 ; hind foot, 15 '4 ; ear
from meatus, 10. For cranial measurements see Table, p. 613.

Specimens examined. — Fifteen, from the following localities: —
Norway : Hurdalen, Gudbrandsdal, 1 ; Fluberg, Hadeland, 2 (B.M. and

U.S.N.M.) ; Eidsvold, Akershus, 2 ; Keen, Ohristiania, 1.

Sweden: Nerike, Wermland, 2 (U.S.N.M.); Upsala, 1 (U.S.N.M.);

Medelpad, W. Norrland, 4 ; no exact locality, 1 (U.S.N.M.) ; Dalarne, 1.

1. Hurdalen, Gudbrandsdal, Dr. R. CoUett (p). 84. 10. 31. 3.
Norway.

6 juv. Fluberg, Hadeland. Ohristiania Museum 98. 3. 1. 18.

(b).

2 ? al. Eidsvold, Akershus. Dr. R. Collett (p). 84. 10. 31. 1-2.

6 juv. Heen, Ohristiania. Ohristiania Museum 93. 3. 1. 19.

(E).

2c5juv. Medelpad, W. Norrland, Lord Lilford (p). 11.1.1.111-114.

2 ? juv. Sweden.
1 al. Dalarne. Stockholm Museum 90. 8. 1. 18.

(e).

Genus LEMMUS Link.

1795. Lemmus Link. Zool. Beytriige, i, pt. ii, p. 75. Type by tautonymy

Mas lemmus Linnaeus.
1811. Myodes Pallas, Zoogr. Rosso-Asiat., i, p. 172 (part).
1877. Myodes Ooues, Monogr. N. Amer. Rodentia, p. 237.
1896. Lemmus Miller, N. Amer. Fauna, No. 12, p. 36, July 23, 1896.

Type species. — Mus lemmus Linnoeus.

Geographical distribution. — Arctic region of both hemispheres ;
in Europe south to southern Norway.

Characters. — General foi'm noticeably heavier and more
robust than in the voles ; legs and tail short ; feet broad, the
palm and sole covered with a dense growth of stiffened hairs,
under which are concealed the rudimentary, functionless
tubercles ; metacarples of third and fourth fingers much shorter
than phalanges ; ungual phalanges of manus greatly enlarged,
slightly longer than first and second phalanges combined, the
claws enlarged but simple; ear well developed though small,

LEMMUS 615

without meatal valve ; skull very bi'oad and low, the zygoma
conspicuously expanded at middle, the mesopterygoid space short
but continued forward over (under when skull is examined from
below) posterior margin of palate ; lower incisor with root lying
entirely on lingual side of cheek-teeth ; molars rootless, the
enamel pattern characterized by great depth of outer re-entrant
angles in maxillary teeth, the points of these angles extending
practically to inner margin of crown ; vv^ with four transverse
loops and no closed triangles ; m.^ with three transverse loops and
one closed triangle.

Hemarks. — Except for the primitive condition of the lower
incisors characteristic of the lemmings as a group, the members
of the genus Lemmus are probably more specialized than those of
any of the related genera. In Dicrostonyx* the ear is more
reduced and the claws reach their maximum of development ;
but the skull and molars are distinctly more generalized in form
and structure. Two species of Lemmus are now known to
inhabit the Old World, while several other forms have been
described from North America. Only one occurs in Europe,
west of Russia.

LEMMUS LEMMUS Linnseus.

1758. [Mils] lemmus Linnaeus, Syst, Nat., i, 10th ed., p. 59 (Sweden).

1808. L[emmus'] lemmus Tiedemann, Zoologie, i, p. 474.

1820. Lemmus borealis Nilsson, Skand. Faun., i, p. 185 (Substitute for

le^nmus).
1822. Lemmus norwcgicus Desmarest, Mammalogie, pt. ii, p. 287 (Norway).
1895. Myodes lemmus CoUett, Christiania Vetenskabs, Selskabs Forhand-

linger, 1895, No. 3, p. 3.

* While no living member of the genus Dicrostonyx is now known to
occur in Europe west of the Gulf of Finland, sub-fossil remains are found
in many parts of central and western Europe associated with those of
Lemmus. Such specimens of the two genera may be distinguished as
follows : —

Lemmus. — Skull very broad and low ; zygomata greatly expanded at
middle ; temporal ridges early uniting to form a knife-like crest in inter-
orbital region ; postorbital process shelf-like ; auditory bullae very large,
their substance conspicuously spongy ; a large foramen below and behind
alveolus of ui.^ ; tooth-rows noticeably converging anteriorly ; upper molars
with outer re-entrant angles conspicuously deeper than inner ; m-, m^, and
?«3 with one or more transverse loops in addition to the terminal loop.

Dicrostonyx. —Skull moderately broad, not specially depressed ; zygo-
mata not greatly expanded at middle ; temporal ridges never uniting, the
interorbital region with a noticeable longitudinal furrow ; postorbital
process peg-like; auditory bullfe not unusually large, their substance scarcely
spongy ; no noticeable foramen below and behind alveolus of m^ ; tooth-
rows nearly parallel ; ux^per molars with outer and inner re-entrant angles
approximately equal ; m", ni^ and m^ without transverse loops in addition
to the terminal loop. Externally Dicrostonyx is distinguished from
Lemmus by the reduction of the ear to a low rim, the absence of any well
developed nail on thumb, and the bifurcate form of claws on third and
fourth fingers in winter.

616

RODENTIA

1896. L[cmmus'] lemmus Miller, North American Fauna, No. 12, p. 37,

July 23, 1896.
1910. Lemmus lemmus Trouessart, Faune I\Iamm. d'Europe, p. 198.

Type Zoca/iV?/.- -Mountains of Lappmark, Sweden.

Geographical distribution. — Northern Scandinavia and Finland,
east to the White Sea, south in the mountains of Norway to
Langfjeld, Christiansand," and in Sweden to northern Werm-
land.t During seasons of abnormal increase the animals wander
to the extreme south of Norway.

Diagnosis. — General character as in the genus ; size medium
(head and body about 13.5 to 145 mm. ; tail, 12 to 14 mm. ; hind
foot, 17 to 20 mm. ; condylobasal length of skull, 29 to 32 mm.) ;
colour yellowish brown, the head, neck and shoulders with a
conspicuous black mantle.

External characters. — Form robust, the head large and very
broad, appearing to be set directly on the shoulders, the legs
short and muscular, the tail very short and thick. Head broader
than deep, its greatest width nearly equal to that of body at
middle, the muzzle bluntly rounded ; ear short, concealed in the
fur, extending barely to eye when laid forward, its outline evenly
rounded, the meatus without trace of valve ; eyes rather small,
their distance apart about equal to that from inner canthus to
nostril ; muzzle pad obscure, the nostrils opening close to broad
short groove which divides narrow upper lip, their inner and
lower margins slightly projecting. Front leg short, very muscular,
the foot broad, with short, robust digits, the innermost of which
is very short, strongly compressed, and serving merely as a base
for the conspicuous strap-shaped nail 4 mm. in length by 3 mm.
wide ; the third is longest, and the fourth, second and fifth are
successively shorter ; palm covered with a dense mass of stiffened
hairs about 3 mm. in length, the tubercles represented by a
rudimentary pad at base of the two longest fingers ; claws
simple but large and robust, about 5 mm. or more in length
(frequently exceeding length of fingers), sharply pointed or blunt
according to condition of wear. Hind foot broad, with short,
strong digits, the three median of which are sub-equal and
longest, the innermost shortest and outermost intermediate ;
claws strong, simple, about 4 mm. in length ; sole densely hairy
like palm, the rudimentary, functionless tubercles completely
hidden ; on cutting away the hair four minute pads may be
detected, one at the base of the inner toe, one at the base of
second, the third between bases of third and fourth toes and the
fourth at base of fifth. Tail short, extending about to middle of
outstretched hind foot, its form somewhat club-shaped, the
diameter at base distinctly less than that just below tip ; hairs
rather short at base, nearl}' as long as vertebra? at tip where they

* Collett, Norges Pattedyr, p. 138, 1911.

t Lilljeborg, Sveriges och Norges Ryggradsdjur, i, p. 326, 1874.

LEMMUS

617

form a dense pencil ; annulations irregular but rather well
detined, about 0*5 mm. wide. Fur full and soft, the hairs of
general body-covering about 10 mm. in length, those of back
and rump increasing rather abruptly to 20 or 30 mm. Mammae :
p2-2,i2-2 = S.

Colour. — Underparts and sides bufF, paler (in some specimens
whitish) on throat and chin, yellower on cheeks and sides of
neck, darker and more ochraceous on belly ; upper parts black
anteriorly, ochraceous posteriorly, the exact tint of the latter
varying between the ochraceous, raw-sienna, and tawny of
Ridgway, the slate-grey under colour appearing irregularly at
surface, especially at sides of longer haired area, where there is
usually an ill-defined blackish stripe, especially noticeable in
summer ; a narrow ill-defined black median stripe may usually
be traced backward from posterior border of black area to middle
of back or to rump ; black area everywhere sharply defined, its
posterior border a short distance behind shoulders, its lateral
border extending almost horizontally along a line beginning at
nostril and passing just below eye and ear (leaving entire upper
lip buff) to side of shoulder, where it curves upward ; an
ochraceous bufty line, about 3 mm. wide, extends backward from
eye to region between ears where it abruptly widens to a
squarish patch, the two patches frequently uniting in median
line to form a conspicuous occipital area 20 mm. wide by
10 mm. long; extreme point of muzzle above naked pad pale
buff ; some trace of a huffy median line usually present along
middle of face ; ear blackish, the hairs immediately surrounding
it mostly tipped or annulated with buify ; feet light buffy with
a silvery gloss, the digits tinged with hair-brown ; claws light
horn-colour ; tail light buff, the upper surface usually sprinkled
with blackish hairs. There is not much seasonal change in
colour. Winter specimens tend to be more yellowish, summer
specimens more brown ; in the latter the median dark stripe on
middle of back reaches its extreme of distinctness, and the dark
border to ochraceous dorsal area is often rather conspicuous.
Newly born young are essentially similar to the adults, though
the buff and ochraceous are dull ; owing to the shortness of the
fur the colour pattern is very sharply defined.

SJmll. — In general form the skull is broad and depressed,
with rather slender rostrum and unusually heavy, abruptly
spreading zygomata. Dorsal profile essentially horizontal from
lambdoid region to anterior base of zygomata, the rostrum bent
abruptly downward at an angle of about 35° ; ventral profile
concave immediately behind incisors, then essentially parallel to
dorsal surface, but curving abruptly upward posteriorly from
middle of bulla to base of paroccipital process. Brain-case
broadly lyrate in outline when viewed from above, owing to
the noticeable constriction just in front of mastoid region, and
the abrupt posterior expansion over widest portion of bulliie ;

618

RODENTIA

postorbital process forming a narrow but evident shelf, extending
from base of zygoma to beyond middle of posterior border of orbit ;
interparietal large, sub-quadrate, its antero-posterior diameter
about two-thirds transverse diameter ; lambdoid crest rather high
and distinct laterally, but scarcely noticeable along posterior
margin of interparietal ; occiput abruptly truncate, nearly
perpendicular below, slightly rounded-off above, the condyles
and paroccipital processes just visible in dorsal view ; seen from
behind the occiput is low and broad, its width noticeably more
than twice height above foramen magnum ; paroccipital processes
projecting more backward and outward than downward, their
lowest point slightly below level of condyles ; floor of brain-case
with no special features, the basioccipital with a moderately
defined median longitudinal ridge, its width along basal suture

Fig. 122.
Letnmut, lemmus. Nat. size.

contained about 2^^ times in median length ; auditory bulla large
but not very highly inflated, the interior nearly filled by a
dense mass of spongy tissue, the ventral border about on level
with cutting surface of molars, the size chiefly due to extension
forward into ectopterygoid fossa nearly to level of posterior
border of molar, width of bulla at level of meatus greater than
width between outer surfaces of occipital condyles. Inter-
orbital region narrow, cylindrical, scarcely as wide as rostrum,
its dorsal surface concave between zygomatic roots, posteriorly
with well marked ridges, which early unite to form a short,
trenchant median crest. Zygoma very heavy anteriorly, broadly
expanded at middle, abruptly slender and weak posteriorly ; ante-
orbital foramen rather small, very narrow below, the plate forming
its outer border slightly developed at base, its anterior border

LEMMtS

61&

vertical. Nasal simple, squarely truncate posteriorly at level of
middle of zygomatic root, scarcely or not exceeded by the very
slender nasal branches of premaxillaries. Incisive foramina long
and very narrow, extending from about 1 • 5 mm. behind incissors
to within about 1 mm. of level of molar alveoli. Palate about as
wide as alveolus anteriorly, about twice as wide posteriorly ;
lateral grooves moderately well marked ; posterior emargination
extending nearly to level of front of m^, its median spine blunt
and short, but evident ; pterygoids short, the hamulars heavy,
curved outward posteriorly, the mesopterygoid space extending
over posterior margin of palate. Mandible rather slender, but
with conspicuously developed ridges for muscular attachment ;
posterior portion with a peculiar thin and sunken aspect, due
partly to the absence of the incisor root from this' region, and
partly to the high masseter ridge and the conspicuous expansion
outward of under surface of large angular process ; articular
process very slender and weak, much compressed ; coronoid
process broad at base, but narrowing rapidly to a delicate
backward-curved point, the extremity of which in uninjured
specimens reaches to level of base of articular facet ; dental
foramen near posterior margin of base of articular process, and
on level with rim of alveolus ; a slightly larger foramen, 1 "5 mm.
below and in front of dental foramen at posterior extremity of
depression between outer alveolar wall and inner surface of
ascending portion of mandible.

TeefJi. — As compared with those of the typical voles, such as
the members of the genera Microtus and Arvicola, the incisors are
weaker and the molars heavier relatively to size of skull. The
molars are further peculiar in the unusual
width of the posterior tooth, which causes
the cutting surface of the tooth-row to be
almost parallel-sided instead of backwardly
tapering ; main axes of upper tooth-rows not
essentially parallel as in the voles, but so
converging anteriorly that if prolonged they
would meet slightly in front of incisors.
Upper incisors rather long and distinctly
curved, not projecting forward, the root pro-
ducing a slight swelling at front of ante-
orbital foramen ; cross-section of shaft ap-
proximately semicircular, the Hatteued border
directed inward, the anterior margin pale
yellow in colour, slightly wider than posterior
margin, the greatest transverse diameter a
little behind middle ; enamel covering anterior
fourth of inner surface and extending to slightly beyond middle of
outer surface ; cutting surface of tooth deeply and conspicuously
hollowed, its anterior margin bent sharply backward internally,
following course of enamel, its posterior border forming a distinct

Lemmus lemmus.
Teeth. X 5.

620

RODENTIA

cusp-like projection on margin of central hollow, slightly above
level of alveolus. Lower incisor more slender than upper tooth,
its root extending to level of middle of m.„ its cross section essen-
tially like that of upper incisor, but greatest transverse diameter
in front of middle, and postero-external border distinctly flattened.
First upper molar with an anterior transverse loop and four
alternating closed triangles. The anterior loop somewhat
triangular in form, owing to the deep re-entrant angle at its
postero-internal border, the two closed triangles on lingual side
of tooth noticeably smaller than the others, their main axis
oblique instead of transverse, the salient angles tending to be
truncate at their extremites ; re-entrant angles four, all well
developed and regularly alternating, two on each side of tooth.
Second upper molar with crescentic anterior loop isolated by the
very deep first oviter re-entrant angle, a second loop essentially
like first loop of rii^, and two closed triangles like the posterior
pair in m^ but slightly smaller ; re-entrant angles three, all well
developed and regularly alternating, two outer and one inner.
Third upper molar with four loops extending completely across
crown, the first crescentic like that of m^, and similarly isolated
by a single deep inner re-entrant angle, the second slightly
narrower and bounded posteriorly by two exactly opposed
re-entrant angles, the third similar to second but bounded
posteriorly by a deep inner re-entrant angle and a very shallow
exactly opposed outer angle, the terminal loop with transverse
diameter slightly less than in the others, its posterior border
with a shallow but noticeable concavity on inner side ; re-entrant
angles five, the first (outer) extending across crown, the second
and third opposite and equal, the last two opposite and unequal,
the inner extending nearly across crown, the outer very shallow.
First lower molar with a narrow anterior loop thrown into a
projecting point in front by the presence of a broad concavity on
both outer and inner border ; closed triangles three, essentially
alike in form, but that on outer side noticeably smaller than the
others ; posterior loop similar to anterior loop of nr and vf but
narrower ; re-entrant angles five, all well developed and alter-
nating, two outer and three inner (in some specimens one or both
of the concavities on anterior loop might be counted as a small
additional re-entrant angle). Second lower molar with a posterior
loop like that of first, and four triangles, the first pair of which
are smaller than the second pair ; re-entrant aqgles four, all well
developed, two on each side. Third lower molar consisting
essentially of three tranverse loops isolated by the two re-entrant
angles extending across crown from inner side, but second loop
with postero-external border indented by a small but well
developed re-entrant angle which cuts off outer exti'emity of loop
as a closed triangle not quite equal in size to the outer closed
triangle of m^ and nr ; terminal loop similar in form to that of
the other mandibular teeth, but slightly larger and less narrowed.

LEMMUS 621

Measurements. — External measurements of two males and a
female from Hjerkin, Dovre, Norway : head and body, 145, 129
and 145; tail, 15, 18 and 17; hind foot, 17, 17 and 18. Two
males and a female from Jemtland, Sweden : head and body,
129, 134 and 130; tail, 18, 19 and 19 ; hind foot, 17, 18 and 18.
For cranial measurements see Table, p. 622.

Specimens examined. — Seventy-seven, from the following localities ; —
Norway: Trondhjem, 4; R5raas, Trondhjem, 5 (B.M. and U.S.N.M.) ;
Brekkebygden, Trondhjem, 5; Hyllingen, Trondhjem, 3; Vigel Fjeld,
Trondhjem, 4; Kvikne, Trondhjem, 2; Dovre Mts., 1 (U.S.N.M.); Hjerkin,
Dovre, 5 ; Fille Fjeld 3 ; IMolmen, northern Gudbrandsdal, 2 ; Tonsast,
Hedemarken, 1 (U.S.N.M.) ; Stor Elvedal, Hedemarken, 1 (U.S.N.M.) ;
Lillehammer, Kristiansamt, 1 (U.S.N.M.) ; Gudbrandsdal, no exact
locality, 1 (U.S.N.M.) ; Gausdal, Gudbrandsdal, 3 (U.S.N.M.) ; no exact
locality, 9.

Sweden: Jemtland, 27 (B.M. and U.S.N.M.).

2. Roraas, Trondhjem, Nor- Christiania Museum 93. 3. 1. 16-17.

way. (e).

3(5,5 9. Trondhjem. G.Barrett-Hamilton 8. lU. 5. 1-8.

2 6. ICvikne, Trondhjem. G.Barrett-Hamilton 8.10.5.12-13

6, 2 9. Hjerkin, Dovre. G. Barrett-Hamilton 8. 10. 5. 9-11.
{J. L. Bonhote.) (p).

6,9, Fille Fjeld. (Brown and E. R. Alston (p). 79.9.25.63-65.

6 juv. Alston.)

2 6. M5lmen, Gudbrandsdal. R. J, Cuniughame 98. 5. 2. 10-11.

(c & p).
2. Norway. F. Du Cane Godman 73.3.21.2-3.

7 al. Norway. Prof. Owen (p). 56. 7. 9. 9-15.

5 <5, 4 9. Jemtland, Sweden. Lord Lilford (p). 8. 10. 19. 19-27.

(G. Kolthoff.)

Note on Lemmus lemmus crassidens Nehring.

On March 3, 1896, Major G. E. H. Barrett-Hamilton exhibited
before the Zoological Society of London " several fresh-looking
skeletons of the Norway Lemming {My odes lemmus), obtained by
Dr. H. Gadow in caves near Athouguia, in Portugal." Commenting
on these specimens he said : " Early in the year 1895 Dr. H. Gadow
handed me for examination some skeletal remains of a species of
small mammal, which, on a first inspection, appeared to be those of
some species of Vole- — Microtus. Thinking the remains were those
of Voles I put them aside for a time ; but later on, when I had an
opportunity of examining them more carefully I found, to my
surprise, that they consisted of some skeletons and detached bones of
the Norway Lemming, Myodes lemmus. When first received by me
the remains consisted of a good many fragments and single bones,
and of two almost complete skeletons. These latter were completely
enveloped in the original skin, which had become so dried and
hardened that in order to enable myself to examine the skeletons I
had to get it remo^•ed. The whole appearance of the specimens was
so fresh that, unaware as I was of their true character, I had the
dried skin, which enveloped them like mummies, removed, so that, I

622

RODENTIA

Observations.

Ridges joined, low.

moderate.

high.

moderate.

low.
,, ,, moderate.
,, nearly joined.
,, joined, low.
,, ,, moderate.

,, low.

•Aioj-moo5

00-^ CM (MOOOOOCOOOdOOOOOOOCD O .COOO i

jBinqipuBjc

t- t- 00 00 oD t- t~ t- t- 00 c- t- 00 t- t- oo ' t- b- '

•AVO.I-l[C(00^

OlC-l CM O^CMCMCMCMOCMOrHOO O^JIOq^CO

.^.reilixBK

OO C» 00 00 CX) 00 QO 00 00 00 CO 00 CO 00 00 CB 00 00 00 CO

•9iqipuBK

-*0 O OCO^CMCO(MO<I^CO<M(>)CO CO OCq

Oth th iHOooa3Oooo-HO05 o loo 1

CM CM (M (MCM(MCqrHCMCMCMCMCMi-lTH iH (MCM

•Btua'jsBiQ;

00^ ^ Ttl^OOCMO^CMOOCOO CO^CDOCM

030 a ooooc3500000 03c:» c^ioocjio

7-1 iHrHiHiHi-iiHT-li-(i-l iH

•IBSBiJ

00 CM 00 CM (M ^ O O '^ O O 00 O CM O CM . C^ -^ .

cries GO C35 OS c» CTi (i) 00 C2 CT) CX) til 00 cjo oo ' oo cx) '

-urejq jo q'jdofi

00"* O ^tOOOCMCMOcMCMOOCOO-l OOCMCOCO ,

t-00 CX) oo CX) t- c» CX3 ci)(i) CX) CX) cis t- CO t- co t- t- '

HUp'Ba.iq

piO^SBK

OOCM O CMCMO^ M^OCMOOOO-* OCMOOCM

-^lO >0 »niO^-^ l-rtilOlOlOlOCO^ ^lOCO-* 1
rHi-l tHiHtHt-ItH iHi-(iHi-lrHrH>H rHiHiHiH

•nonou^suoo
[■B^iqio.is:)!!!

00 O 00 O O O O O O O O O O CM O CM O 00 00 1

CO -^ CO 4)(4}i(r)-^4t(4j<-^4tfTi(-.a(-;j<4ti 4jf4tHcocM '

•qipiJS-iq
oi^BiuoS-tz

OOCM -^ ^^OO^^OCMOOOCMO O OOO

CX) cb cb th (b cb o 00 cb cb cb <b --I cis o cb loco 1

tHCM cm CMCM(MC^lT^C<ICM<MC^ICMT-^C^^ T-H CM 1-1

•mSnai
IBSTjqoi^puoo

OOO ■* CM O 00 to -^ 00 00 CM CO CM CM ■* CM O CO ■*

cb iH tH cm tH Ol O en O iH tH O rH C31 05 05 0 05 05 1
COCO CO CO CO CM CO C<1 CO CO CO CO CO CM CM CM CO CM CM

»0 •O O 'O CM- 'O "^ *« <^ 'O 'C 0+ CH- O

Number.

85. 5. 2. 10
8. 10. 5. 2

154541

8. 10. 5. 9

154542
8. 10. 5. 10
8. 10. 19. 25

153134

8. 10. 19. 23

153138

153133

8. 10. 19. 24

153136
8. 10. 19. 26
8. 10. 19. 27

8G6c, 2 L*

,, iL

Locality.

' — ' — '

L. lemmus lemmus.

Norway : I\l61men .

Brekkebygden
T6nsa3t, Hede

marken
Hjerkin, Dovre

Sweden : Jemtland

L. lemmus "crassidens.

Portugal ?

EVOTOMYS

623

regret to say, not one of these most interesting specimens has been
pi-eserved in the condition in which I received it. Some of the
vertebne, however, are still connected together by the dried remains
of the ligaments. This, and the whiteness and excellent preservation
of the bones, will show how easy it was to be deceived as to their
natiire, and to come to the belief that they were of recent origin and
perhaps unimportant . . . The present skulls resemble those of recent
Lemmings very closely indeed...! cannot find any characters
sufficiently important to enable me to separate the two specifically "
(Proc. Zool. Soc. London, 1896, pp. 304-305). Three years later,
however, they were examined by Nehring and distinguished from the
Norway Lemming as Myodes Icmmus crassidens, principally on
account of their relatively large molar teeth and certain fancied
peculiarities in the form of the coronoid process of the mandible
(Wiegmann's Archiv fiir Naturgeschichte, lvi, i, p. 174, June, 1899).

Through the courtesy of the authorities of the Cambridge Museum
I have been able to compare the original crassidens material with a
series of seventeen skulls of true Icmmus in the British Museum. As
submitted to me this material consisted of three perfect skulls
(including the type), one skull without lower jaw, one rostral portion
of skull together with palate and maxillary teeth, two complete lower
jaws, and one single mandible, the whole representing not less than
six nor more than eight individuals. The specimens are well
preserved, with most of the teeth in place, and have all the
appearance of fresh material. In size and form the skulls show no
peculiarities. No old individuals are represented ; the measurements
therefore do not attain the maximum (s'ee Table opposite). The
coronoid processes are broken at the tip, giving them the short,
blunt appearance noted by Nehring. In one complete skull and the
odd mandible the teeth are of the same size as in ordinary average
specimens of true lemmus ; in the type they equal those of the
largest-toothed Norwegian specimen in the series (from Molmeu,
northern Gudbrandsdal) ; in one complete skull, one skull without
lower jaw, and one odd lower jaw they are slightly above the
maximum of lemmus ; while in the fragmentary skull and two odd
jaws they are decidedly above the maximum. That is, assuming
that eight individuals are represented, three are perfectly matched
among the seventeen Scandinavian specimens, while five are not.
This makes it impossible at present to synonymize crassidens with
Icmmus. On the other hand, it seems almost equally impossible to
believe that the specimens came from Portugal, and that some
confusion of material did not take place during the period when
Dr. Gadow's Athouguia mummies, regarded as "unimportant," were
"put aside."

Genus EVOTOMYS Coues.

1839. Myodes de Selys-Longchamps, Etudes de Micromamm., p. 87

(section).

1840. Hypudxxis Keyserling and Blasius, Wirbelth. Europas, p. 34 (sub-

genus: type Mas glarcohis Schreber), not Hypiuhnis Uliger, 1811.
1858. Hypiidxus Blasius, Siiugethiere Deutschlands, p. 336 (sub-genus).
1874. Evotomys Coues, Proc. Acad. Nat. Sci. Philadelphia, p. 186 (genus,

type Mus rutilus Pallas).
1896. Evotomys MiUer, North Amer. Fauna, No. 12, p. 42, July 23, 1896.

624 RODENTIA

1900. Craseomys Miller, Proc. Washington Acad. Sci., ii, p. 87 (sub-gonus
of Evotomys : type Hypuclxus rufocanus Sundevall).

1907. Craseomys Thomas, Proc. Zool. Soc, London, 1906, p. 8G3, April 11,
1907 (genus).

Ti/pe species. — Mus rutilus Pallas.

Geographical distribution. — Boreal portions of both hemi-
spheres ; south in Europe to the Pyrenees and the mountains of
southern Italy.

Characters. — Bony palate terminating posteriorly in a simple,
thin-edged, horizontal shelf ; lower incisor with root extending
to outer side of molar-roots, but not rising above level of cutting
surface of molars and not forming any protuberance on outer
surface of ascending portion of mandible ; molars with two well
developed prongs or roots in adult, the crown disappearing by
wear in extreme old age ; pattern of enamel folding characterized,
as compared with that of other European voles, by a tendency
toward roundness or bluntness of the salient angles ; external
form without special peculiarities.

Rcma7-Jis. — In the shortness of the mandibular incisor root, a
peculiarity which it shares with the American Phenacomys, the
genus Evotomys stands nearer to the Lemnii than any of the other
European members of the sub-family. Its rooted molars must
also be regarded as a primitive character, since the same
peculiarity is found in many of the related fossil genera. About
tifty forms are now recognized in the genus, thirteen of which
occur in northern Europe. These have recently been grouped in
three sub-genera,* two of which have even been regarded as
genera ; but the discovery of intermediate species makes this
course seem no longer tenable, f

KEY TO THE EUEOPEAN FORMS OF EVOTOMYS.

Tail densely haired, the pencil at least one-fourth

as long as vertebrae (Arctic Europe) E. rutilus, p. 646.

Tail moderately haired, the pencil much less than
one-fourth as long as vertebrte.
Skull of adult massive, its size large, the condylo-
basal length frequently more than 26 lum.
(25 to 27-6 mm.); teeth heavy, length of
tooth-rows 6 mm. or more.
Red of dorsal area conspicuously contrasted
with grey of sides ; ear noticeably over-
topping fur; vi^ normally without third
re-entrant angle on inner side (Northern
and mountainous portions of Scandinavia) E. rufocanus, p. 648.

* True Evotomys, Craseomys Miller, and Phaulomys Thomas (Ann.
and Mag. Nat. Hist., 7th ser., xv, p. 493, May, 1905. Type Evotomys
smithii Thomas, from Kobe, Hondo, Japan).

t For observations on the lack of distinctions between Phaulomys and
Craseomys see Anderson, Ann. and Mag. Nat. Hist., 8th ser., iv, p. 318,
October, 1909. For notes on the status of Craseomys see account of
Evotomys rufocanus, below, p. 658.

EVOTOMYS

625

Red of dorsal area not conspicuously contrasted
with yellowish hrown of sides ; ear scarcely
overtopping fur ; m^ normally with third
re-entrant angle on inner side (Channel

Islands) .' E. cxsarms, p. 645.

Skull of adult delicate or slightly massive, its size
small or moderate, the condylobasal length
rarely 2G mm. (21-5 to 26-2 mm.); teeth
light or moderately heavy, the length of
tooth-rows rarely 6 mm. (4-8 to 6 mm.) ; ear
decidedly overtopping fur.
Nasal decidedly longer than diastema [hind foot
18 to 19 mm. ; condylobasal length of skull
24-8 to 25-8 mm.; colour bright, the line
of demarcation along sides of body notice-
able] (Skomer Island, Wales) E.skomerensis, p. 644.

Nasal about equal to diastema, rarely much

longer -E. glareolus, p. 626.

Size smaller, hind foot usually less than 18 mm.
(15-4 to 18 mm.); condylobasal length
of skull usually less than 24-5 mm.
(21-5 to 24-5 mm.). Characteristically
lowland forms.
General colour above light and bright

(Drainage system of the Danube) E. g. istericus, p. 637.

General colour above dark or dull.

Red of dorsal area showing little if any
tendency to suffuse sides and rump
(Central and southern Sweden ;

southern and eastern Norway) E. g. sxtecictis, p. 6.36.

Red of dorsal area tending to suffuse sides
and rump.
Size medium, hind foot 16 "6 to 17*6
• mm., condylobasal length of skull
23 to 24-6 mm. ; colour dark and
usually bright (West-central conti-
nental Europe) E.g. glareolus, p. 632.

Size smallest, hind foot often less than
16-6 mm. (15 -4 to 17 mm.), condy-
lobasal length of skull often less
than 23 mm. (21-5 to 24-2 mm.) ;

colour dark and dull (Great Britain) E. g. hritannicvs, p. 634.
Size larger, hind foot usually more than
18 mm. (17 to 20 mm.) ; condylobasal
length of skull usually more than 24 • 5 mm.
(23 to 26-2 mm.).
Third upper molar usually with two re-
entrant angles on inner side.
Red dorsal area broad, not well-defined
from buffy grey sides (Western

Norway) -£'• </• iwrvegicvs, p. 638.

Red dorsal area narrow, well defined from

dull grey sides (Pyrenees) E. g. vasconioe, p. 639.

Third upper molar usually with three re-
entrant angles on inner side.
Size smaller (hind foot 17 to 19 mm.,
condylobasal length of skull 23 to
25 "4 mm.); colour lighter (Jura to

south-western Alps) E. g. helveticus, p. 640.

2 s

626 RODENTIA

Size larger (hiud foot 18 -G to 21 mm.,

condylobasal length of skull 25 to

26 • 2 mm.) ; colour darker.

Teeth weak (normal) ; incisive foramina

narrow (normal), the width of the

two together about J length (Alps,

except western portion) E. g. nageri, p. 641.

Teeth heavy ; incisive foramina wide,
the width of the two together more
than J length (Southern Italy) E. g. haUucalis, p. 643.

EVOTOMYS GLAREOLUS Sclireber.
(Synonymy under subspecies.)

Geographical distribution. — Europe from the Pyrenees and
southern Italy to Scotland and central Scandinavia, west to
Great Britain, east into Siberia. Eastern limit of range not
known.

Diagnosis. — Tail about half as long as head and body or
somewhat more, moderately haired, the pencil always much less
than one-fourth as long as vertebrae ; skull of adult delicate or
moderately heavy, the condylobasal length rarely 26 mm.
(21 '5 to 26*2 mm.); nasal usually about as long as diastema;
teeth light or moderately heavy, the length of tooth-rows rarely
6 mm. (4 • 8 to 6 mm.) ; colour variable but never so bright as in
Evotoinys rutilus, that of the larger local forms always tending
to be dull.

External characters. — Form rather heavy, the head short and
blunt, with rounded muzzle and small eye. Ear slightly over-
topping the fur, extending barely to eye when laid forward, its
outline sub-circular ; meatus with well developed semilunar valve,
both surfaces of which are naked ; a tuft of long hairs in angle
between outer extremity of valve and outer margin of ear-conch ;
surface of ear densely pubescent except on basal half of inner
side. Muzzle pad small and inconspicuous, the aperture of the
nostril slit-like, wider anteriorly than posteriorly, the margin
swollen, particularly in front ; upper lip with narrow median
cleft continued upward across pad. Fore foot with thumb
reduced to a minute tubercle scarcely half as large as the
smallest pad on palm, its nail rudimentary and closely appressed,
but easily visible with a lens ; outer digit extending to base of
fourth ; second about half as long as third ; fourth and third sub-
equal ; palmar tubercles five, large, sub-equal, crowded, occupying
nearly the entire surface of palm, the skin between them finely
tuberculo-reticulate. Hind foot with inner digit extending to
base of second, outer slightly beyond base of fourth, the second,
third and fourth sub-equal and about one-third as long as sole ;
surface of sole densely pubescent behind tubercles, naked and
tuberculo-reticulate between them ; tubercles six, well defined,
moderately large, essentially alike in form, their outline ovate ;

EVOTOMYS

627

postero-external tubercle smaller than the others (ahout half as
large as postero-internal). Tail about half as long as head
and body or slightly more, its annulations ill defined and
completely concealed by the dense covering of hair ; pencil
well developed, usually about one-sixth as long as vertebra?.
Mammte : p2 — 2, i2 — 2 = 8.

Colour. — Upper parts brown with a decided reddish or
ochraceous suffusion, especially along back ; sides and rump
usually lighter than back, often with a tinge of grey, but never
forming a decided contrast as in Evotomiis rufocanus; under-
parts light grey with or without a yellowish brown wash ; feet
greyish ; tail obscurely bicolor.

Skull. — The skull of Evotomi/s glareolus is small and lightly
built, the surface smooth and rounded, without noticeable ridges
or depressions for muscular attachment. Dorsal protile in
general slightly convex throughout,
though flattened or even a little con-
cave in interorbital region, the nasals
sloping rather abruptly forward, their
surface nearly flat ; there is usually
no sharp distinction between dorsal
and posterior outlines, the profile
merely bending abruptly downward
posteriorly over somewhat obliquely
truncate occipital region ; ventral
profile shallow-concave anteriorly, then
essentially straight to lower surface of
bulla ; depth of skull considerably
greater posteriorly than anteriorly, so
that the general outline formed by
the two profiles is distinctly wedge-
shaped. Brain-case sub-circular or
very broadly oval in outline when viewed from above, sometimes
a little squared at sides or in front ; the surface smooth except
for the small but evident postorbital processes, a distinct trace
of the lambdoid crest extending downward from outer extremity
of interparietal to meatus, and a shorter but more noticeable
vertical ridge along upper part of suture between occipital and
petrous portion of temporal, this ridge lying when skull is
viewed from above in axis of concavity between paroccipital
process and occipital condyle ; in old individuals there is also a
slightly indicated angularity extending along sides of brain-case
and across median lambdoid region ; interparietal convex behind,
double concave in front, its lateral extremities acute or slightly
truncate, its greatest antero posterior diameter about one-half to
one-third transverse diameter ; occiput when viewed from behind
broadly ai'ched, >slightly flattened above and at sides, the median
portion of brain-case not rising conspicuously above lambdoid
level ; paroccipital processes slender but well developed, free

Fig. 124.
Evotomi/s glareolus. Nat. size.

628 EODENTIA

from bulli<?, their points extending to slightly below level of
lower lip of foramen magnum. Floor of brain-case smooth
except for a median ridge on anterior half of basioccipital ; width
of basioccipital along anterior border about one-third median
length. Auditory bull.ie large, rounded and evenly inflated, their
surface smooth, their walls thin and translucent ; anteriorly they
overlap pterygoids for a distance of about 1 mm., though without
much encroaching on pterygoid fossa or developing a noticeable
beak. Interorbital region wide and smooth, its least breadth
conspicuously greater than that of rostrum at region of greatest
width of nasals, its upper surface with broad and shallow but
noticeable longitudinal depression. Zygoma slender, scarcely
expanded at middle, the arches when viewed from above strongly
convex in front but spreading abruptly enough to bring the
greatest zygomatic breadth a little in front of middle of orbit ;
anterior zygomatic root with faint concavity over infraorbital
foramen. Rostrum short and weak, tapering noticeably forward
when viewed either from above, below, or the side ; protuberances
over roots of incisors slight ; nasals short, bluntly truncate
posteriorly at level a little in front of middle of zygomatic root,
usually somewhat exceeded by nasal branches of premaxillaries ;
outer border of nasal when viewed from above essentially
straight from posterior border to widest region. Incisive
foramina long and rather wide, extending from about 2 mm.
behind alveolus of incisor to within 1 mm. of level of m^. Palate
rather short and narrow, slightly concave laterally, its posterior
border, at level of anterior loop of m^, variable in form but
usually with a slight median convexity and a slight concavity at
each side ; mesopterygoid space long, nearly parallel-sided, the
hamular processes very slightly bent outward, applied against
inner surface of bullaj. Mandible slender and weak, particularly
the hinder portion, into which the root of incisor penetrates so
slightly as to produce no evident thickening ; masseter ridge low
and inconspicuous except in old individuals ; coronoid process
rather long, noticeably curved backward, its extremity nearly
reaching level of condyle ; articular process long, slender and
compressed, the dental foramen lying below its base at level of
cutting surface of molars ; angular process slender and rather
long, convex externally, concave internally, its main axis curving
outward and upward.

Teeth. — Compared with those of the European species of
Microtus the incisors are slender and weak, and the molars are
small, with both salient and re-entrant angles rounded and ill-
defined. Upper incisor projecting downward, the cross-section
of shaft bluntly triangular, the inner surface widest, the anterior
and postero-external surfaces narrower and sub-equal ; enamel
well defined, extending over entire anterior surface and extreme
antei'ior portion of inner surface ; cutting surface of tooth
distinctly hollowed but not sufficiently to produce any suggestion

EVOTOMYS

629

of the weakened appearance characteristic of Lemmns and
Myopus, its posterior border fading gradually into surface of
tooth or marked oif by a slight angle. Lower incisor noticeably
more slender than upper tooth, its root extending beneath that
of 7H3, but not rising to level of cutting surface of molars, its
shaft nearly semicircular in cross section, the flattened surface
directed inward, the enamel extending over anterior half of
curved surface ; anterior surface of both upper and lower incisors
yellowish brown, posterior surface whitish. Molars relatively
small and weak, at first rootless and growing from a pulp as in
Lemmus and Microtus, the prisms extending uninterruptedly to
base. At a slightly later stage the extreme base is seen to be
smooth, forming a ring below prisms ; this ring then contracts
at middle and divides into two ; as the cutting surface of the
crown wears away and the tooth is pushed upward in the
alveolus, each of these secondary rings narrows and lengthens
into a prong or root standing in axis of tooth-row, the anterior

Pig. 125.
Evotomi/s glareolus. Enamel pattern in two individuals.

prong of each tooth somewhat larger than the posterior ; with
advancing age the prongs become relatively longer and the
crowns lower, so that in the final stage the crowns may com-
pletely wear away, leaving in their place the flattened heads of
the six roots. It follows that the enamel pattern, instead of
remaining essentially constant throughout life, as in the genera
with rootless molars, undergoes marked changes in the later
stages of crown-wear, gradually losing its definiteness and finally
disappearing. As compared with that of European voles of
other genera the enamel pattern is characterized by a general
lack of sharp angularity, especially in the salient angles on
inner side of the maxillary teeth, where the loops assume the
form of circles or semicircles rather than that of closed triangles,
and by the arrangement of the prisms of the mandibular teeth
in imperfectly alternating pairs, so that completely isolated and
closed triangles are the exception rather than the rule. In
crown length m^ slightly exceeds m\ which in turn is usually
somewhat longer than «r, the discrepancy in neither case very

G30 RODENTIA

noticeable ; iHj nearly equal to wi., and m.^ comLinerl, m.^ slightly
shorter than }/(,,. First upper molar with a transverse, evenly
and broadly crescentic anterior loop and four alternating,
approximately equal closed triangles, those on outer side slightly
the larger ; each side with three salient and two re-entrant
angles. In young individuals the outer re-entrant angles are
conspicuously longer than those on inner side, but much more
oblique in direction, so that they do not extend appreciably
further beyond middle of crown ; the outer salient folds are at
this stage rather acvite, and the inner folds though blunt are
distinctly angular. With increasing age first the inner folds
and later the outer folds lose their angularity and assume an
indefinite rounded or semicircular form, while the discrepancy
in depth between the outer and inner re-entrant angles becomes
less marked. Second upper molar with anterior transverse loop,
one inner and two outer closed triangles ; outer side with two
re-entrant and three salient angles, inner side with one re-entrant
and two salient angles. Anterior loop unsymmetrical, broadly
rounded internally, narrowly pointed and slightly curved back-
ward externally ; rest of crown essentially like last three prisms
of m}, but discrepancies in form between outer and inner
elements less marked in youth and earlier disappearing with age.
Third upper molar with anterior transverse loop similar to that
of m^ but with outer, narrowed portion relatively less developed ;
following this are an inner and two outer triangles, closed or
variously opened, and a posterior longitudinal loop of variable
form ; inner triangle the largest of the three, though usually
not much exceeding antero-external triangle ; postero-external
triangle decidedly smaller than either of the others ; all three
triangles may be completely isolated from each other and from
terminal loops, or all may be open at both sides, leaving a
continuous dentine area along middle of crown ; usually the
enamel of the two sides comes in contact at one or more points,
and in the majority of specimens the inner triangle is completely
or essentially closed. The terminal loop when at its smallest is
short and perfectly simple in form, its greatest diameter oblique
to the tooth-row and equal to a little more than half that of
anterior loop ; when at its largest its greatest diameter distinctly
exceeds that of anterior loop, a shallow but evident re-entrant
angle is developed at its antero-external base just behind second
outer closed triangle, and its inner side is cut by a deep
re-entrant angle extending across to enamel of outer side.*
In its most simple form the tooth, has three salient and
two re-entrant angles on each side ; in its most complicated
form it has three salient and three re-entrant angles on outer
side, four salient and four re-entrant angles on inner side. First

* This angle is best developed during the earlier and middle stages of
crowu-wear; later it tends to become obsolete at a less advanced stage
than those forming part of the strictly normal pattern. The frequency of

EVOTOMYS

631

lowev inolar with a narrowly crescentic, somewhat unsym-
metrical posterior transverse loop, three inner and three outer
closed or partly open triangles, and a short rounded anterior
loop ; outer side with three well defined salient angles and the
same number of re-entrant angles, inner side with four well
defined salient angles and four re-entrant angles ; the number
of salient angles on either or both sides may be increased by
one through the assumption of a distinctly angular form by the
base of the anterior loop, this occurring more frequently on the
outer than on the inner side. The folds are moderately angular,
those of inner side slightly the larger, but the contrast in size
and form is much less evident than in the upper teeth. Second
and third lower molars essentially alike in form, each consisting
of three transverse loops, three inner and three outer salient
angles, and two inner and two outer re-entrant angles, these last
better developed in ui.^ than in m.,, in which the anterior re-entrant
angle is sometimes obsolete. The first and second loops of m.,
and the second loop of mg often show a tendency to become
partly or almost completely divided into two triangles, a
peculiarity more evident in the second tooth than in the third.

Bemarks. — Evotomys glareolus is the common, widely-dis-
tributed European representative of the genus. It is divided
into nine geographical races. The forms inhabiting Great
Britain and the central portion of the Continent are small,
while those of western Norway, the Alps, Pyrenees and the
mountains of southern Italy are noticeably larger. There seems
little doubt, however, that intergradation between the small
and large forms takes place in every instance, though as yet it
can be satisfactorily demonstrated in the case of glareolus and
nageri only.

presence or absence is

also largely a

racial character,

iS may be seen

Q the following tabular statement : —

Percentage of

Number of

Angle

Angle

specimens with

specimens.

present.

absent.

angle present.

E. glareolus glareolus...

100 .

. 51 .

. 49

.. 51-0

E. glareolus britannicu.

43 .

. 11 .

. 32

.. 25-5

E. glareolus stiecicus....

37 .

. 12 .

. 25

.. 32-4

E. glareolus istericus....

56

. 36 .
5 .

. 20
15

64-3

E. glareolus norvcgicus.

20 .

.. 25-0

E. glareolus vasconiir ...

29 .

8 .

. 21

.. 27-6

E. glareolus hclveticus..

.... 100 .

. 68 .

. 32

.. 08-0

E. glareolus nageri

.... 100 .

. 85 .

. 15

.. 85-0

E. glareolus hallucalis..

1 .

1 .

—

—

E. skomerensis

15 .

. 14 .

1

.. 93-3

E. csesarius

22

. 22 .

. 16 .

1

0

0

15

100

E. ruti lus

16

100

E. rufocamcs

16 .

6-2

63:

RODENTIA

EVOTOMYS GLAREOLUS GLAREOLUS Schrebei".

1780. Mas glareolus Schreber, Saugthiere, iv, p. 680, pi. cxc b (LoUand,

Denmark).
1792. Mus rutilus minor Kerr, Anim. Kingd., p. 237 (Casan and Goettingen).

Based on Pallas, Glires, p. 247. Not Mus rattus viinor Kerr,

Anim. Kingd., p. 229.
1792. ^Mus rutilus] /3 minor Donndorft', Zool. Beytriigo, i, p. -152. Based

ou Pallas, Glires, p. 247.
1803. Lcvimics arvalis Geoffrey, Catal. Mammif. du ]\Ius. Nat. d'Hist. Nat.,

p. 185 (Meudon, Seine, France). Not Mtts arvalis Pallas.
1828. Arvicola fulvus Millet, Faune de Maine-et-Loire, ii, p. 40 (Angers,

Maine-et-Loire, France). Not Lemimis fulvus Geoffroy, 1803.
1831. Hypudxus hercynicus IMehlis, Isis, p. 876 (Harz Mountains, Germany).
1884. Lemmiis ruhidus Baillon, M^m. Soc. Royale d'Emulation d'Abbeville,

1833, 15. 54 (Abbeville, Somme, France).
1834. H[ypud;vus'] glareolus Melchior, Den Danske Stats og Norges

Pattedyr, p. 116.
1836. Arvicola rufcsccns de S^lj-s-Longchamps, Essai Monogr. sur les

Campagnols des Environs de Liege, p. 13 (Longchamps-sur-Ger,

Belgium).
1842. Arvicola iwatensis F. Cuvier, Hist. Nat. des Mammif., vii, Tabl.

Gen. et M6th. Described and figured in Livr. 68 of same work,

1834 (Abbeville, Somme, France).
1857. Arvicola glareolus a. Blasius, Saugethiere Deutschlands, p. 337 (part).
1896. E[votomys'] glareolus Miller, North American Fauna, No. 12, p. 44,

July 23, 1890.
1900. Evotomys hercynicus hercynicus Miller, Proc. Washington Acad.

Sci., II, p. 100, July 26, 1900 (part).
1900. Evotomys hercynicus ruhidus Miller, Proc. Washington Acad. Sci.,

II, p. 102, July 26, 1900.

1909. Evotomys glareolus Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

pp. 419-420, May, 1909.

1910. Evotomys glareolus Trouessart, Faune Mamm. d'Europe, p. 170.

Type locality. — Island of Lolland, Denmark.

Geographical distrihution. — West-central Europe north of the
Alps and Pyrenees and south of the Baltic ; from the Atlantic
coast east to Silesia ; north-eastern limit of range not shown.

Diagnosis. — General colour above dark, but usually rather
bright, the dorsal reddish area broad and diffuse ; usual measure-
ments of adults : hind foot, 16 '6 to 18 mm., condylobasal
length of skull, 23 to 24-6.

Colour. — Winter pelage : red dorsal area diffuse and ill-
defined, extending from eyes nearly to base of tail and tending to
spread slightly over sides. In general colour it is very nearly
the mars-brown or prouts-brown of Ridgway, visually with a
tinge of russet, the exact shade resulting from vaiuous combina-
tions of cinnamon-rufous, vinaceous-rufous and black. Sides and
cheeks dull brownish buff thickly sprinkled with black. The
colour of sides merges insensibly into that of back and passes
abruptly into the strongly buff-tinged grey of belly. The buff
wash on ventral surface is variable, but usually conspicuous.

f:voTOMYS 633

Hairs everywhere slaty at base, this colour appearing irregularly
at surface on sides and underparts. Feet whitish, distinctly
tinged with brown. Ear like dorsal red area. Tail sharply
bicolor, dark bi'own above, soiled white below.

Skull and teeth. — Skull small, nearly the minimum for the
species (see Table of measurements, p. 650) ; auditory bulhe not
abruptly inflated on inner side, a character scarcely appreciable,
however, except on comparison with E. glareoliis istericus. Third
upper molar with third inner re-entrant angle present in about
one-half the specimens examined (see p. 631).

Measurements. — External measurements of adult male and
nearly adult female from Nystad, Lolland, Denmark : head and
body, 102 and 96; tail, 51 and 47; hind foot, 17*6 and 17*6.
Adult male from Hillerod, Zealand, Denmark : head and
body, 98; tail, 44; hind foot, 16' 5; ear from meatus, 13-5.
Average and extremes of ten adults from Brunswick, Germany :
head and body, 102-5 (98-111); tail, 47-3 (40-54) ; hind foot,
17' 1 (16* 6-17 -6). Two adult males from Miiuseklipise, Harz
Mts., Germany : head and body, 108 and 102 ; tail, 45 and 52 ;
hind foot, 16*6 and 17. Adult male and female from Guines,
Pas-de-Calais, France : head and body, 101 and 97 ; tail, 57
and 59; hind foot, 17*5 and 18 "5. Two adult males from
Barbizon, Seine-et-Marne, France : head and body, 103 and 108 ;
tail, 54 and 47 ; hind foot, 17 '6 and 17 '8. For cranial
measurements see Table, ]). 650.

Specimens examined. — One hundred and eighty-six, from the following
localities : —

Denmark: Hillerod, Zealand, 7; Nystad, Lollaud, 7 (U.S.N.M.);
Odense, Fyu, 2.

Holland: Oosterbeek, Guelderland, 1; near Leiden, 2 (U.S.N.M.).

Belgium: Waremme, Li^ge, 28 (U.S.N.M.); Maredsous, Li6ge, 15; no
exact locality, 1.

Fhance : Guines, Pas-de-Calais, 2 ; Pont de Briques, Pas-de-Calais, 5 ;
Dinau, C6tes-du-Nord, 4 ; Barbizon, Seine-et-Marne, 9 ; Mont-Dore, Puy-
de-D6me, 3; Lignieres- Sonueville, Charente, 16 (Mottaz) ; Cadillac,
Gironde, 1 (U.S.N.M.) ; Foret de Bouconue, Gers, 1 ; Lyons, Rhone, 1.

Germany: Brunswick, 44 (B.M. and U.S.N.M.); Bahrenberg, Harz
Mts., 2 (U.S.N.M.); Miiuseklippe, Bodethal, Harz Mts., 6 (U.S.N.M.)
Ingelheim, Rheinhessen, 6 ; Strassburg, 4 ; near Magdeburg, Saxony, 5
Tharandt, Saxony, 1 ; Saxony, no exact locality, 1 ; Niesky, Silesia, 6
Gross Henuersdorf, Silesia, 2 ; Wolfshau, Silesia, 4 (U.S.N.M.).

Austria-Hungary : Haida, Arva, Bohemia, 1 (not typical).

2 (J, 5 ?. Hillerod, Zealand, Den- 0. Thomas (c & p). 98. 6. 7. 13-19.
mark.

1 Kt. Odense, Fyn. Prof. Sundeval (p). 49. 11. 1. 5.

9. Oosterbeek, Guelderland, 0. Thomas (c & p). 98. 2. 1. 15.
Holland.

4 (5, 4 9. Maredsous, Liege, Bel- Rev. L. Warnau (p). 8. 10. 18. 1-8.
giuna.

1. Belgium. E. de S61ys Long- 45. 7. 58.

champs (p).

(5,9. Guines, Pas - de - Calais, 0. Thomas (c & p). 94.0.6.16-17.

France.

6, 4 9. Boulogne, Pas-de-Calais. O. Thomas (c & p). 98. 1. 9. 16-20.

634 RODENTIA

4:6. Diuan, Brittany. G.Barrett-Hamilton 8.9.29.4-5.

(c & p). 11. 1. 2. 43-44.

3 6,3 9. Barbizon,Seine-et-Marne. G. S. Miller (c). 8.8.4.242-247.

1. Lyons, Rhone. Purchased. 4.3. 10. 14. 9.
6. Foret de Bouconne, Gars. 0. Thomas (p). C. 4. 1. 86.

(A. Robert.)

2 c5, 1 ?. Brunswick, Germany. G. Barrett-Hamilton 8. 9. 29. G-7.

(c & p). 11. 1. 2. 45.

2 (J, 4 ?. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2.46-51.

2 6,2 9. Strassburg, Alsace. 0. Thomas (p). 8. 8. 10. 122-

(C. Mottm.) 127.

3. Magdeburg, Saxony. Dr. W. Wolterstorff 92. 12. 1. 18-20.

(c & P).

2. Herrenkrug, Magdeburg. Lord Lilford (p). 8. 10. 19. 4-5.
9 al. Tharandt, Saxony. Berlin Museum (e). 93. 1. 1. 21.

{Nitsche.)

1. Saxony. Dr. Thienemann (p). 45. 11. 15. 18.

2 6, 9. Niesky, Silesia, Germany. Dr. E. Hamilton (p). 97. 12. 4. 24-26.

(IF. Baer.)

2 6, 9. Niesky, Silesia. {W. Baer.) Lord Lilford (p). 99. 1. 9. 23, 24,

26.

6, 9. Gross Hennersdorf, Si- Lord Lilford (p). 99. 1. 9. 22, 25.
lesia. {W. Baer.)

6. Haida, Bohemia. Lord Lilford (p). 8. 10. 19. 1.

EVOTOMYS GLAREOLUS BRITANNICUS Miller.

1832. Arvicola riparius Yarrell, Proc. Zool. Soc, London, p. 109 (England,

no exact locality). Not of Ord, 1825.
1900. Evotomys hercynicus britannicus Miller, Proc. Washington Acad.

Sci., II, p. 103, July 26, 1900 (Basingstoke, Hampshire, England).

Type in British IMuseum.
1910. Evotomys glareolus britannicus Trouessart, Faune Mamm. d'Europe,

p. 170.

Ty]pe locaUtij. — Basingstoke, Hampshire, England.

Geographical distribution. — Great Britain.

Diagnosis. — Like Evotomys glareolus glareolus, but colour
usually less intense, and average size smaller (hind foot, 15 '4 to
17 mm. ; condylobasal length of skull, 21 "4 to 24*2 mm.).

Measurements. — Type (adult male, from Basingstoke, Hamp-
shire, England) : head and body, 105 ; tail, 51 ; hind foot, 16 "6.
Average and extremes of eight adults from the type locality :
head and body, 103-4 (98-107) ; tail, 49-3 (44-54); hind foot,
16 "3 (15*4-17). Two adult females from Aberia, Merioneth-
shire : head and body, 100 and 101 ; tail, 48 and 45 ; hind
foot, 16 '4 and 15*8. Two adult females from Dumphail, Elgin,
Scotland : head and body, 102 and 110 ; tail vertebrse, 55 and 51 ;
hind foot, 16 and 16*8. For cranial measurements see Table,
p. 652.

Specimens examined. — Que hundred and eighty - three, from the
following localities : —

Scotland: Birnie, Morayshire, 2; Dunphail, Elgin, 5; Fochabers,
Elgin (Edinburgh), 1 ; Lhanbride, Elgin, 1 ; Cortachy, Forfar, 1 (Wilson) ;
Dunkeld, Perthshire, 1 ; Loch Earn Head, Perthshire, 1 ; Windygates,
Fife, 12 (Edinburgh) ; Dalmeney Park, Edinburgh, 2 ; Grantown-on-Spey,

EVOTOMYS

635

Edinburgh, 1 (Wilson) ; Cromlix, Dunblane, 5 ; Blackwood, Lanarkshire, 5 ;
Stockbriggs, Lanarkshire, 2 ; Drumlaurig Woods, Dumfriesshire, 2 ; Kirtlo
Bridge, l5umfriesshire, 1 ; Dumfriesshire, no exact locality, 1 ; Craigmere,
Galloway, 3 (Edinburgh).

England : Filey, Yorkshire, 1 ; Lanfair, Anglesey, 3 ; Llansligo, Angle-
sey, 2 ; Plas Maclog, Anglesey, 1 ; Parkgate West, Cheshire, 1 ; Higher
Sutton, Macclesfield, Cheshire, 1 ; Northenden, Cheshire, 5 ; Aberia,
^Merionethshire, 7 ; Langharne, Carmarthenshire, 4 ; Bridgend, Glamorgan-
shire, 1 ; Swansea, Glamorganshire, 1 ; Colwyn Bay, Denbighshire, 1 ;
Grimsby, Lincolnshire, 18 ; Waltham, Lincolnshire, 1 ; Herefordshire, 2 ;
Arley, Staffordshire, 2 ; Cheadle, Staffordshire, 1 ; Swithland, Leicester-
shire, 3;'Lilford, Northamptonshire, 3; Oundle, Northamptonshire, 1;
Histon, Cambridgeshire, 1 ; Worcestershire, 1 ; Wormsley, Oxfordshire, 4 ;
Luton, Bedfordshire, 2 ; Toddington, Bedfordshire, 1 ; Welwyn, Hertford-
shire, 1 ; Lowestoft, Suffolk, 2 ; Chelmsford, Essex, 1 ; Hillingdon, Middle-
sex, 6 ; Betchworth, Surrey, 2 ; Bletchingley, Surrey, 7 ; Godalming,
Surrey, 2 ; Kingston, Surrey, 1 ; Sussex, 1 ; Basingstoke, Hampshire, 13 ;
New Forest, Hampshire, 2; Eversley, Hampshire, 8; Bembridge, Isle of
Wight, 4 ; St. Helens, Isle of Wight, 1 ; Salisbury, Wiltshire, 1 ; Pill-on-
Avon, Somerset, 1 ; Blandford, Dorset, 1 ; Bridport, Dorset, 9 ; Northlew,
Devonshire, 6.

6.

B i r n i e, Morayshire,
Scotland.

E. R. Alston (v).

79. 9. 25. 58.

?al.

Birnie, Morayshire.

Rev. G. Gordon

(c & p).
W. R. Ogilvie-Grant

88. 11. 22. 1.

i,19.

Dunphail, Elgin.

11. 1. 3. 209-210.

(c & p).

11.1.3.239-241.

?.

Llanbride, Elgin.

W. Taylor (c & p).

11. 1. 3. 238.

2 9.

Dunblane, Perthshire.

W. R. Ogilvie-Grant

(c & P).
W. Evans (c & p).

11.1.3.211-212.

2 6.

Dalmeny. Edinburgh.

11.1.3. 235-23G.

6,2 9.

Cromlix, Stirlingshire.

W. R. Ogilvie-Grant

(c & P).

11.1.3.243-245.

2 6.

Stockbriggs, Lanark-
shire.

E. R. Alston (c & p).

79. 9. 25, 56-57.

2.

Drumlaurig Woods,
Dumfriesshire.

Jardine Collection.

86. 7. 2. 9-10.

6.

Kirtle Bridge, Dum-
friesshire.

Miss D. Bate (c & p).

11. 1. 3. 242.

6, 1.

Lanfair, Anglesey,
Wales.

C. Oldham (c & p).

11.1.3.229-230.

6.

Plas Maclog, Anglesey.

T. A. Coward (c & p).

11. 1. 3. 234.

1.

Northenden, Cheshire,

G. Barrett-Hamilton

11. 1. 3. 237.

England.

(P)-

6, ? al.

Aberia, Merionethshire,

G. H. Caton Haigh

94. 6. 5. 1-2.

Wales.

(c & P).

16,39.

Aberia, Merionethshire.

G. Caton Haigh

11. 1. 3. 231.

(c & P).

253-256.

4?.

Langharne, Carmarthen-

W. E. de Winton

11. 1. 3. 257-260.

shire.

(c & P).

?.

Bridgend, Glamorgan-
shire.

R. I. Pocock (c & p).

11. 1. 3. 232.

6.

Swansea, Glamorgan-
shire.

C. Oldham (c & p).

11. 1. 3. 233.

9.

Waltham, Lincolnshire,

G. H. Caton Haigh

11. 1. 3. 213.

England.

(c & P).

lal.

Herefordshire.

Rev. S. 0. Ridley
(c & p).

82. 1. 28. 1.

1 al.

Bishopstone, Hereford-

Rev. S. 0. Ridley

81. 1. 1. 3.

shire.

(c & P).

6.

Worcestershire.

J. S. Elliott (c & p).

11. 1. 3. 214.

636 llODENTIA

(J, 9. Wormslev, Oxfordshire. W. R. Ogilvie-Grant 11.1.3.215-216.

(c & p).
9. Toddington, Bedford- Joseph S c r i v n e r 98. 2. 27. 1.

(albino) shire. (c & i>).

6. Welwyn, Hertfordshire. W. R. Ogilvie-Graut 11. 1. 3. 217.

(c & p).
2juv.al. Hillingdon, Middlesex. 0. Thomas (c & p). 80.10.14.1-2.
6, 3 9. Hillingdon, Middlesex. O. Thomas (c & p). 11. 1. 3. 219.

250-252.
6. Surrey. W. R. Ogilvie-Grant 11. 1. 3. 218.

(c & p).
9. Kingston, Surrey. C. H. B. Grant (p). 11. 1. 3. 404.

9. Susses. W. R. Ogilvie-Grant 11. 1. 3. 220.

(c & p).
2 9. Basingstoke, Hampshire. G. S. Miller (c). 7. 7. 7. 2943-4.

(7. 7. 7. 2944. Type of subspecies.)
2 6,9 9. Basingstoke, Hampshire. G. S. Miller (c). ' 7. 7. 7. 2947-55,

2976-7.
6, 9. New Forest, Hampshire. G. S. Miller (c). 7. 7. 7. 2969,

2937.
4 J, 4 9. Eversley, Hampshire. G. S. Miller (c). 7. 7. 7. 2939-40,

2956-7, 2972-
3, 2978-9.

2 6,2 9. Bembridge, I. of Wight. W. R. Ogilvie-Grant 11. 1. 3. 221-224.

(c & p).

9. St. Helens, I. of Wight. 0. Thomas (c & p). 11. 1. 3. 225.

juv. Salisbury, Wiltshire. E. R. Alston (p). 79. 9. 25. 59.

9. Pill-on-Avon, Somerset. R. I. Pocock (p). 11. 1. 3. 226.

3 (5, 3 9. Northlew, Devonshire. R. C. Wroughton (p). 11. 1. 3. 227

228, 261-264.

EvoTOMvs GLAREOLUS suEcicus Miller.

1900. Evotomys licrcynicus sueclcus Miller, Proc. Washington Acad. Sci.,
II, p. 101, July 26, 1900 (Upsala, Sweden). Type in U.S. National
Museum.

1910. Evotomys glarcolus succicus Trouessart, Faune Mamm. d'Europe,
p. 171.

Type locality. — Upsala, Sweden.

GeograjjJncal distribution. — Lowlands of Sweden, Finland and
south-eastern watershed of Norway.

Diagnosis. — Similar to Evotomys glareolus glareolus, but red
dorsal area narrower and less diffuse, and sides, face, and rump
more grey.

Colour. — ^The elements of the colour are essentially as in E.
glareolus glareolus, but the reddish brown is more strictly confined
to the back, and the grey element on sides is more conspicuous.

Skull and teeth. — The skull and teeth resemble those of the
typical race, except that third re-entrant angle on inner side of
m^ appears to be considerably less often present (see Table,
p. 631).

Measurements. — Type (young adult female from Upsala,
Sweden) : head and body, 96 ; tail, 42 ; hind foot, 18. Average
and extremes of six specimens from the type locality : head
and body, 94-3 (92-103); tail, 41 (38-43); hind foot, 16-9

EVOTOMYS

637

(IG-i-lT • 6). Average and extremes of eight specimens from
Eggedal, Buskerud, Norway : head and body, 97-5 (9'2-102-5) ;
tail, 54-6 (52-59); hind foot, 16-G (16-4-17). For cranial
measurements see Table, p. 650.

Specimens examined. — One hundred and four, from the following
localities : —

Norway: Asker, Christiania, 7 (B.M. and U.S.N.M.); Lillehammer,
Hedemarken, 18 (U.S.N.M.) ; Eggedal, Buskerud, 27 (U.S.N.M.); Spjosod,
Telemarken, 33 (U.S.N.M.) ; Molmen, Kristiansamt, 2 ; Holaaker, Kris-
tiansamt, 2.

Sweden : Kvickjock, Norbotten, 2 (B.M. and U.S.N.M.), identification
not positive; Upsala, 7 (U.S.N.M.); Medelpad, west Norrland, 2 ; Sma-
land, 2.

6, 9. Medelpad, W. Norrland, Lord Lilford (p). 8. 10. 19. 2-3.

Sweden.

2. Smalaud. Prof. Sundevall (c). 49. 11. 1. 3, 5.

1. Kvickjock, Norbotten. Prof. Sundevall (c). 45. 10. 25. 7.

2 £. Holaaker, 1900 ft. R. J. Cuninghame 98. 2. 28. 8-9.

(c & p).

2 S. Molmen. R. J. Cuninghame 98. 5. 2. 3-4.

(c & p).

S, 9. Asker, Christiania. Christiania Museum 93. 3. 1. 14-15.

(E).

EvOTOMYS GLARKOLUS ISTERICUS Miller.

1900. Evotomys hercynicus hercynicus Miller, Proc. Washington Acad. Sci.,
II, p. 100, July 26, 1900 (part ; specimens from Marxheim, Bavaria,
and Bustenari, Roumania).

1909. Evotomys glareolus istericus Miller, Ann. and Mag. Nat. Hist.,

8th ser., iii, p. 419, May, 1909. Type in British Museum.

1910. Evotomys glareolus istericus Trouessart, Faune Mamm. d'Europe

p. 172.

Type locality. — Bustenari, Prahova, Roumania (in the Car-
pathians, north-west of Bucharest ; altitude 480 m.).

Geograpliical distribution. — Drainage basin of the Danube,
from Bavaria through Hungary to Roumania and probably to
Bulgaria and the coast of the Black Sea.

Diagnosis. — Auditory bullse more abruptly inflated on inner
side than in the typical subspecies ; colour lighter and more
yellow than in the other races, the red of rather narrow dorsal
area a clear yellowish rufous.

Cokmr. — Summer pelage : dorsal stripe narrow and well
defined, not tending to spread over sides. It is rufovis slightly
varied with yellowish wood-brown, and rather thickly sprinkled
with black-tipped hairs. Face, cheeks, and sides pale yellowish
wood-brown, tinged with grey ; underparts varying from creamy
white to a yellowish cream-bufi". Rump like sides, rather
strongly contrasted with dorsal stripe. Feet greyish white. A
dark shade at ankle. Ears thinly haired, like dorsal stripe in
colour. Tail sharply bicolor, brown above becoming darker at
tip, soiled white below. Winter pelage : dorsal stripe slightly
less sharply defined than in summer, the rufous paler but warmer,

638 RODENTIA

considerably varied with wood-brown, but very inconspicuously
sprinkled with black-tipped hairs. Face, cheeks and sides more
yellowish wood-brown than in summer, and scarcely tinged with
grey. Rump slightly suffused with colour of dorsal area, and
therefore less contrasted with back than in summer. Feet
nearly pure white.

SlcuU and teeth. — The skull resembles that of the typical race
except that the auditory bullae are larger, their inner margins
more nearly approaching each other, and rising more abruptly
above level of basioccipital. Teeth as in E. glareolus glareohts ;
third upper molar with third re-entrant angle on inner side
present in about 64 per cent of fifty-six skulls examined.

Measurements. — T3^pe (adult male), from Bustenari, Roumania :
head and body, 96; tail, 43-5; hind foot, 17-4; ear from
meatus, 10 '5. Average and extremes of five adults from
Hatszeg, Hunyad, Hungary: head and body, 105 "4 (99-111);
tail, 45-4 (40-49); hind foot, 17-8 (17-4-18). Average and
extremes of ten adults from Marxheim, Bavaria, Germany :
head and body, 96-0 (93-103) ; tail, 46-0 (43-49); hind foot,
17*7 (17-18). For cranial measui'ements see Table, p. 651.

Specimens examined. — Sixty-six, from the following localities : — ■

Germany : Strass, near Burgheim, Bavaria, 5 ; Marxheim, Bavaria, 48
(B.M. and U.S.N.M.).

Austria-Hungary: Csall6koz-Somorja,Pressburg, Hungary,!; Hatszeg,
Hunyad, Hungary, 11.

BouMANiA : Bustenari, Prahova, 1.

'66. Strass, Burgheim, Lord Lilford (p). 8.10.19.6-8.

Bavaria. {Korbicz.)
5 6,5 9. Marxheim, Bavaria. Lord Lilford (p). 8. 10. 19. 9-18.

1. Csallokoz-Somorja, Press- Budapest Mus. (k). 94. 3. 1. 59.
burg, Hun gar V.
8 (5, 2 9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 2. 2. .35-44.

sylvania.
6. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 34.

sylvania.
6. Bustenari, Prahova, Rou- Lord Lilford (p). 4.4.6.72.

mania. {W. Dodson.) {Type of subspecies.)

EVOTOMYS GLA.EE0LUS NORVEGICUS Miller.

1900. Evotomys norvcgicus Miller, Proc. Washington Acad. Sci., ii, p. 93,

July 26, 1900. Type in U.S. National Museum.
1910. Evotomys norvegicus Trouessart, Faune Mamm. d'Europe, p. 166.

Type locality. — Bergen, Norway.

Geographical distribution. — Western Norway, north at least
to Nordland ; apparently confined to the western watershed.

Diagnosis. — Larger than Evotomys glareolus glareolus (hind
foot in adults, 18-4 to 19 mm., condylobasal length of skull,
24-2 to 26*2 mm.) ; skull noticeably more robust than in the
small races, the zygomata very abruptly spreading anteriorly ;
third upper molar usually with two re-entrant angles on inner

EVOTOMYS

639

side ; red dorsal area broad, not well defined from bufly grey
sides of body.

Colour. — Summer pelage : dorsal stripe dull ferruginous
slightly varied with light wood-brown and much darkened by a
uniform sprinkling of black-tipped hairs, fading rather abruptly
into the light wood-brown of face, cheeks and sides ; rump wood-
brown tinged with red in median line and forming no noticeable
contrast with colour of back ; underparts pale drab-grey, washed
to a varying degree with yellowish brown and irregularly
darkened by appearance at surface of slaty under colour ; feet
dull white, a dark shade at inner side of ankle ; tail sharply
bicolor, dark brown above, whitish below.

SJcull. — The skull is decidedly larger and more heavily built
than that of Evotomys glareolus ylareolus and the other small
races. Zygomata heavy, abruptly spreading antei'iorly, the
greatest zygomatic width at level of front of molar series.
Postorbital processes distinct but very small. Brain-case
moderately high and rounded, distinctly rectangular in outline
when viewed from above. Nasals squarely truncate posteriorly a
little in front of tips of nasal branches of premaxillaries.

Teeth. — The teeth are larger than those of true Evotomys
glareolus, and the third upper molar usually has only two
re-entrant angles on inner side. Otherwise they show no
peculiarities.

Measurements. — Type (adult male from Bergen, Norway) :
head and body, 108 ; tail, 57 ; hind foot, 18*6. Average and
extremes of eight adults from Graven, Hardanger, Norway :
head and body, 109-8 (104- 5-115) ; tail, 57-5 (55-60-5); hind
foot, 18-7 (18-4-19). For cranial measurement see Appendix.

Specimens examined. — Tv?eiity-six, from the following localities in
western Norway: Mo, Ranen, Nordland, 2 (U.S.N.M.) ; Saltdalen, Bodo,
Nordland, 3 ; SOndfjord, 1 ; Bergen, 4 (U.S.N.M.) ; Opheim, Bergen, 2 ;
Graven, Hardanger, 14.

3. Saltdalen, Bodo, Norway. Dr. R. CoUett (p). 84. 10. 31. 8 -10.

Skull. Sondfjord. Dr. R. Collett (p). 84. 10. 13. 12.

6, ?. Graven, Hardanger. U.S. Nat. Mus. (e). 5. 8. 5. 1-2.
(Miss Stejncger.)

Evotomys glaeeolus vASCoNiiE Miller.

1900. Evotomys vasconix Miller, Proc. Washington Acad. Sci., ii, p. 96,

July 26, 1900. Type in U.S. National Museum.
1910. Evotomys vasconix Trouessart, Faune Mamm. d'Europe, p. 168.

Type locality. — Montrejeau, Haute-Garonne, France.

Geographical distribution. — Pyrenees and region at their
immediate base (at present known from the French side only).

Diagnosis. — Size as in Evotomys glareolus norvegicus or slightly
greater (hind foot in adults, 18-6 to 19-6 mm., condylobasal

G40

RODENTIA

length of- skull, 25 to 26 mm.) ; third upper molar usually with
two re-entrant angles on inner side (see Table, p. 631) ; red of
dorsal area narrower and not so bright, and sides less huffy.

3Ieasuremenfs. — Type (adult male) : head and body, 107 ;
tail, 53 ; hind foot, 19. Adult female from Caterille, Haute-
Garonne, France : head and body, 97 ; tail, 44 ; hind foot, 18*8.
Adult male and female from I'Hospitalet, Ariege, France : head
and body, 112 and 110; tail, 54 and 60; hind foot, 19 -G and
18 '8. Two adult females from Porte, Pyrenees-Orientales,
France : head and body, 110 and 112 ; tail, 55 and 56 ; hind
foot, 19 and 19. For cranial measurements see Table, p. 654.

Spccime7is examined. — Thirty-four, from the following localities iu
France : Port6, Pyrenees-Orientales, 11 ; I'Hospitalet, Ariege, 5 ; Ax-les-
Thermes, Ariege, 4 ; Caterille, Haute-Garonne, G ; Luchon, Haute-Garonne, 1 ;
Montr6jeau, Haute-Garonne, 2 (U.S.N.M.); Bareges, Hautes-Pyr6n6es, 5.

.5,2?.

Port6, Pyr6n(^es-0rien-
tales, France.

G.

S. Miller (c).

8.

8.

4. 248-250.

2(5,9.

Port6, Pyr6n6es-Orien-

tales. {A. Robert.)

0.

Thomas (p).

8.

9.

1. 59-Gl.

i, 9.

L'Hospitalet, Ariege.
(.4. PMhert.)

0.

Thomas (p).

8.

9.

1. G2-63.

2 S.

Ax-les-Thermes, Ariege.

V.

Builles (p).

8.

3.

27. 12-13.

6, 9.

Ax-les-Thermes, Ariege.

G.

S. Miller (c).

8.

8.

4. 252-253.

5 6,9.

Caterille, Haute-
Garonne. (^4. Robert.)

0.

Thomas (p).

6.

4.

1. 87-92.

9.

Luchon, Haute-
Garonne. (A. Robert.)

0.

Thomas (p).

G.

4.

1.93.

9.

Bareges, Haute s-
Pyreness.

G.

S. Miller (c).

8.

8.

4. 251.

EVOTOMYS GLAREOLUS HELVETICUS Miller.

1900. Evotoynys hercynicus hclveticus Miller, Proc. Washington Acad. Sci.,
II, p. 98, July 26, 1900. Type in British Museum.

1910. Evotomys glarcolus helveticus Trouessart, Faune Mamm. d'Europe,
p. 171.

Type localiti/. — Montauban, Haute-Savoie, France (near
Geneva, Switzerland). Altitude, 900 m.

Geographical distribution. — Jura Mountains ; south through
the non- Alpine parts of Switzerland and along the lower western
portion of the French Alps.

Diagnosis. — Intermediate between Ewtomys glareolus glareolus
and E. g. nageri in size (hind foot, 17 to 19 mm., condylobasal
length of skull, 23 to 25-4 mm.), but colour somewhat paler and
more buffy greyish than in eithei-.

Measurements. — Type (young adult male) : head and body, 95 ;
tail, 45 ; hind foot, 18 ; ear, 15. Adult male and female from
the type locality : head and body, 100 and 101 ; tail, 43 and 51 ;
hind foot, 19 and 19. Average and extremes of ten adults from
Cranves-Sales, Haute-Savoie : head and body, 101 (98-105) ; tail,
49-2 (48-52); hind foot, 18-3 (17-6-^19). Average and

EVOTOMYS

641

extremes of ten adults from Barcelonnette, Basses- Alpes, France :
head and body, 110-4 (103-117) ; tail, 55-3 (49-58) ; hind foot,
18-5 (17 • 6-18* 6). Five adults from Geneva, Switzerland:
head and body, 114-4 (103-122) ; tail, 47-8 (45-53); hind foot,
17-9 (17-18). Seven adults from St. Cergues, Vaud, Switzer-
land: head and body, 107 (101-111); tail, 46 (42-50); hind
foot, 17-8 (17-4-18-2). For cranial measurements see Table,
p. 654.

Specimens examined. — One hundred and twenty-nine, from the following
localities : —

Switzerland : St. Cergues, Vaud, 27 (U.S.N.M. and Mottaz) ; Geneva,
23 (U.S.N.M. and Mottaz).

France : Montauban, Haute-Savoie, 21 ; Lucinges, Haute-Savoie, 12 ;
Cranves-Sales, Haute-Savoie, 12 ; Scieutriers, Haute-Savoie, 8 (Mottaz) ;
Barcelonnette, Basses-Alpes, 15 (B.M. and U.S.N.M.) ; Saint Paul, near
Barcelonnette, Basses-Alpes, 11.

6. Montauban, Haute- O. Thomas (p). 2.6.7.1.

Savoie,900m. Prance. (T^^^e of subspecies.)

(A. Robert.)
6 6,5 9. Cranves - Sales, Haute- 0. Thomas (p). 5. 11. 18. 33-4-3.

Savoie. (A. Robert.)
6, 9. Barcelonnette, Basses- U.S. Nat. Mus. (e). 5. 8. 5. 3-4.
Alpes.
4c5, 2 9. St. Paul, Basses-Alpes. 0. Thomas (p). 8.8.10.116-121,

(C. Mottaz.)

EvOTOMYS GLAEEOLUS NAGERI Schinz.

1845. Hypludxus] nageri Schinz, Synops. Mamm., ii, p. 237 (Oberalpsee,

Urserenthal, Uri, Switzerland).
1857. Arvicola glareolus b. Blasius, Siiugethiere Deutschlands, p. 337.
1862. Myodes bicolor Fatio, Eev. et Mag. de ZooL, 2nd ser., xiv, p. 257,

July, 1862 (Genthal, Berne, Switzerland). Type in Geneva

Museum.
1900. Evotomys nageri Miller, Proc. Washington Acad. Sci., ii, p. 94,

July 26, 1900.
1910. Evotomys nageri Trouessart, Faune Mamm. d'Europe, p. 167.

Tyjpe locality. — Oberalpsee, near Andermatt, Uri, Switzerland.

Geographical distrihution. — -Alps (except westernmost portion)
and mountains of northern Italy.

Diagnosis. — Size slightly larger than in Evotomys norvegicus
(usual measurements of adults: hind foot, 18-8 to 20 mm.;
condylobasal length of skull, 25 to 26-2 mm.) ; third upper
molar usually with three re-entrant angles on inner side ; dorsal
red area narrow and dark, the sides of body dull greyish in
evident though not conspicuous contrast.

Colour. — Dorsal stripe rather well detined, extending from
forehead to rump, but not showing much tendency to spread
laterally. It is cinnamon-rufous slightly varied with pale
broccoli-brown and inconspicuously darkened by a sprinkling of
black-tipped hairs ; the resulting colour approaching chestnut, or
something lighter and more yellowish. Face, cheeks and sides

2 T

642 RODENTIA

light hair-brown fading to smoky grey on lower part of sides.
Rump light broccoli-brown tinged with reddish brown in median
line, and forming in most specimens a distinct though not very
noticeable contrast with dorsal stripe. Whole ventral surface
pale smoke-grey, washed to a varying degree with yellowish
brown, especially along median line, the slate-grey bases of the
hairs appearing irregularly at surface. Feet dull white ; a dark
shade at inner side of ankle. Ears thinly haired, dull reddish
brown. Tail bicolor, dark brown above, buffy whitish below.
The winter pelage tends to be darker than the summer pelage,
with less contrast between colours of dorsal stripe and sides ; but
dull, brownish individuals are not infrequently met with in
summer.

SJmU and teeth. — Skull largest among the Continental Euro-
pean members of the group, iJ. glareolus hallucalis perhaps excepted.
In form it shows no sjjecial peculiarities. Brain-case relatively
large, a peculiarity moi'e noticeable in smooth, young-adult skulls
than in those which have become angular with advancing age.
Zygomata not very abruptly spreading in front, the middle
portion of the two arches usually almost parallel. Incisive
foramina narrow (normal), the width of the two together about
one-quarter length. Molars distinctly larger than in E. glareolus
glareolus, but pattern of enamel folding with no other peculiarity
than that the third re-entrant angle on inner side of m^ is usually
present (see Table, p. 631).

Measurements. — Adult male from Les Plans, Vaud, Switzer-
land : head and body, 105; tail, 65; hind foot, 19*8. Average
and extremes of four adults from Briinig, Switzerland : head and
body, 105-7; tail, 61-7; hind foot, 19-3 (18-8-20). Average
and extremes of ten adults from Andermatt, Uri, Switzerland :
head and body, 106-3 (101-112); tail, 64*5 (59-72); hind foot,
18-8 (18-6-19). Average and extremes of four adults from
Goschenen, Uri, Switzerland: head and body, 117-7 (114-123);
tail, 64-2 (62-68;; hind foot, 19-6 (19-20). Two adult males
from Gordola, Locarno, Ticino, Switzerland : head and body,
103 and 103; tail, 62 and 65; hind foot, 19-6 and 19-3; ear
from meatus, 13-5 and 13. Adult male and female from
Lugano, Ticino, Switzerland : head and body, 106 and 107 ; tail,
48 and 49 ; hind foot, 19-8 and 20. For cranial measurements
see Table, p. 653.

Specimens exawmed.^Oue hundred and seventy-five, from the following
localities : —

Prance : Chamonix, Haute-Savoie, 8 (U.S.N.M.).

Switzerland: Les Plans, Vaud, 7 (U.S.N.M.) ; Zermatt, Valais, 10
(U.S.N.M.) ; Briinig, Berne, 20 (U.S.N.M.) ; Meiringen, Berne, 11 (U.S.N.M.) ;
Genthal, Berne, 1 (Geneva ; type of bicolor Fatio) ; Lucerne, 2 ; Vitznau,
Lucerne, 10 (B.M. and U.S.N.M.) ; Goschenen, Uri, 8 (U.S.N.M.) ; Ander-
matt, Uri, 49 (U.S.N.M.); Vulpera-Tarasp, Grisons, 14 (Rothschild);
Campfer, Grisons, 5 (Rothschild) ; Untervatz, Grisons, 3 ; Poschiavo,
Grisons, 1 (Mottaz) ; Faido, Ticino, 1 (U.S.N.M.) ; Lugano, Ticino, 5
(U.S.N.M. and Mottaz) ; Gordola, Locarno, Ticino, 3.

EVOTOMYS

643

Italy : Padola, Cadore, 1 (Turin) ; Casamazzagno, Cadore, 1 (Turin) ;
Ceresole d'Alba, Turin, 2 ; Stupiuigi, Turin, 1 (Turin) ; INIoncalieri, Turin,
9 (U.S.N.M. and Turin) ; Ambonasco, in mountains above Chiavari,
Liguria, 2 (Genoa) ; no exact locality, 1.

6,4:9.

Vitznau, Lucerne, Swit-
zerland.

0. Thomas (c & i>).

5. 8. 3. 22-2C.

3 6.

Untervatz, Grisons.

Lord Lilford (p).

1. 11. 7. 15-17.

2 6,9.

Locarno, Ticino.

0. Thomas (c & p).

5. 8. 2. 15-17.

6, 9.

Turin, Italy.

Dr. E. Festa (c & p).

8. 8. 1. .3-4.

1.

Italy.

Purchased (Brandt).

53. 11. 19. 2, 4.

EvOTOMYS GLAREOLUS HALLUCALIS ThomaS.

1882. Arvicola glarcolus Cavanna, Bull. Soc. Entomol. Ital., xiv, p. 87

(Serra Crispo, Monte PoUino, Basilicata, Italy).
1906. Evotomys nageri haUucalis Thomas, Ann. and Mag. Nat. Hist

7th ser., xviii, p. 221, September, 1906. Type in British Museum
1910. Evotomys nageri haUucalis Trouessart, Faune Mamm. d'Europe,

p. 168.

Type locality. — Santa Eufemia d'Aspromonte, Calabria, Italy.

Geographical distribution. — High mountains of southern Italy.

Diagnosis. — Externally similar to Evotomys glareolus nageri,
but underparts less washed with yellowish brown ; skull with
longer, narrower brain-case and rather shorter rostrum than in
nageri, the incisive foramina shorter and wider than in the
related forms ; teeth larger than in any other European species
except E. csesarius and E. rufocanus.

Colour. — Except that the underparts are a light grey washed
with whitish cream-buif the colour is similar to that of Evotomys
glareolus nageri. Line of demarcation along sides ill defined.
Feet dull whitish, somewhat lighter than usual in E. g. nageri.

Skull. — Although the only known skull is somewhat imperfect
its form appears to be notably different from that of Evotomys
glareolus nageri and the other members of the group in the
decided elongation of the brain-case and a slight shortening of
the rostrum. Length of brain-case measured from back of inter-
parietal to line joining tips of postorbital processes nearly equal
to zygomatic breadth, instead of decidedly less than zygomatic
breadth as in the related large forms. Interorbital region
rather wide and smooth, a character probably due in part at
least to the comparative youth of the specimen (roots of molars
less tham 1 mm. long). Rostrum relatively shorter than in
E. g. nageri, and incisive foramina much shorter and wider, the
greatest breadth of the two together decidedly more than one-
third length of foramen instead of about one-fourth as in the
related species.

Teeth. — The incisors are shghtly more slender than usual in
Evotomys glareolus nageri, though a few specimens from the Alps
are essentially the same. Molars heavier than in E. g. nageri,
though not peculiar in form, the length of the upper series more

2 T 2

644 RODENTIA

nearly approaching length of diastema than in any other European
species except the similarly heavy-toothed E. csesarius. Third
upper molar with well developed third inner re-entrant angle.

Measurements. — Type (young adult male) : head and body,
115; tail, 66; hind foot, 21 ; ear from meatus, 13. For cranial
measurements see Table, p. 653.

Specimen examined. — The type.

Bemarhs. — Although at present very imperfectly known the
south Italian Evotomys appears to be a well-characterized race.

Aspromonte, Calabria. (A. Robert.) 0. Thomas (p). 6. 8. 4. 9.

{Type of subspecies.)

EVOTOMYS SKOMERENSIS Barrett-Hamilton.

1903. Evotomys skom,erensis Barrett-Hamilton, Proc. Royal Irish Acad.,

XXIV, p. 316. Type in British Museum.
1910. Evotomys sliomerensis Trouessart, Faune ]\Iamm. d'Europe, p. 167.

Type locality. — Skomer Island, off coast of Pembrokeshire,
Wales.

Geographical distribution. — Skomer Island.

Diagnosis. — Size as in the other large European members of
the genus ; tail about half as long as head and body ; colour
above unusually light and bright, sharply and conspicuously
contrasted with buffy white of underparts ; skull about as large
as that of E. nageri, but with brain-case unusually ridged and
angular, and nasals decidedly longer than diastema, rather
abruptly narrowed near middle.

Colour. — Winter pelage : upper parts a bright light reddish
brown approaching the orange-rufous of Ridgway, a narrow area
on sides paler and with a distinct buffy suffusion ; underparts
and entire fore leg strongly contrasted whitish grey, faintly
tinged with pale buff (the combination much paler than the
cream-buff of Ridgway), the line of demarcation along sides well
defined ; tail sharply bicolor, brownish above, pale cream-buff
below ; feet buffy white above, pale hair-brown on furred portion
of sole. Summer pelage not known.

Sktill. — The skull differs notably from that of Evotomys
glareolus nageri and E. g. norvegicus in the short, broad, rather
strongly ridged and angled brain-case, the conspicuous develop-
ment of the mastoid region, which stands out more strongly from
general outline of brain-case than in any of the related species,
and the unusual length of the nasals, which distinctly exceed
the diastema. Instead of diminishing gradually in width from
before backward, as in the related species, the nasals contract
rather abruptly near middle, so that their general outline is
almost spatulate. The nasals are more abruptly bent downward
anterioi'ly than in E. glareolus nageri and E. g. norvegicus.
Otherwise the dorsal profile is nearly flat, like that of the Alpine

EVOTOMYS 645

animal. The depth of the entire skull from occipital region to
anterior base of zygomata is relatively greater than in any of the
other European species except E. cdesarms. Postorbital processes
small but unusually prominent and well defined, a well developed
ridge extending obliquely backward and upward from each
process nearly to anterior edge of parietal. Rostrum and
incisive foramina normal. Zygomata rather abruptly flaring,
about as in _B. glareolus norvegicus. Auditory bullae relatively
larger than in the large forms of Evotomys glareolus.

Teeth. — The teeth are essentially like those of Evotomys
glareolus nageri ; thii'd upper molar normally with three re-
entrant angles on inner side (see Table, p. 631).

Measurements.- — Type (adult male): head and body, 108;
tail, 59; hind foot, 18; ear from meatus, 13. Average and
extremes of ten adults: head and body, 109 "7 (105-121); tail,
55-5 (50-61); hind foot, 18-4 (18-19); ear from meatus,
13*5 (13-14). For cranial measurements see Table, p. 656.

Specimens examined. — Thirteen, all from Skomer Island (B.IM. and
U.S.N.M.).

5 (5, 2 9. Skomer Island, S.W. Y. H. Mills (c & p). 3. 7. 4. 1-7.
Wales. (3. 7. 4. 3. Type of species.)

1 al. Skomer Island. R. Drane (c & p). 4. 8. 5. 1.

2. Skomer Island. E. Drane (c & p). 11. 1. 3. 265-266.

EVOTOMYS CiESARIUS Miller.

1896. Evotomys glareolus Barrett-Hamilton, The Zoologist, 3rd ser., xx,

p. 98, March, 1896 (Jersey, Channel Islands).
1908. Evotomys cxsarius Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 195, February, 1908. Type in British Museum.
1910. Evotomys cxsariiis Trouessart, Faune Mamm. d'Europe, p. 169.

Tijpe locality. — St. Helier, Jersey, Channel Islands.

Geographical distribution. — ^Known only from the island of
Jersey, Channel Islands.

Diagnosis. — General size as in the other large European
forms, but tail distinctly less than half as long as head and body,
and ear unusually shortened ; colour dark and rather dull ;
essentially no line of demarcation on sides ; skull larger and
more massive than in any of the Continental members of the
genus, E. rufocanus not excepted.

Colour. — Winter pelage : upper parts a rich, dark, reddish
brown, approaching the cinnamon-rufous of Ridgway, but not so
vivid, the sides of body and outer surface of fore leg lighter and
suffused with dull buff, though not sufficiently to produce any
marked contrast with back ; underj^arts a clear rich buff (between
the buff and cream-buff of Ridgway), the line of demarcation
along sides ill-defined ; tail sharply and conspicuously bicolor,
blackish above, concolor with belly below ; feet a dusky grey
above, rather dark brown on furred portion of sole. Summer
pelage : red area restricted to back, and noticeably browner and

646

RODENTIA

duller than in winter, the exact shade nearly hazel ; sides bioccoli-
brown, in rather strong contrast with back, but fading insensibly
into the dull buft" of underparts. Tail and feet as in winter.

Shidl. — The skull is larger than that of any of the other
European Evotomys* In form it differs from the others in its
greater depth, more convex dorsal profile, and especially in the
greater angle at which the nasals are bent downward. In the
last character it is approached by E. sJcomerensis ; but the dorsal
surface of brain-case is not flattened as in the Skomer vole.
Interorbital region wide, not tending to assume a cylindrical
form as in E. rufocanits, the lateral ridges noticeable in old age
but remaining wide apart, with a broad trough-like median
groove. Brain-case relatively shorter and broader than in the
Continental species, but less ridged and angular than in
E. sJcomerensis. Postorbital processes rather large, but less
Microtus-like than in E. rufocanus and with no trace of a ridge
extending obliquely backward toward parietal such as occurs in
E. sJcomerensis. Nasals about as long as diastema, moderately
spatulate in outline. Rostrum more robust than in any of the
other European species ; incisive foramina normal. Auditory bullae
large, not peculiar in form. Zygomata not very abruptly spread-
ing, the median portion of the two arches parallel.

Teetli. — Except for their larger size the teeth are essentially
as in Emtomys glareolus. Third upper molar with third inner
re-entrant angle normally present (see Table, p. 631). Anterior
loop of wij very short.

Measurements. — External measurements of type (not fully
adult male) : head and body, 96; tail, 49; hind foot, 18 ; ear
from meatus, 1 1 . Average and extremes of ten adults : head
and body, 111-5 (107-115); tail, 50-9 (48-58) ; hind foot, 19-3
(18-5-20); ear from meatus, 11-4 (11-12). For cranial
measurements see Table, p. 657.

Specimens examined. — Eighteen, all from the island of Jersey.

2 6. St. Helier, Jersey. G. Barrett-Hamilton 3. 2. 11. 1-2.

(c & p). (3. 2. 11. 2. Typeoi species.)

5 (5, 2 ?. Trinity, Jersey. O. Thomas (p). 8. 9. 2. 12-17.

[R. H. Bunting.)

EVOTOMYS RUTILUS Pallas.

1778. M'us rutilus Pallas, Nov. Spec. Quadr. Glir. Ord., p. 246.

1874. Evotomys rutilus Goues, Proc. Acad. Nat. Sci., Philadelphia, p. 187.

1900. Evotomys rutilus Miller, Proc. Washington Acad. Sci., n, p. 91,

July 26, 1900.
1910. Evotomys rtdihis Trouessart, Faune Mamm. d'Europe, p. 169.

Ty^pe locality. — Siberia east of the Obi.

GeofjrajjJiical distribution. — Arctic Asia and Europe, south in
Norway to Tromso, and in Sweden to Norbotten.

* With the possible exception of E. glareolus hallucalis, adults of which
have not been examined.

EVOTOMYS 647

Diagnosis. — Size medium (hind foot about 18 mm. ; condylo-
basal length of skull, 25 mm.) ; tail noticeably less than half as
long as head and body, very densely haired, its pencil at least
one-quarter as long as vertebraj ; dorsal area clear bright chest-
nut, sharply but not conspicuously contrasted with colour of
sides.

Colour. — Dorsal stripe a clear bright reddish brown approach-
ing the chestnut of Ridgway, faintly darkened by a slight
admixture of black-tipped hairs. Sides and face ochraceous-buff.
Underparts cream-colour or dull whitish, darkened by the slate-
grey bases of the hairs, which show through irregularly at
surface. Feet whitish. Ear concolor with dorsal stripe. Tail
sharply bicolor, brownish tinged with red above, dirty white below.

Sktdl. — In general the skull resembles that of Evotomijs
glareolus. Brain-case shorter and more squarish in outline when
viewed from above, its upper surface, together with that of
rather wide interorbital region somewhat flattened. Postorbital
processes obsolete. Palate and mesopterygoid space relatively
narrower than in E. glareolus ; posterior termination of palate
usually not an entire-edged shelf, since the lateral groove on each
side habitually opens freely into posterior nares, thus isolating
median portion of shelf from the outer extremities adjoining
alveoli.

Teeth. — Molars relatively smaller and weaker than in Evotomijs
glareolus. Enamel pattern essentially as in E. glareolus but
re-entrant angles tending to be deeper, those of the opposite
sides more conspicuously alternating, and salient angles more
acute, so that inner triangles of maxillary teeth
seldom if ever assume a circular outline. An-
terior outer re-entrant angle of m^ fully as
deep as second, the anterior portion of the
tooth therefore never assuming the Alticola-\\\ie
aspect characteristic of that region in E. glareo-
lus ; inner side of m^ always with three deep
sub-equal re-entrant angles. Second lower molar
nearly or quite divided into four alternating
closed triangles and a posterior closed loop ; fiq. 126.

W3 showing a decided tendency to assume a Evotomys rutuu,.
similar form owing to the depth and distinct Enamel pattern, x 5.
alternation of its re-entrant angles.

Measurements. — External measurements of an adult from
Karesuando, Sweden (from well-made skin) : head and body, 98 ;
tail, 23 ; pencil, 12 ; hind foot, 18. Adult from Lappmark,
Sweden (no exact locality) : hind foot, 17. For cranial measure-
ments see Table, p. 657.

Specimens examined. — Seventeen, from the following localities : —
Norway: Sydvaranger, 3 (U.S.N.M.) ; Porsanger, 2 (U.S.N.M.).
Sweden: Lapland, no exact locality, 7 (B.M. and U.S.N.M.); Tornea,
Lappmark, 3 (B.M. and U.S.N.M. ; Karesuando, Norbotten, 2 (U.S.N.M.).

648 RODENTIA

Bemarhs. — Externally Evotomys rutilus is distinguishable from
the other European members of the genus by its short, abun-
dantly-haired tail, and by the bright colour of its dorsal area.
The weak molar rows and the peculiarities of the third upper
molar are also diagnostic. Notwithstanding the weakness of the
teeth the pattern of the enamel folding is characterized by
unusually definite angularity and general firmness of line.

1. Tornea, Lapland. Prof. Sundevall (p). 45. 10. 25. 9.

2. Lapland. Prof. Sundevall (p). 49. 11. 1. 7-8.
skull. Varanger Fjord, Fin- Dr. R. Collett (p). 84. 10. 31. 5.

mark.

EVOTOMYS RUFOCANUS Sundevall.

1846. Hyimdfeus rufocanus Sundevall, Ofversigt af Kongl. Vetensk. Akad.

Forhandl., iii, p. 122.
1897. Evotomys rufocamis Bailey, Proc. Biol. Soc, Washington, xi, p. 122,

May 13, 1897.
1900. Evotomys mfocanns Miller, Proc. Washington Acad. Sci., ii, p. 89,

July 26, 1900.
1910. Evotomys (Craseomys) rufocamis Trouessart, Faune Mamm. d'Europe,

p. 173.

Type locality. — Lappmark, Sweden.

Geographical distribution. — Northern Europe, south in the
mountains of Norway to Dovre, east into Asia.

Diagnosis. — Form essentially as in Evotomys glareolus, neither
tail nor ear shortened ; size large, as in E. csesarius (hind foot,
18 to 19, condylobasal length of skull, 26-6 to 27*6 mm.) ; skull
and teeth heavy, the upper molars with salient angles more
pointed than in the other European members of the genus,
E. rutilus excepted ; m^ normally with only two re-entrant angles
on inner side ; dorsal stripe narrow, conspicuously contrasted
with grey of sides.

Colour. — Dorsal stripe well defined, narrow, extending from
between eyes nearly to base of tail. Its colour is dark but
rather bright, usually intermediate between hazel and cinnamon-
rufous, inconspicuously sprinkled with black-tipped hairs. Sides
a clear light grey formed by an intimate blending of hair-brown,
whitish, black, and slate-colour. Whole underparts soiled buffy
white, irregularly dulled by the slate-grey under colour. Cheeks
and muzzle similar to sides but slightly darker. Ear like dorsal
stripe. Feet dirty white. Tail sharply bicolor, brownish
above, dirty white below. In immature specimens the colour is
dull, the dorsal stripe bister scarcely tinged with red, the sides
less conspicuously grey.

Skull. — The skull exceeds in size and massiveness that of any
of the forms of Evotomys glareolus, in this respect essentially
agreeing with that of E. csesarius. Interorbital region narrower
and more cylindrical than in any of the related species, zygomata
heavier and more abruptly and conspicuously expanded in front

EVOTOMYS

649

of middle, and postorbital processes tending to spread laterally
along posterior border of orbit as in Microtus rather than to
assume the bluntly peg-like form characteristic of Evotomys.
Brain-case rather long, the occipital region extended backward
by the unusually broad interparietal. Basioccipital narrow as
in E. glareolus, not widened as in the large E. csesarius, its width

Fig. 127.
Evotomys rufocamis.

Nat. size.

along anterior suture equal to about one-third length from suture
to foramen magnum. Auditory bullse rather large, slightly more
narrowed antero-internally than in E. glareolus. Palate rather
narrow but not peculiar in form. Anteorbital foramina relatively
smaller (narrower) than in any of the other European members
of the genus. Mandible without special peculiarities.

Teeth. — Relatively to size of skull the teeth are larger and
heavier than in the other European members
of the genus, a peculiarity which is to the
eye exaggerated by the contrast with narrow
palate. The enamel pattern of the molars
agrees with that of Evotomys glareolus in
number and arrangement of the elements ;
m^ normally with only two re-entrant angles
on inner side (see Table, p. 631), and with
first outer re-entrant angle tending to be
deep, as in E. rutilus. It is peculiar, how-
ever, in a certain tendency to acuteness of
the salient angles, a character especially
noticeable in not fully adult individuals, and
in the constancy with the triangles of all
three upper molars and of 7h, become completely closed.
Anterior loop of m^ with inner reentrant angle relatively deeper

Emtomys rufocamiH.
Enamel pattern. X 5.

650

RODENTIA

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than in E. glareobis, so that main axis of loop appears to curve
inward instead of outward. Roots of molars developing later
in life than in the other European species, the position of the
inner extremity of upper teeth indicated by an evident capsule.
Basal portion of m^ usually well outlined on inner surface of
mandible.

Measurements. — External measurements of adult male from
Lappmark, Sweden: head and body, 114; tail, 37; hind foot
(dry), 18*4. Adult from Kvickjock, Sweden (well-made skin):
head and body, 110; tail, 40; hind foot, 18, Eor cranial
measurements see Table, p. 657.

Specimens examined. — Twenty-four, from the following localities : —
Norway : Varangerfjord, 1 ; Sudvaranger, 1 (U.S.N.M.) ; Stabursnaes,
Porsanger, 2 (B.M. and U.S.N.M.) ; Kishand, Porsanger, 1 ; Porsanger, 1
(U.S.N.M.) ; Molmen, Kristiansamt, 1 ; Sundal Fjeld, 1 ; Hyllingen,
Trondhjem, 1 ; Tanen, Finnmarken, 2 (U.S.N.M.).

Sweden : Karesuando, Norbotten, 3 (B.M. and U.S.N.M.) ; Lappmark
(no exact locality), 1 ; Tornea, Lap.pmark, 2 (U.S.N.M.) ; Kvickjock, Nor-
botten, 6 (B.M. and U.S.N.M) ; no exact locality, 1.

Remarks. — Evotomys rufocanus is so readily distinguishable
from the other European members of the genus as to require no
special comparison. So different is the animal from the better
known species that it has been made the type of a special sub-
genus Craseomys. The discovery of additional species, notably
Evotoynys csesarius, intermediate in character between true
Emtomys and Craseomys, and the fuller comprehension of the
dental peculiarities of Evotomys ruiilus, make such a division of
the genus Evotomys appear to be no longer warranted.

1 skull. Varangerfjord, Finmark, Dr. R. Collett (p). 84. 10. 31. 4-5.
Norway.

6. Stabursnaes, Porsanger. Dr. R. Collett (p). 84. 10. 31. 7.

i skull. Kishand, Porsanger. Dr. R. Collett (p). 84. 10. 31. 6.

6. Molmen, Kristiansamt. R. J. Cuninghame 98. 5. 2. 5.

(c & p).

Sundal Fjeld. E. Lort Phillips 94. 10. 22. 2.

(c & p).

9. Hyllingen, Trondhjem. G. Barrett-Hamilton 11. 1. 2. 114.

6. Karesuando, Norbotten, Stockholm Museum 90. 8. 1. 15.

Sweden. (e).

1. Kvickjock, Norbotten. Prof. Sundevall (p). 49. 11. 1. 11.

S. Lappmark. (G. Kolthoff.) Lord Lilford (p). 11. 1. 1. 153.

1. Sweden. " Stockholm Museum 46. 6. 2. 70.

(E).

Genus MICROTUS Schrank.

(Synonymy under sub-genera.)

Geographical distribution. — Northern portions of northern
hemisphere south in America to southern Mexico and in Eui-ope
to the Mediterranean coast.

MICROTUS 659

Characters. — Bony palate terminating posteriorly in a sloping
median ridge and two lateral pits ; lower incisor with root
extending to outer side of molar-roots, rising high above level of
cutting surface of molars, and forming a noticeable protuberance
on outer surface of ascending portion of mandible at base of
articular process ; molars permanently rootless, growing con-
tinuously from a persistent pulp and not wearing away in old
age ; pattern of enamel folding characterized by acuteness of
salient angles ; mammjv, 8 ; plantar tubercles, 6 ; neither fur,
external form, nor skull modified for aquatic or subterranean life.

Itemarlts. — The genus Microtus as here understood, restricted
to the species with normal skull, palate and enamel folding,
8 mammiie, 6 plantar tubercles, and no special modifications of
external form, is the largest and most widely distributed group
in the sub-family. About 100 species are now recognized, a
number which will be materially increased as the fauna of the
interior of Asia becomes better known. Fifteen species are here
recognized as occurring in western Europe. Among them are
represented two sub-genera.

Sub-Genus MICROTUS Schrank.

1798. Microtus Schrank, Fauna Boica, i, p. 72 {Microtus terrestris Schrank

= Mjis arvalis Pallas).
1817. Mynomes Rafinesque, Amer. Monthly Magazine, ii, p. 45 (Mynomes

pratensis Rafinesque = Arvicola pennsylvanicus Ord).
1836. Heniiotomys de Selys-Longchamps, Essai monographique sur les

Campagnols des environs de Li6ge, p. 7. Part {arvalis and

terrestris = scherman).
1857. Paludicola Blasius, Saugethiere Deutsclilands, p. 333. Part {amphi-

bius = scherman, nivalis and ratticeps). Not of Wagler, 1830.
1857. Agricola Blasius, Saugethiere Deutschlands, p. 334 {agrestis).
1867. Praticola Fatio, Les Campagnols du Bassin du L6man, p. 34

(included ampJiibius — scherman, nivalis, arvalis, ratticeps and

campestris = agrestis). Not of Swainson, 1837.
1867. Sylvicola Fatio, Les Campagnols du Bassin du L6man, p. 63

{agrestis). Not of Harris, 1782.
1883, Microttis Lataste, Le Naturaliste, ii, p. 348.
1890. Cavipicola Schulze, Schriften Naturwiss. Viereins d. Harzes in

Wernigerode, v, p. 24. Part {arvalis, subterraneus and campestris

= agrestis).
1894. Tetramerodon Rhoads, Proc. Acad. Nat. Sci., Philadelphia, p. 282

{Arvicola tetramerus Rhoads).
1896. Microtus Miller, North American Fauna, No. 12, p. 62, July 23, 1896.
1899. Euarvicola Acloque, Faune de France, Mammiferes, p. 49 {agrestis).

Type species. — Microtus terrestris Schrank = Mus arvalis
Pallas.

Geographical distribution. — Northern portions of both hemi-
spheres, south in America to southern Mexico, and in the Old
World to Portugal, central Spain, the Mediterranean coast of

2 u 2

660

RODENTIA

France, northern Italy, the central poi-tion of the Balkan Penin-
vsula, Asia Minor and northern India.

Diagnosis. — Principal characters as in the genus ; third upper
molar with three re-entrant angles on inner side ; skull with
brain-case narrow and high, or broader and more flattened, its
surface ridged and angular ; interorbital region narrow, the
temporal ridges well developed ; elements of posterior edge of
palate well delined.

Hemarhs. — The sub-genus Microtus contains most of the
members of the genus. Twelve species are now known to occur
in western Europe.

KEY TO EUROPEAN FORMS OP THE SUB-GENUS MICROTUS.

Second upper molar with two closed triangles on

inner side M. agrestis, p. 662.

Condylobasal length of skull in largest individuals
27 to 28-4 mm.
First upper molar with third inner triangle
usually present; underparts heavily washed

with wood-brown (Hebrides) M. a. exsul, p. 669.

First upper molar with third inner triangle
usually absent ; underparts slightly or not
washed with wood-brown.
Brain-case relatively long, the distance from
, condyle to back of interorbital constric-
tion decidedly greater than zygomatic
breadth (Alps and their immediate
neighbourhood ; marshes near mouth

of Rhone) M. a. levemedii, p. 671.

Brain-case relatively short, the distance from
condyle to back of interorbital constric-
tion about equal to zygomatic breadth

(Scandinavia) M. a. agrestis, p. 668.

Condylobasal length of skull in largest individuals

24-6 to 26-6 mm.

Auditory bulla small and flattened, its greatest

diameter (from paroccipital process to tip

of anterior spine) contained about 3^ times

in condylobasal length of skull (Portugal

and north-western Spain) M. a. rozianus, p. 680.

Auditory bulla large and inflated, its greatest
diameter contained about 3 times in condy-
lobasal length of skull.
Condylobasal length of skull in largest indi-
viduals 25 to 26 mm. (England and

southern Scotland) M. a. hirtm,-g. 673.

Condylobasal length of skull in largest indi-
viduals 25 "4 to 26*6 mm.
General colour of upper parts darker,
usually with decided tinge of russet

(Northern and central Scotland) .1/. a. neglectm, p. 675.

General colour of upper parts lighter,

usually with no decided tinge of russet

(West-central Continental Europe).... M. a. bailloni, p. 672.

Second iipper molar with one closed triangle on

inner side.

First lower molar with three re-entrant angles

on outer side M. ratticeps, p. 708.

MICROTUS 661

First lower molar with four or five re-entrant
angles on outer side.
Nasals moderately cuneate, their combined
breadth posteriorly a little more than half
their greatest breadth ; diastema notice-
ably shorter than nasals.
Condylobasal length of skull about 27 mm. ;
upper tooth-row about 7 mm. , the teeth

not unusually robust (Central Spain) M. cabrern', p. 701.

Condylobasal length of skull about 30 mm. ;
upper tooth-row about 8 mm., the teeth

unusually robust (South-eastern Spain) M. clentatus, p. 703.
Nasals strongly cuneate, their combined breadth
posteriorly much less than half that ante-
riorly ; diastema usually longer than nasal,
never noticeably shorter.
Largest skulls with condylobasal length
about 27 to 30 mm.
Kostrum very short and deep (distance
from front of zygoma to tip of nasal
less than depth of rostrum through
posterior extremity of nasal) ; nasal
forming very conspicuous angle (about
34"^) with dorsal surface of interorbital

region (Southern Hungary) M. angiUaris, p. 706.

Rostrum normal in form (distance from
front of zygoma to tip of nasal greater
than depth of rostrum through pos-
terior extremity of nasal) ; nasal not
forming very conspicuous angle (about
20°) with dorsal surface of interorbital
region.
Auditory bullae unusually large ; upper
molars with osteodentine spaces re-
duced (Thessaly) M. hartituji, p. 704.

Auditory bullae not unusually large;
upper molars with osteodentine
spaces of normal width.
General form of skull narrow and
agrestis-like ; median ridge of pos-
terior termination of bony palate
without evident groove along su-
ture ; zygomata not conspicuously
expanded at middle ; underparts
pale clear grey (Channel Islands). M. sarnius, p. 700.
General form of skull broad, not
agrestis-\\kti ; median ridge of pos-
terior termination of bony palate
with evident groove along suture ;
zygomata conspicuously expanded
at middle in adults ; underparts
brownish buff or buffy grey (Ork-
ney Islands).
Brain-case not flattened, the ratio
of occipital depth to width
about 57 ; condylobasal length
of largest skulls 28 to 29 '6 mm.

(South Orkney Islands) M. orcadensis, p. 694.

Brain-case flattened, the ratio of
occipital depth to width about
58 ; condylobasal length of

662 RODENTIA

largest skulls 26-6 to 27 "6 mm.

(North Orkney Islands) M. sandayensis, p. 696.

First lower molar \Yith anterior
outer re-entrant angle obso-
lete; back decidedly greyish ;
underparts scarcely tinged
with yellowish brown (Sanday

Island) M.S. sa»rfa)/e»si«, p. 697.

First lower molar with anterior
outer re-entrant angle usually
well developed ; back not de-
cidedly greyish ; underparts
strongly washed with yellow-
ish brown (Westray Island)... M. s. westriv, p. 698.
Largest skulls with condylobasal length less
than 27 mm.
Brain-case with occipital region noticeably
flattened ; skull in general tending to

assume a fossorial aspect (Alps) M. inccrtus, p. 690.

Brain-case with occipital region not notice-
ably flattened ; skull in general not
tending to assume a fossorial aspect.
Interorbital region much elevated, so
that dorsal profile of skull is strongly
convex (Asturias and Sierra de Gua-

darrama) M. asturiatuis, p. 693.

Interorbital region low, so that dorsal
profile of skull is slightly convex or

nearly flat M. arvalis, p. 681.

Skull narrow and rounded ; brain-case
so long that distance from condyle
to back of interorbital constric-
tion is greater than zygomatic
breadth (Roumania and southern

Hungary) M. a. levis, p. 687.

Skull broad and rather angular ;
brain-case so short that distance
from condyle to back of inter-
orbital constriction is at most
equal to zygomatic breadth.
General colour greyish or brownish,
the sides of body not conspicu-
ously huffy (Eastern Germany) M. a. dupHcatus, p. 686.
General colour a decidedly yellowish
brown, the sides of body con-
spicuously suffused with buff.
Size slightly smaller and colour a
little less yellowish (Central

Europe) M. a. arvalis, 'p. 683.

Size slightly larger and colour a
little more yellowish (Pyre-
nees) M. a. meridiainis, p. 686.

MICROTUS AGRESTIS Linnjeus.

(Synonymy under subspecies.)

Geographical distribution. — Northern and central Europe from
northern Scandinavia to the Alps, Pyrenees, Galicia and Portugal,
and from England, Scotland and the Hebrides eastward.

MICROTUS 663

Diagnosis. — Size small or medium (hind foot, 17 to 21 mm.;
condylobasal length of skull in largest individuals, 24 • 6 to 28*4
mm.) ; second upper molar with well developed postero-internal
triangle ; first lower molar with four re-entrant angles on outer
side ; skull moderately broad (ratio of zygomatic breadth to
condylobasal length ranging from 53 to 60), the intertemporal
region developing a distinct ridge with age ; length of brain- case
measiu-ed from interorbital constriction to condyle equal to or
greater than zygomatic breadth.

External characters. — General form robust, the neck short
and thick, the head large, wide and blunt, the ears inconspicuous ;
legs short ; tail about one-third as long as head and body. Head
large, broad posteriorly, tapering slightly to the blunt muzzle ;
ear scarcely appearing above fur, and extending, when laid
forward, barely half-way to eye, its outline evenly rounded ;
meatal lobe well developed, its height at middle about 3 mm.
Eye small, not very prominent, situated distinctly nearer to
muzzle than to base of ear, its diameter contained about ?>h times
in distance from inner canthus to muzzle. Nostril pad small,
not very well defined, divided down middle by a narrow but
evident groove continuous with that crossing upper lip ; nostril
opening almost directly outward, its anterior margin noticeably
swollen. Mouth small, the upper teeth very slightly projecting.
Fore foot with inner digit reduced to a minute tubercle, whose
dorsal surface is mostly covered by the closely appressed, highly
arched nail ; third and fourth digits sub-equal and longest, fifth
extending barely beyond base of fourth, second intermediate
between fifth and third ; palmar tubercles five, well developed,
occupying considerably more than half surface of palm, that at
base of thumb is slightly the largest (more than twice as large
as thumb itself), the others are sub-equal ; surface of palm between
tubercles slightly and inconspicuously granular. Hind foot
moderately long and slender, the toes relatively short (length of
third digit a little less than one-third length of sole) ; inner digit
extending scarcely to base of second, outer slightly beyond base
of fourth ; second, third and fourth sub-equal ; plantar tubercles
six, all well developed, occupying distinctly more than half of
surface of region in which they lie ; with the exception of the
sixth which is sub-circular in outline and only about half as large
as the others, they are sub-equal in size and all are irregularly
pyriform in outline ; surface of integument between tubercles
finely granular ; behind last tubercle the sole is densely covered
with short hair. Claws simple, moderately curved, those of hind
feet, toe for toe, slightly the longer. Tail extending slightly
beyond outstretched hind foot, its annulations pronounced though
often somewhat irregular, about 20 to the centimeter at middle ;
hairs not sufticiently abundant to conceal annulations, but form-
ing an evident though thin pencil, their length usually equal to
width of two or three rings. Fur without special peculiarities of

664

ROPENTIA

distribution or texture, its quality somewhat harsh and loose.
Mamma; : p 2-2, « 2-2 = 8.

Sl-ull. — The skull is moderately broad and rather deep, attain-
ing a marked degree of angularity in old age. Dorsal profile
moderately convex throughout, the interorbital region usually
flattish, the nasals sloping forward at an angle of about 18°.
Occiput sufficiently oblique for the condyles to be visible when
skull is viewed from above. Ventral profile with no special
peculiarities ; contrast between occipital depth and that of rostrum
very pronounced. Brain-case oblong, well squared anteriorly by
the prominent postorbital processes, its lateral borders nearly
parallel except for the break caused by the zygomatic roots ;
outer edges of parietals marked in old age by well developed
longitudinal ridges which are continuous posteriorly with lambdal
crest and which come together abruptly in front to join the

interorbital ridge. No trace of
sagittal crest ; lambdal crest repre-
sented by the outer extremities
only. Interparietal large, its area
nearly equal to that of parietal,
its form somewhat variable, but
posterior border usually straight
or nearly so, anterior border bicon-
vex with median projection, the
outer extremities rather abruptly
truncate ; greatest antero-posterior
diameter, exclusive of median pro-
jection sensibly less than half
transverse diameter. Depth of
occiput when viewed from behind
more than half width, the brain-
case rising slightly above occipital
level ; paroccipital processes well
developed, their tips extending
noticeably beyond level of lower lip of foramen magnum, their
bases continued upward as conspicuous ridges along suture
between occipital and squamosal. Floor of brain-case without
special features, the basioccipital with ill-defined median ridge ;
width of basioccipital along anterior suture contained about
three times in median length. Auditory bulla? large, evenly
inflated, without much spongy tissue, the surface smooth and
shining ; beak moderately developed, closely applied to outer side
of base of pterygoid ; region near meatus slightly flattened.
Interorbital region narrow, sub-cylindrical, with distinct temporal
ridges which eventually unite to form a definite median crest
extending well forward to nasals. Zygomata wide spreading
and short, a small portion near middle essentially parallel with
main axis of skull ; median expansion evident but not very wide ;
anteorbital foramen narrow and high, the plate forming outer

Fig. 120.
Microtus agrestis. Nat. size.

MICROTUS 665

wall of canal I'udimentary below, absent above. Rostrum slender,
scarcely wider than interorbital region, rather deep proxinially
but shallow anteriorly, the least depth behind incisors slightly
greater than width in same region ; nasals broad anteriorly,
abruptly narrowed at middle, the posterior termination pointed,
blunt, or angular-emarginate, seldom extending behind level of
middle of zygomatic root, and usually a little exceeded by nasal
branches of premaxillaries ; incisive foramina long and narrow,
slightly narrower posteriorly than anteriorly, extending from about
3 mm. behind incisors almost or quite to level of molar alveoli.
Palate rather narrow (less than twice width of alveolus), marked
by two longitudinal grooves continued back from incisive foramina
to lateral bridges of posterior border ; median posterior ridge and
lateral pits well defined, the anterior border of pits on level w^ith
first inner re-entrant angle of m^. Mesopterygoid space narrow,
about 2^ times as long as wide, the pterygoids and hamulars
straight. Ectopterygoid plate well developed, the ectopterygoid
pit large and deep, not encroached on by bulla. Mandible robust,
the portion in front of molars short ; masseter ridge well developed ;
coronoid process large, its base broad, its extremity rising to level
of condyle and distinctly curved backward ; articular process
abruptly bent inward at level of base of incisor, the root of
which does not as a rule, however, produce any marked swelling
on outer side of process ; angular process well developed though
rather slender, curved strongly outward.

Teeth. — Upper incisors robust, strongly curved, the shaft
forming almost exactly half a circle, the root producing a slight
protuberance in lower portion of anteorbital foramen, the exposed
part of tooth directed downward with a slight backward curve,
the anterior face barely visible when skull is
viewed from above ; section of shaft obscurely
triangular with broadly rounded angles, the
anterior face longest, the inner face nearly as
long as anterior face, the postero-external
face noticeably shorter than either of the
others ; enamel covering entire anterior face
and extreme anterior portion of inner face ;
cutting area rather deeply but simply hol-
lowed, the anterior (enamel) edge entire and
nearly straight when viewed from in front,
obliquely flattened and with rather abruptly y^^ -^g^,

backward-curved extremities when viewed Microtus agrestic.
from below, the posterior edge usually with Enamel pattern, x 5.
some irregular notches cut by the points of
the lower teeth. Lower incisors much less strongly curved
than upper incisors, their roots extending into base of articular
process of mandible but not producing any noticeable swelling
on outer surface ; section of shaft much like that of upper
incisor, but antero-posterior diameter relatively greater, and

666

RODENTIA

anterior border more strongly and evenly rounded, so that
triangular outline is less evident ; distribution of enamel essen-
tially as in upper tooth. Molars moderately large relatively
to size of skull, their enamel pattern well dehned, with sharp
salient and re-entrant angles ; upper teeth without noticeable
contrasts in length of crowns, but m^ decidedly longer than
either of the succeeding mandibular teeth. First upper molar
normally with an anterior transverse loop and four alter-
nating closed triangles ; rarely a postero-internal loop similar
to that in m'^ may be present ;* two re-entrant angles are
normally present on each side (a third on inner side when
postero-internal loop is present), they are sharply pointed and
well defined, their points extending beyond median line of
crown, those of outer side slightly narrower and deeper than
those of inner side ; anterior loop crescentic in outline, its inner
limb longer and narrower than outer limb (though its apex does
not extend to level of those of inner triangles), its length
decidedly less than greatest width of tooth, its main axis oblique
to that of tooth-row ; inner and outer triangles essentially alike
in form, those of inner side slightly the larger ; postero-internal
loop when fully developed similar to that of m'^, but more often
rudimentary. Second upper molar with an anterior transverse
loop, two outer and one inner closed triangles, a postero-internal
rounded loop about half as large as the largest triangle, and two
outer and two inner re-entrant angles ; anterior loop much like
that of m^, but wider externally than internally, its long axis
nearly transverse to tooth-row, its length nearly equal to greatest
Avidth of tooth ; closed triangles similar to last three of preceding
tooth ; postero-internal loop oval or elliptical, its long axis
directed slightly forward. Third upper molar with anterior loop
and three closed triangles essentially like those of m^, except that
closed triangles are smaller and transverse loop is nearly
symmetrical and about as wide as crown at middle ; behind
closed triangles lies a long terminal loop subtended externally
by a slight notch and internally by a deep re-entrant angle ;
this loop in its simplest form is indented at middle by a deep
re-entrant angle which imparts to it a strongly curved crescentic
outline, the anterior limb longer and more pointed than the
posterior limb ; outer convex surface usually marked by a slight
salient angle just behind point of attachment to second outer
closed triangle ; in this, the most usual form, the tooth has three
well developed re-entrant angles and four well developed salient
angles on inner side, two well developed re-entrant angles and a
notch and three well developed salient angles and a slight pro-
jection on outer side ; in the most complicated form of terminal

* The presence of this loop is a normal character in the Hebridean
form, M. agresiis exsul, some trace of it being visible in ten of the fourteen
specimens examined. Among the other races it appears to be uniformly
uncommon, occurring in about five per cent, of the skulls seen.

MICROTUS 66 <

loop the inner re-entrant angle extends across to outer side,
isolating a second inner closed triangle and leaving the posterior
limb of crescent as a broad, rounded, backward-projecting, simple
loop, the inner side of which sometimes bears an incipient
re-entrant angle. First lower molar with posterior transverse
loop, three inner and two outer closed triangles, and a large
anterior loop indented by a deep re-entrant angle on inner side
and a shallow, more posterior reentrant angle on outer side, the
latter sometimes deep enough to meet point of inner re-entrant
angle subtending the terminal loop, thus isolating a third outer
triangle and reducing the loop to a crescent much like that
terminating m^ in its simpler form ; rarely the extreme antero-
internal border of loop shows traces of a supplemental re-entrant
angle ; inner side of tooth with five well developed salient
angles and a rounded anterior projection (rarely showing a
tendency to divide into two), and five well developed re-entrant
angles (rarely a rudimentary sixth) ; outer side of tooth with
three deep, definite re-entrant angles and a shallow fourth, and
four well developed salient angles and a broadly rounded anterior
convexity which sometimes develops a slight basal projection ;
outer re-entrant angles wider and shallower than those of inner
side, their points curved forward, the contrast between the two
sides much more noticeable than in the maxillary teeth ; outer
triangles about equal in size to those on inner side of m^, inner
triangles decidedly larger and more transversely elongated ;
posterior loop somewhat oblique, narrowly crescentic, slightly
curved, its inner limb much the longer, its transverse diameter
about equal to that of greatest width of crown. Second lower
molar with transverse posterior loop, two inner and two outer
closed triangles, two re-entrant and three salient angles on each
side, the elements essentially like the corresponding portion of
m^ but slightly smaller ; occasionally the anterior (external)
triangle develops on inner side a slight basal projection which
may become sufficiently pronounced to add a rudimentary salient
and re-entrant angle to those normally present. Third lower
molar with three transverse loops, each side with three salient
and two re-entrant angles, those of inner side much the deeper,
the antero-external re-entrant angle often obsolete ; posterior
-Oop larger than either of the others, more broadly crescentic in
outline than terminal loops of 7h, and m^, though not essentially
different in form, the outer limb practically absent ; first and
second loops somewhat irregular and variable in form, the first
usuUy truncate externally, the second usually with an angular
projection at outer side.

Remarlis. — This species is recognizable among the European
voles by the presence of the well developed postero-internal loop
in the second upper molar. Externally it cannot always be
distinguished with certainty, particularly in dry specimens.
Throughout the greater part of its range, where the only other

668 RODENTIA

member of the sub-genus is Microtus arvalis, fully adult
individuals may be i-ecognized by their larger size, more hairy
ears, larger tubercles on sole, and by the browner, less buffy
coloration, the dark and light elements of which are more
coarsely mixed. As might be anticipated from the animals wide
range, numerous local forms have been developed. Seven of
these are now known.

Microtus agrkstis agrestis Linnseus.

1761. Mils agrestis Linnaeus, Fauna Suecica, 2nd ed., p. 11 (Upsala,

Sweden).
1766. [Mas] gregarius Linnaeus, Syst. Nat., i, 12th ed., p. 84 (Germany

and Sweden).
1792. Mus arvalis nigricans Kerr, Anim. Kingd., p. 239 (Renaming of Mas

agrestis).
1820. Lemmiis arvalis Nilsson, Skand. Fauna, i, p. 189 (not Mns arvalis

PaUas).
1841. Arvicola agrestis de S61ys-Longchamps, Bull, de I'Acad. Royale des

Sci. des Arts et Belles-Lettres de Braxelles, viii, pt. 2, p. 236.
1844. L[cmviN.s] insularis Nilsson, Ofversigt af Kongl. Vetensk.-Akad.

Forhandl., Stockholm, i, p. 34, March 20, 1844. (Ostgotha

Skargard, Sweden.)
1857. Arvicola agrestis a. Blasius, Saugethiere Deutschlands, p. 369 (part).
1884. Microtus agrestis Lataste, Ann. IMus. Civ. Stor. Nat. Geneva, xx,

p. 255, March, 1884.
1896. Microtus agrestis Barrett-Hamilton, Proc. Zool. Soc, London, p. 602.
1910. Micjvtus agrestis Trouessart, Faune Mamm. d'Europe, p. 175 (part).

Type locality. — -Upsala, Sweden.

Geographical distribution. — Scandinavian Peninsula ; eastward
into Finland. Exact limits of range not known.

Diagnosis. — Size large (hind foot, 18 to 19 "4 mm.; condylo-
basal length of skulls in largest individuals, 27 to 28*5 mm.):
skull with brain-case tending to be relatively short, the distance
from condyle to back of interorbital constriction usually about
equal to zygomatic breadth ; first upper molar rarely with small
postero-internal loop ; general colour above a light bister, below
greyish with faint buffy cast.

Colour. — Hairs of upper parts slate-black basally, those of
underfur tipped (about 2 mm.) with a dull buff' intermediate
between the ochraceous-buff and cream-buff of Ridgway (the
extreme tips often dark), the longer hairs iridescent black. The
general effect is a clear brown, rather lighter than bister along
back and becoming somewhat buffy on sides, everywhere rather
conspicuously "lined" by the longer black hairs. Underparts
well-defined light silvery grey faintly washed with buffy and
darkened irregularly by the slaty bases of the hairs. Feet and
under surface of tail concolor with belly ; dorsal surface of tail
dark brown. In winter the colour above is usually not so dark
as in summer, and the "lining" on back is less conspicuous;

MICROTUS 669

occasionally the shoulders are suffused with grey ; tail more
noticeably bicolor than in summer, and underparts of body more
silvery.

Skull and teeth. — Skull large, that of old individuals becoming
strongly angular and developing a knife-like ridge in interorbital
region. Brain-case showing no tendency to lengthen out
posteriorly, the distance from back of interorbital constriction to
posterior surface of condyle barely equal to zygomatic breadth.
Teeth with no special peculiarities ; in^ rarely"' showing any trace
of postero-internal loop.

Measurements. — Average and extremes of five adults from
Upsala, Sweden: head and body, 121-8 (118-127); tail, 35-8
(33-38); hind foot, 18 "4 (18-19). Average and extremes of
five adults from Medstugan, Jemtland, Sweden : head and body,
126-8(123-132); tail, 35- 6 (31-39) ; hind foot, 18-6 (18-19-4).
Adult male from Porsanger, Norway : hind foot, 18. For cranial
measurements see Table, p. 676.

Specimens examined. — Seventy-three, from the following localities : —
Norway : Porsanger, 1 (U.S.N.M.) ; Brekkebygden, Trondhjem, 2 ;
Molmen, Kristiansamt, 4; Holaaker, Kristiansamt, 2; Hjerkin, Kris-
tiansamt, 4 ; Sundal Fjeld, 1 ; Alverstrom, Bergen, 2 (U.S.N.M.) ; Bergen, 2
(U.S.N.M.); Graven, Bergen, 3 (U.S.N.M.); Gausdal, 3 (B.M. and
U.S.N.M.) ; Asker, Kristiania, 1 (U.S.N.M.) ; Smaalenene, 1 (U.S.N.M.).

Sweden: Medstugan, Jemtland, 11; Upsala, 34 (B.M. and U.S.N.M.);
near Stockholm, 2.

4 6. Molmen, Kristiansamt, R. J. Guninghame 98. 5. 2. 6-9.

Norvyay. (c & p).

2 6. Holaaker, Kristiansamt. R. J. Guninghame 98. 2. 28. 6-7.

(c & p).
2 6. Hjerkin, Kristiansamt. G. Barrett-Hamilton 11. 1. 2. 46-47.
{N. F. Ticehurst.) (c & p).

1. Sundal Fjeld. E. Lort Phillips 94. 10. 22. 1.

(c & p).
1. Gausdal. Christiania Museum 93. 3. 1. 12.

(E).

1. Gausdal. Miller Collection. 7. 7. 7. 4537.

3 6,5 9. Mandal. R. J. Guninghame 8. 8. 9. 30-37.

(c & p).

1. Jemtland, Sweden. Baron E. de S^lys- 45. 7. 5. 12.

Longchamps (p).

4 6. Upsala. {G. Kolthoff.) Lord Lilford (p). 0. 5. 15. 2-5.

6, ?. Stockholm. " Prof. Sundevall (c). 45. 10. 25. 5-6.

MiCROTUs AGRESTis EXSUL Miller.

1908. Microtus agrestis exsul Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 201, February, 1908. Type in British Museum.

1909. Microtus agrestis insul Lydekker, Zool. Record, xlv (1908), Mamm.,

p. 74 (Accidental renaming of exsul).

1910. Microtus agrestis exsul Trouessart, Faune Mamm. d'Europe, p. 176.

Ty^je locality. — -North Uist, Hebrides, Scotland.
Geographical distribution. — North and South Uist, Hebrides.

* In a little more than five per cent, of the specimens examined.

670

RODENTIA

Diagnosis. — Size and cranial characters as in the large
Microtus agrestis agrestis of Norway and Sweden ; first upper molar
with small third inner loop usually present (in ten among fourteen
skulls examined), almost as well developed as postero-internal
loop in second tooth ; colour of underparts more brownish than
in the other races.

Colour. — Upper parts and sides darker and clearer (less
russet) brown than in M. agrestis hirtus and M. a. neglectus,
essentially as in true agrestis ; underparts rather heavily washed
with brownish buff, this suffusion in many specimens becoming a
definite, rather light ochraceous-buff.

Skull and teeth. — The skull is not distinguishable with
certainty from that of true Microtus agrestis,
but is at once recognizable among those of
the British races by its large size. Teeth
with no peculiarities other than the unusual
frequency with which a definite postero-
internal loop is present in m^. This occurs
in no less than ten of fourteen skulls ex-
amined, while in specimens of the other
races it is present only eight times among
136 skulls.

Measurements. — External measurements of

type (adult female) : head and body, 123 ;

tail, 44 ; hind foot, 19 ; ear from meatus, 12.

A second female from the same locality :

head and body, 111 ; tail, 39 ; hind foot,

18" 5; ear from meatus, 12. Adult female from Loch Boisdale,

South Uist : head and body, 111; tail, 44 ; hind foot, 18-5; ear

from meatus, 12. For cranial measurements see Table, p. 677.

Specimens examined. — Fourteen, from the following localities in the
Hebrides : North Uist, 11 (B.M. and Edinburgh) ; Loch Boisdale, South
Uist, 3 (Edinburgh).

BemarJcs. — This form is of unusual interest on account of the
presence as a normal character of a peculiarity of the enamel
pattern occurring elsewhere in the species as a rather rare
anomaly.* Its general unlikeness to the other British forms and
similarity in certain respects to true agrestis of Scandinavia is
also worthy of special note.f

S, 3 9. N. Uist, Hebrides. J. F. Davison (c & p). 6. 3. 1. 1-4.

(6. 3. 1. 3. Ty2ye of subspecies.)
1 al. N. Uist. J. A. Harvie Brown (p). 79. 9. 18. 2.

* A postero-internal loop in m* is sometimes found in the American
Microtus pennsylvanicus, a species with the same enamel pattern as
M. agrestis. Such a tooth is figured in North American Fauna, No. 12,
fig. 1. July 23, 1896.

t For a discussion of the general problems of distribution involved in
this animal's presence in the Hebrides, see Stejneger, Smithsonian Miscell.
Coll., XLViii, pp. 458-512, May 4, 1907, and Naturen (Bergen), xxxii,
pp. 193-202, 269-277, July-September, 1908.

Microtus agrestis exsitl.
Enamel pattern. X 5.

MICROTUS

671

MlCROTUS AGRESTIS LEVERNEDII Crespon.

1844. A[_rvicola] levernedii Crespon, Faune M6ridionale, i, p. 73 (Marshes

between St. Gilles and Aigues-Mortes, Gard, France). Type in

Nimes Museum.
1857. Arvicola agrestis a. Blasius, Saugethiere Deutschlands, p. 369 (part).
1869. [Arvicola agrestis] var. nigra. Fatio, Faune Vert. Suisse, i, p. 241

(Engstlen, Berne, Switzerland. Alt. 1750 m.). Type in Geneva

Museum.
1900. [Arvicola agrestis] rufa Fatio, Revue Suisse de Zool., viii, p. 472

(Geneva, Switzerland). Type in Geneva Museum.
1905. Arv[icola^ agrestis angustifrons Fatio, Arch. Sci. Phys. et Nat.,

Geneve, 4th ser., xix, p. 191, February 15, 1905 (Meiringen, Berne,

Switzerland. Alt. 650 m.). Type in Geneva Museum.
1905. Arv[icola] agrestis latifrons Fatio, Arch. Sci. Phys. et Nat., Geneve,

4th ser., xix, p. 194, February 15, 1905 (Geneva, Switzerland).

Type in Geneva Museum.
1910. Microtus agrestis Trouessart, Faune Mamm. d'Europe, p. 175 (part).

Type locality. — Marshes between St. Gilles and Aigues-Mortes,
Gard, France.

GeograpMcal distribution. — Alps, Jura, and neighbouring
portions of Switzerland and France ; also in the extensive marshes
on Mediterranean coast of France at mouth of the Rhone.

Diagnosis. — Similar to Microtus agrestis agrestis, but skull
with brain-case tending to be longer and narrower, the distance
from condyle to back of interorbital constriction usually greater
than zygomatic breadth.

Measurements. — Type (adult female) : hind foot (dry), 20.
Adult male from the type locality : head and body, 131 ; tail,
46; hind foot (fresh), 20*6, (dry), 20; ear, 11-8. Immature
male from the type locality : head and body, 107 ; tail, 39 ; hind
foot (dry), 20 ; ear, 11*8. Three adult males from St. Cergues,
Vaud, Switzerland : head and body, 131, 132 and 133 ; tail, 37,
33 and 38 ; hind foot (dry), 18 '6, 19 and 19. Adult male from
Meiringen, Berne, Switzerland: head and body, 127; tail, 40;
hind foot, 18 '6. For cranial measurements see Table, p. 676.

Specimens examined. — Sixty-six, from the following localities : —
France: Near St. Gilles, Gard, 6 (B.M. and Nimes); Montauban,
Haute-Savoie, 1 ; Cranves-Sales, Haute-Savoie, 8 ; Lucinges, Haute-
Savoie, 2; Chamonix, Haute-Savoie, 1 (U.S.N.M.) ; Etupes, Doubs, 1.

Switzerland : Geneva, 9 (Mottaz and Geneva ; types of latifrons Patio
and rufa Patio among latter) ; St. Cergues, Vaud, 5 (U.S.N.M.) ; Chesieres,
Vaud, 3 (Mottaz) ; Valine de Joux, Vaud, 3 (Mottaz) ; Les Plans, Vaud, 1
(U.S.N.M.); Grindelwald, Berne, 1 (U.S.N.M.); Meiringen, Berne, 20
(B.M., U.S.N.M. and Geneva; type of angustifrons Fatio among latter);
Briinig, Berne, 1 ; Engstlen, Berne, 1 (Geneva ; type of niger Fatio) ;
Degersheim, St. Gallen, 1 (U.S.N.M.); Chur, Grisons, 1 (U.S.N.M.);
Oberholsen Valley, 1.

Bemarhs. — Though not well differentiated from true Microtus
agrestis the Alpine form is usually distinguishable by its relatively
more elongate brain-case, a character often more evident in sub-
adult individuals than in those of more advanced age. The

672

RODBNTIA

material representing the marsh-inhabiting animal to which the
name levernedii was originally applied is too meagre to permit
any final decision as to the identity of this form with that
occurring in Switzerland. The brain-case in the type as well as
in the five other sjaecimens examined is of the elongated form
characteristic of the Alpine race as compared with true agrestis.
The hind foot, however, appears to be rather constantly larger
than in the Swiss animal. Eventually it may prove that the
name levernedii must be restricted to this semi-aquatic vole, in
which event the name niger of Fatio would be available for the
Swiss form.

4 6. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4.233-236.

6. Montauban, Haute-Savoie, 0. Thomas (p). 6. 4. 2. 7.

900 m. {A. Robert.)

6,9. C r a n V e s - Sales, Haute- A. Robert (c & p). 5. 4. 9. 8-9.
Savoie, 900 m.

5 &, ?. Cranves-Sales, Haute-Savoie, 0. Thomas (p). 5. 11. 8. 24-27,

1200 m. {A. Robert.) 29, 82.

?. Lucinges, Haute-Savoie, 0. Thomas (p). 6.4.2.7-8.

900 m. (A. Robert.)

9. Etupes, Doubs. (C. Mottaz.) 0. Thomas (p). 8. 8. 10. 115.

2. Meiringen, Berne, Switzer- Tomes Collection. 7. 1. 1. 139-140.

land.

1. Oberholsen Valley. Tomes Collection. 7. 1. 1. 138.

MiCROTUS AGRESTis BAiLLONi de Selys-Longchamps.

1841. Arvlicola] baillonide S61ys-Longchamps, Atti della seconda Riunione

degli Scienziati Italiani, Torino, 1840, p. 225 (Northern France ;

Abbeville, Somme, from context).
1845. ? Alrvicola] intermedia Bonaparte, Atti della sesta Riunione degli

Scienziati Italiani, Torino, 1844, p. 350 (cited as doubtful synonym

of bailloni, with de Selys-Longchamps as authority). Nomen

nudum.
1857. Arvicola agrestis a. Blasius, Saugethiere Deutschlands, p. 369 (part).
1896. Microtus agrestis neglectus Barrett-Hamilton, Proc. Zool. Soc,

London, p. 602 (part).
1910. Microtus agrestis neglectus Trouessart, Faune ]\Iamm. d'Europe,

p. 176 (part).

Type locality. — Abbeville, Somme, France.

Geographical distribution. — Western Continental Europe from
the shores of the Baltic southward into central Germany and
south-western France. Southern and eastern limits of range not
known.

Diagnosis. — Simiiar to Microtus agrestis agrestis but smaller
(hind foot, 17 to 18 "6 mm.; cotidylobasal length of skull in
largest individuals, 24 • 5 to 26*6 mm.) ; skull with brain-case
tending to assume ^ulie narrower form characteristic of M. agrestis
levernedii.

Measurements^' — Adult male from Hillerod, Zealand, Denmark :
head and body, 113; tail, 3i) ; hind foot, 18- 2; ear, 13-5.

MICROTUS 673

Average aud extremes of seven adults from Brunswick, Germany :
head and body, 116-2 (109-123) ; tail, 38-8 (36-44) ; hind foot,
17*2 (16 "6-18). Young-adult male from Guines, Pas-de-Calais,
France: head and body, 103; tail, 34; hind foot, 17. Young-
adult male from Barbizon, Seine-et-Marne, France : head and
body, 98; tail, 32; hind foot, 17 "6. Adult male from Porte,
Pyrenees-Orientales, France : head and body, 120 ; tail, 41 ; hind
foot, 17. For cranial measurements see Table, p. 678.

Specimens examined. — Forty-five, from the following localities : —

Denmark: Hillerod, Zealand, 3; Skansen, Lolland, 1 (U.S.N.M.).

Germany : Brunswick, 28 (U.S.N.M.).

France : Guines, Pas-de-Calais, 1 ; Dinan, C6tes-du-Nord, 1 ; Manon-
ville, Meurthe-et-Mosellc, 1 ; Barbizon, Seine-et-Marne, 3 ; Mont Dore,
Puy-de-D6me, 1 ; Solferino, Landes, 1 ; Foret-de-Bouconne, Gors, 1 ;
Porte, Pyr6nees-0rientales, 2 ; I'Hospitalet, Ariege, 1 ; Pyrenees, no exact
locality, 1.

6, 2 9. Hillerod, Zealand, Den- 0. Thomas (c & v). 98. C. 7. 20-22.
mark.
6. Guines, Pas - de - Calais, 0. Thomas (c & p). 94. 6. 6. 18.

France.
6. Manonville, Meurthe-et- Lord Lilford (p). 1. 11. 7. 12.

]\Ioselle. (Lomont.)
6, 9. Barbizon, Seine-et-Marne. G. S. Miller (c). 8. 8. 4. 231-232.

6. Solferino, Landes. 0. Thomas (p). 6. 4. 1. 75.

(A. Robert.)
?. Foret de Bouconne, Gers. 0. Thomas (p). 6. 4. 1. 76.

(.4. Bohcrt.)
6, 9. Port6, Pyrenees-Orientales. G. S. Miller (c). 8. 8. 4. 229-230.

9. L'Hospitalet, Ariege. G. S. Miller (c). 8. 8. 4. 228.

6 juv. Mt. Dore, Puy-de-D6me, G. S. Miller (c). 8. 8. 4. 227.

4500 ft.

MiCROTUS AGRESTis iiiRTUS Bellamy.

1839. Arvicola hirta Bellamy, Nat. Hist. South Devon, p. 373 (Devonshire,
England).

1847. A{rvicola] britannicns de S61ys-Longchamps, Revue Zoologique,
1847, p. 307, October, 1847 (England).

1847. [Arvicola^ britannicus de S^lys-Longchamps, Atti della ottona
Riunione degli Scienziati Italiani, Genova, 1846, p. 495.

1857. Arvicola agrestis h. Blasius, Saugethiere Deutschlands, p. 369 (part).

1896. Microtus agrestis neglectus Barrett-Hamilton, Proc. Zool. Soc,
London, p. 602 (part).

1910. Microtus agrestis neglectus Trouessart, Faune IMamm. d'Europe,
p. 176 (part).

Tyj?e locality. — Yealmpton, Devonshire, England.

Geograjjhical distrihufion. — England and the lowlands of
southern Scotland.

Diagnosis. — Size as in Microtus agrestis hailloni or slightly
smaller (hind foot, 17 to 18 mm. ; condylobasal length of skull
in largest individuals, 24 '5 to 26 mm.); skull with brain-case

2 X

674

RODENTIA

I'IG. 132.
Microtus wjrestis fiirtus. Nat. size.

tending to assume the sliort squarish form characteristic of
true a(jrestis ; upper parts noticeably tinged with russet, and
belly heavily washed with wood-brown.

Colour. — General effect of upper
parts a tawny russet very indistinctly
"lined" with black, the individual
hairs slate-black basally, the light
portion closely approaching orange-
buff, though not so bright ; sides paler
and with a noticeable buffy tinge :
underparts light ochraceous-buff irre-
gularly darkened by the slaty bases of
the hairs ; feet and under surface of
tail concolor with belly ; upper sur-
face of tail dark brown.

Measurements. — Average and ex-
tremes of five adults from the New
Forest, Hampshire : head and body,
113-4(109-118); tail, 35-2 (33-39);
hind foot, 17-3 (16-6-18); ear,
12*2 (11-13). Average and extremes of five adults from Graf-
tonbury, Hereford : head and body, 113-2 (108-120) ; tail, 31-5
(29-33); hind foot, 16-7 (16-18). Average and extremes of
four adults from Great Grimsby, Lincolnshire : head and body,
110 (108-112); tail, 29-2 (27-32); hind foot, 16-5 (16-18).
For cranial measurements see Table, p. 678.

Specimens examined. — One hundred and thirty-four, from the following
localities : —

Scotland : Windygates, Fife, 4 ; Stockbriggs, Lanarkshire, 2 ; Black-
wood, Lanark, 7 ; Criefi, Perthshire, 1 ; Drumlaurig Woods, Dumfries, 1 ;
Kirtle Bridge, Dumfries, 4 ; Wyseby, Dumfries, 3 ; Hawick, Roxburgh-
shire, 1.

England : Marsham, Yorkshire, 1; Dunham Park, Bowdon, Cheshire, 1 ;
Longendale East, Cheshire, 1; Grimsby, Lincolnshire, 19; Grainsby Hall,
Lincolnshire, 5 ; Cheadle, Staffordshire, 1 ; Anglesey, Carnarvonshire, 1 ;
St. Bride's, Pembrokeshire, 1 ; Sandringham, Norfolk, 1 ; Methwold Fen,
Norfolk, 3 ; Lowestoft, Suffolk, 1 ; Rugby, Warwickshire, 1 ; Lilford,
Northamptonshire, 3 ; Oundle, Northamptonshire, 3 ; Shelford, Cambridge-
shire, 1 ; Cambridgeshire, no exact locality, 2 ; Graftonbury, Hereford-
shire, 17 ; Boxmoor, Hertfordshire, 1 ; Hampton, Middlesex, 2 ; Godalming,
Surrey, 1 ; Earlsfield, Surrey, 7 ; Southerndown, near Bridgend, Glamorgan-
shire,"l ; New Forest, Hampshire, 20 ; Eversley, Hampshire, 1 ; St. Helens,
Isle of Wight, 2; Alum Bay, Isle of Wight, 2; Bonchurch, Isle of
Wight, 1 ; St. Leonards, Sussex, 1.

Miss D. Bate (c & p). 11. 1. 8. 267-269.

2 6, 9. Wyseby, Dumfriesshire,
Scotland.

1. Drumlaurig, Dumfries-
shire.

?. Crieff, Perthshire.

Sir W. Jardine (c). 86. 7. 2. 11.

W. R. Ogilvie-Grant 88. 5. 30. 1.
(c & v).

6,i]\xv. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 54-55.

?. Hawick, Roxburghshire. J. E. Harting (c & p). 11. 1. 3. 289.

? imm. St. Brides, Pembroke- Hon. C. Edwards 90. 12. 5. 2.
(albino) ghire, Wales, (c & p).

MICROTUS 675

9 al. M a r s h a m, Yorkshire, W. B. Tegetmeicr (i>). 93. 8. 31. 1.

England. {J. Carter.)

2 6. Bowdon, Cheshire. G. Barrett-Hamilton 11. 1. 2. 48-49.

(T. A. Coward.) (p).

3 6, 9. Grimsby, Lincolnshire. G. Barrett-Hamilton 11. 1. 2. 50-53.

(G. H. Caton Haigh.) (p).

2 .5, 2 9. Grimsbj'. ((?. H. Caton W. E. de Winton (p). 11. 1. 3. 246-249.
Haigh.)

6, 9. Cheadle, Staffordshire. E. W. H. Blogg (p). 11. 1. 3. 287-288.

<^. Anglesey. T. A. Coward (c & p). 11. 1. 3. 270.

2 J, 19. Oundle, Northampton- Lord Lilford (c & p). 11.1.3.271-273.

shire.

2 6,29. Graftonbury, Hereford- W. E. de Winton 11.1.3.274-277.

shire. (c & p).

6. Shelford, Cambridge- Dr. Hans Gadow 5. 1. 21. 1.

shire. (c & p).

2 St. Cambridgeshire. J. Baker (c & p). 39. 9. 29. 4-5.

3c5,4 9. Earlsfield, Surrey. C. H. B. Grant (c & p). 11.1.3.278-284.

2 6. Alum Bay, Isle of Wight. 0. Thomas (c & p). 11.1. 3. 285-286.

1. Bonchurch, Isle of Wight. Rev. C. A. Bury 44.9.2.5.

(c & p).

19. New Forest, Hampshire. G. S. MiUer (c). 7. 7. 7. 2906-

2924.

MiCROTUS AGRESTIS NEGLECTUS JenyilS.

1841. Arvicola neglectus Jenyns, Ann. and Mag. Nat. Hist., 1st ser , vii

p. 270, June, 1841.
1857. Arvicola agrestis b. Blasius, Saugethiere Deutschlands, p. 369 (part).
1896. Microtus agrestis neglectus Barrett-Hamilton, Proc. Zool. Soc,

London, p. 602 (part).
1910. Microtus agrestis fieglectus Trouessart, Faune Mamm. d'Europe,

p. 176 (part).

Type locality. — Moors near Megarnie Castle, Pertlishire,
Scotland.

Geographical distribution. — Highlands of Scotland.

Diagnosis. — Like Microtus agrestis hirtus but size not so small
(condylobasal length of largest skulls, 25-4 to 26-6 mm.), and
colour of upper parts darker and more brownish.

Colour. — The general hue of the upper parts is noticeably-
darker and browner than in the English form, closely resembling
that of the Continental M. agrestis hailloni. The actual colour
of the light element approximates ochraceous-buff with a faint
suggestion of tawny, and the general effect may perhaps be best
described as prouts-brown slightly tinged with raw umber.
Underparts varying from a dull silvery grey faintly tinged with
buffy to a light bufly wood-brown.

Skull and teeth. — The skull averages distinctly larger than
that of M. agrestis hirtus, though it shows no peculiarities of form.
Some of the larger skulls might readily be confused with those
of small individuals of M. agrestis agrestis or M. agrestis exsul ;
but their true identity is shown by the small auditory bullse,
which retain the small size characteristic of the British races
and M. agrestis hailloni. The teeth show no peculiarities ; m^

676

RODENTIA

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with postero-internal loop present in the usual small percentage
of individuals.

Measurements. — Two adult males from Kinloch, Rannoch,
Perthshire : head and body, 120 and 120 ; tail, — and 43 ; hind
foot, 18 and 18*6 ; ear, 12 and 12. Adult female from the same
locality : head and body, 121 ; tail, 4G ; hind foot, 18*6. Two
adult females from Beauly, Inverness : tail, 37 and 38 ; hind
foot, 18 and 18. For cranial measurements see Table, p. 679.

Specimens examined. — Thirteen, from the following localities in Scotland :
Black Island, Cromarty, 1 ; Elgin, 1 ; Dunphail, Elgin, 1 ; Beauly, Inver-
ness, 3 ; summit of Ben Nevis, Inverness, 2 ; Speam Bridge, Inverness, 1 ;
Stonehaven, Kincardineshire, 1 ; Kinloch, Bannoch, Perthshire, 3.

c?, 2 9. Beauly, Inverness-shire, Hon. Margaret Eraser 8.1.4.1-3.

Scotland. (c & p).

2 al. Ben Nevis, Inverness. W. R. Ogilvie-Grant 82. 10. G. 1-2.

(c & p).
1 al. Stonehaven, Kincardine- E. Lort Phillips 85. 10. 5. 1.

shire. (c & p).

2 6, 9. Bannoch, Perthshire. Hon. N. C. Roths- 8. 9. 14. 1-3.

{F. J. Cox.) child (p).

9. Dunphail, Elgin. W. R. Ogilvie-Grant 11. 1. 3. 405.

(c & p).
9. Elgin. W. R. Ogilvie-Grant 11. 1. 3. 404.

(c & p).

MiCROTUS AGRESTIS ROZIANUS Bocage.

1865. Arvicola rozianus Bocage, Mem. Acad. Real das Scieneias de Lishoa,

N.S., III, pt. 2, p. 7 (articles separately paged).
1910. Microtus agrestis rosianus Trouessart, Faune Mamm. d'Europe,

p. 177.

Type localitji. — Geria, near Coimbra, Portugal.

Geographical distrihtition. — Portugal and north-western Spain.
Limits of range unknown.

Diagnosis. — Size and general appearance, so far as is known,*
as in Microtus agrestis hailloni ; skull with auditory bullae notice-
ably flattened, and so reduced in size that the greatest diameter
(from paroccipital process to tip of anterior spine) is contained
3 1 times instead of three times in condylobasal length ; teeth
relatively somewhat larger than in the other small forms.

Colour. — Unknown (the only specimens seen have been injured
by long immersion in alcohol).

Skull. — In general foi'm the skull agrees with that of the
other small races of Microtus agrestis, but it is immediately
recognizable by the small, slightly inflated auditory bulla?, no
approach to which has been seen among the considerable number
of skulls examined from other parts of Europe. The peculiar-
ities of the bulla are best seen in side view.

* The colour of the only skin examined has heen altered by the action
of alcohol.

MicROTns 681

Teeth. — While in no way essentially difterent from those of
the other small races of Microtus acjrestis the teeth of M. a.
rozianus appear to be relatively a little larger (see measurements,
p. 679), though the difference may prove to be merely an
individual peculiarity in the two skulls seen. In one of these
specimens there is a minute postero-internal triangle in the first
upper molar.

Measurements. — External measurements of adult female from
Villalba, Lugo, Spain : head and body, 100 ; tail, 39 ; hind foot,
18' 8, For cranial measurements see Table, p. 679.

Specimens examined. — Three, from tlie following localities in Spain :
La Corufia, 1 ; Villalba, Lugo, 2.

2 al. Villalba, Spain. Dr. V. L. Seoane (c & p). 94. 1. 1. 14-15.
1 al. Corufia. Dr. V. L. Seoane (c & p). 95. 4. 29. 3.

MICROTUS ARVALIS Pallas.
(Synonymy under subspecies.)

Geographical distribution. — Continental Europe from the
Baltic to the Pyrenees and northern Italy, and from the Atlantic
coast eastward.

Diagnosis. — Size less than in Microtus agrestis (hind foot, 15
to 18 '6 mm.; condylobasal length of skull in fully adult
individuals, 23 "4 to 26 6 mm.); plantar and palmar tubercles
relatively smaller ; second upper molar without small postero-
internal loop ; first lower molar with four re-entrant angles on
inner side ; skull slender or moderately broad (ratio of zygomatic
breadth to condylobasal length ranging from 50 to 57), never
conspicuously flattened and never assuming a " fossorial " aspect,
the upper incisors nearly perpendicular.

External characters. — In general not essentially different from
Microtus agrestis, but ear less hairy and with meatal lobe much
less developed, barely half as high ; feet as in agrestis except that
the tubercles on both palm and sole are relatively smaller, those
on fore foot occupying scarcely more than half area of palm (that
at base of thumb barely more than twice as large as thumb itself),
those on hind foot occupying distinctly less than half region in
which they occur ; in form the tubercles all tend to be more
evenly i-ounded than in M. agrestis, owing to the less degree of
crowding.

Skull. — Apart from its smaller size the skull differs from that
of Microtus agrestis in the broader, shorter, more depressed brain-
case, the outline of which when viewed from above is distinctly
rounded both in front, behind, and at the sides, seldom if ever
showing any trace of the squaring characteristic of the larger
animal ; postorbital process low, not distinctly angular ; inter-
parietal with antero-posterior diameter, exclusive of median
pi'ojection, obviously more than half transverse diameter, the

682

RODENTIA

Fig. 133.
Microtus arvalis. Nat. size.

exact proportion of length to breadth varying in the different
races ; auditory bullae frequently though not always larger than in

M. agrestis {proportionately to size
of skull) ; nasals nari'owing more
gradually backward, not abruptly
contracted at middle ; mandible with
coronoid process usually less notice-
ably curved backward, and articular
process marked on outer side by a
more obvious protuberance over base
of incisor-root.

Teeth. — Incisors as in Microtus
agrestis except that the front face
of upper teeth is more nearly flat,
that is, less obliquely rounded off
at outer side. Molars both above
and below differing from those of
31. agrestis in a general tendency
toward wider re-entrant angles and
smaller closed triangles, which gives the pattern as a whole
a less compact appearance. In details of enamel folding the
only important difference between the two
animals is the complete absence in M. arvalis
of a postero-internal loop to 7n'^. The third
upper molar has exactly the same elements
as that of M. agrestis, and is subject to
similar variations in form. First lower
molar with re-entrant angle on inner side of
anterior loop usually less deA'eloped than in
31. agrestis, that on outer side somewhat
deeper, so that the two are approximately
equal ; this causes the loop to appear to
project forward, or to turn outward instead
of inward.

BemarJcs. — Among European voles 3Iicrotus arvalis is dis-
tinguished by its enamel pattern combined with the perfectly
normal skull and small or medium size. As in the case of
31. agrestis, dry specimens, particularly those that are faded and
distorted, cannot always be positively determined. Though
always conforming to the tetramerodont type common to the
majority of species of true Microtus occurring in both the Old
World and America, the enamel pattern shows a somewhat
unusual tendency toward individual variation in exact details of
form.* These variations appear to be in no way characteristic
of local races ; and the five subspecies here recognized are based
on other characters.

* The variations in the enamel pattern have been studied and figured
in great detail by Rorig and Borner, Arbeiten aus der Kaiserlichen Biolog-
ischen Anstalt fiir Land- uud Forstwirtschaft, V, Heft ii, i^p. 37-89,
XdIs. iv-vi, 1905.

Fig. 134.

Microtxis arvalis.
Enamel pattern. X 5.

683

MiCROTUS ARVALIS ARVALIS PallaS.

1778. Mus arvalis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 78 (German}').
1801. ilf[Hs] arv[alis'] albus Bechstein, Gemeinn. Naturgesch. Deutschlauds,

I, 2nd ed., p. 998 (Thiiriugen, Germany).
1803. Lemmus fulvus GeoSroy, Catal. Mammif. du Mus. Nat. d'Hist. Nat.,

Paris, p. 187 (France).
1822. Arvicola vulgaris Desmarest, Mammalogie, pt. ii, p. 282.

1840. Arvicola arvensis Schiuz, Europ. Fauna, i, p. 60 (Substitute for

arvalis) .

1841. [Arvicola] arvalis de Selys-Longchamps, Bull, de I'Acad. Roj'ale des

Sci. des Arts et Belles-Lettres de Bruxelles, viii, pt. 2, p. 235
(arvalis for the first time clearly distinguished from agrestis).

1845. ? [Arvicola arvalis] var. atcr de Selys-Longchamps, Atti della sesta
Riunione degli Scienziati Italiani, Torino, 1844, p. 321 (Nomen
nudum).

1847. ?? Arvicola cunicularius Ray, Rev. ZooL, p. 312, October, 1847
(Riceys, near Troyes, Aube, France).

1853. Arvicola campestris Blasius, Gelehrte Anzeigen, Miinchen, xxxvii,
p. 106, July 29, 1853 (Brunswick, Germany). For identification
with arvalis see Rorig and Borner, Arbeiten aus der Kaiserlichen
Biologischen Anstalt fiir Land- und Forstwirtschaft, v. Heft ii,
pp. 74-75, 1905.

1857. Arvicola campestris Blasius, Saugethiere Deutschlands, p. 375.

1857. Arvicola arvalis Blasius, Saugethiere Deutschlands, p. 379.

1884. Microtus arvalis Lataste, Ann. IMus. Civ. Stor. Nat., Geneva, xx,
p. 259, March, 1884.

1905. Arvicola arvalis, galliardi Fatio, Arch. Sci. Phys. et Nat., Geneve,
4th ser., xix, p. 197, February 15, 1905 (BuUe, Fribourg, Switzer-
land). Type in Geneva Museum.

1905. [Arvicola arvalis] forma variabilis Rorig and Borner, Arbeiten aus
der Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirt-
schaft, v. Heft II, p. 73 (Wahlstatt, near Liegnitz, Silesia, Germany).

1905. [Arvicola arvalis] forma contigua Rorig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirtschaft,
V, Heft II, p. 76 (Rothenburg, Silesia, Germany).

1905. [Arvicola arvalis] forma assimilis Rorig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirtschaft,
V, Heft II, p. 77 (Darmstadt, Hessen, Germany).

1905. [Arvicola arvalis] forma cleprcssa Rorig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- unci Forstwirtschaft,
V, Heft II, p. 88 (Bautzen, Saxony, Germany).

1905. [Arvicola arvalis] forma simplex Rorig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirtschaft,
V, Heft II, pi. V (Gransee, Brandenburg, Germany).

1905. [Arvicola arvalis] forma p)-incipalis Rorig and Borner, Arbeiten aus
der Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirt-
schaft, V, Heft II, pi. V (Burghessler, near Kosen, Thiiringen,
Germany).

1910. Microtus arvalis Trouessart, Faune Mamm. d'Europe, p. 173 (part).

1910. Microtus agrestis campestris Trouessart, Faune Mamm. d'Europe,
p. 176.

Type locality. — Germany. (Under the name arvalis Pallas
included the small voles of the arvalis and agrestis type occurring
in the region extending from England into western Siberia.

684

RODENTIA

Germany is one of the localities mentioned in the text (p. 78),
and to a form occurring in Germany the name arwaZ/s was applied
by de Selys-Longchamps, the first author to distinguish between
this species and agrestis.)

Diagnosis. — Size small (hind foot, 15 to 17 mm. ; condylobasal
length of fully adult skulls, 23 to 25 mm.); colour a nearly
uniform, distinctly yellowish brown, the underparts usually
suflFused with buffy ; interparietal ligulate in outline, its antero-
posterior diameter (exclusive of median spine) not conspicuously
more than half transverse diameter.

Colour. — Upper parts a nearly uniform yellowish brown or
brownish bufi", the sides more yellowish than back ; light tips to
hairs of underfur usually dull buff, rarely, except in abraded
pelage, approaching cream-bufF; black tips to longer hairs not
sufficiently contrasted to produce any noticeable effect of " lining "
or grizzling. Underparts varying from silvery grey to dull buff",
always clouded to a varying degree by the slate-grey under-colour.
Feet dull buffy white. Tail bicolor though not conspicuously
so, brownish or blackish above, buffy or whitish below.

Slull and teeth. — The skull of typical Microtus arvalis may
best be described as lacking any special peculiarities. It is
moderately long and narrow, the brain-case is not particularly
deepened, and the dorsal profile is without marked convexity ;
interparietal tending to be somewhat ligulate in outline, its
antero-posterior diameter in most instances not conspicuously
more than half transverse diameter ; nasals sloping forward at an
angle of about 23° ; interorbital ridges uniting to form an evident
crest in fully adult individuals ; auditory bulla3 rather large,
smoothly inflated. Teeth small, the enamel pattern normal,
though showing all the phases of individual variation known in
the European forms.

Measurements. — Average and extremes of ten adults from
Brunswick, Germany: head and body, 104 (100-111); tail, 40
(35-45) ; hind foot, 15*4 (15-16). Two adult males from near
base of the Dole, Vaud, Switzerland: head and body, 110 and
116 ; tail, — and 36 ; hind foot, 15 '6 and 16. Adult male and
female from Fiorentina, Bologna, Italy : head and body, 111 and
104; tail, 40 and 32 ; hind foot, 15-8 and 15-6 ; ear, 12 and 12.
For cranial measurements see Table, p. 688.

Specimens examined. — One hundred and seventy-nine, from the following
localities : —

Belgium: Li6ge, 1; Waremme, Li^ge, 6 (U.S.N. M.) ; Maredsous,
Namur, 3.

France : Etupes, Doubs, 21 (B.M., U.S.N.M. and Mottaz) ; Lucinges,
Haute-Savoie, 2 ; Montauban, Haute-Savoie, 2 ; Cranves-Sales, Haute-
Savoie, 5 ; Scientriers, Haute-Savoie, 2 (Mottaz) ; Poret de Bouconne,
Gars, 1.

Germaity: Brunswick, 16 (U.S.N.M.); Kalbe, Saxony, 1; Schwarzburg,
Thiiringen, 2 ; Ummerstadt, Thiiringen, 3 ; Rudolstadt, Thiiringen, 1 ;
Kalbe, Saxony, 1 ; Magdeburg, Saxony, 11 ; Moritzburg, Saxony, 5
(U.S.N.M.); Dresden, Saxony, 3 (U.S.N.M.) ; Tharandt, Saxony, 1 ; Niesky,

685

Silesia, 5; Wolfshau, Riesengebirge, Silesia, 2 (U.S.N.M.) ; Marxheim,
near Monheim, Bavaria, 2 ; Strassburg, 3 ; Mark Brandenburg, 2.

Austria-Hungary : Prag, Bohemia, 3 ; Csallokoz-Somorja, Pressburg,
Hungary, 7.

Switzerland: Geneva, 16 (B.M., U.S.N.M. and Mottaz) ; near base of
the Dole, Vaud, 17 (U.S.N.M. and Mottaz) ; Vallee de Joux, Vaud, 1 (Mottaz) ;
Chesieres, Vaud, 6 (Mottaz) ; Lausanne, Vaud, 4 (U.S.N.M.) ; Neuchatel, 1
(U.S.N.M.); BuUe, Fribourg, 1 (Geneva; type of galliardi Fatio) ; Mei-
ringen, Berne, 1 (U.S.N.M.) ; Vitznau, Lucerne, 1 ; Ziiberwangen, St.
Gallen, 11 (U.S.N.M. and Mottaz) ; Mels, St. Gallen, 3 (U.S.N.M.) ; Val
Tours, Grisons, 1 (U.S.N.M.).

Italy : Milan, 1 ; Fiorentina, Bologna, 8 ; Ferrara, 1.

1. Liege, Belgium.

2 6, 9. Maredsous, Namur.
5 6, 9. Etupes, Doubs, France.
(C. Mottaz.)
2 6. Cranves - Sales, Haute-
Savoie.
6, 2 9. Cranves - Sales, Haute-
Savoie. {A. Robert.)
9. Foret de Bouconne, Gers.
(.4. Robert.)
2 6. Schwarzburg, Thiiringen,
Germany. {SchiicJiardt.)

2 9. Ummerstadt, Thiiringen.

(Schuchardt.)
6. Rudolstadt, Thiiringen.

(Schuchardt.)
6. Kalbe, Saxony.
9,2juv. Magdeburg, Saxony.

al.
2(5,3 9. Magdeburg. (Dr. Wolter-
storff.)

1 al. Tharandt, Saxony.

(Nitsche.)
6, 2 9. Niesky, Silesia.

2. Niesky, Silesia. (W. Bacr.)

6. Marxheim, Bavaria. (Dr.

Woltcrstorff.y

3 6. Strassburg, Alsace.

(C. Mottaz.)

2 a]. Colpiu, Mark Brandenburg.

7. Csallokoz-Somorja, Press-

burg, Austria-Hungary.
6,29 al. Prag, Bohemia.

6. Geneva, Switzerland.

S. Vitznau, Lucerne.
6, 2 9. Fiorentina, Bologna, Italy.

1. Milan.

Baron E. de Selys-
Longchamps (p).
Rev. G. Fournier(p).
O. Thomas (i>).

A. Robert (c & p).

0. Thomas (p).

0. Thomas (p).

Lord Lilford (p).

Lord Lilford (p).

Lord Lilford (p).

Lord Lilford (p).
Dr. W. Wolterstorff

(c & p).
Lord Lilford (p).

Berlin Museum (e).

Dr. E.Hamilton (p).
Lord Lilford (p).
Lord Lilford (p).

0. Thomas (p).

Dr. H. Gadow(c&p).
Budapest Museum

(E).

V. Fric (p).
E. R. Alston (p).
O. Thomas (c & p).
Genoa Museum (e).

Baron E. de Selys-
Longchamps (p).

37. 1. 3. 173.

3. 3. 30. 3-5.
8.8.10.106-111.

5. 4. 1). 10-11.

5. 11. 18. 28,
30-31.

6. 4. 1. 77.

95. 4. 18. 15-16.

1. 11. 7- 7-8.
1. 7. 7. 10.

1. 7. 7. 9.

92. 12. 1. 21-23.

1. 11. 7. 1-5.

93. 1. 1. 20.

97. 12. 4. 28-30.
99. 1. 9. 20-21.
1. 11. 7. 6.

8.8.10.112-114.

82. 7. 31. 3-4.

94. 3. 1. 66-72.

90. 1. 30. 2-4.
79. 9. 25. 52.
5. 8. 3. 21.

8. 7. 18. 8.
8. 8. 2. 3-4.
45. 7. 5. 11.

686

MlCEOTUS AHVALIS MEKIDIANUH Miller.

1908. Microtus arvalis vicridianus Miller, Ann. and Mag. Nat. Hist.

8tb ser., i, p. 197, February, 1908. Type in British Museum.
1910. Microtus arvalis mcridia7ins Trouessart, Fauna Mamm. d'Europe,

p. 174.

Tyj)e locality. — Near Biarritz, Basses-Pyrenees, France.

Geographical distribution. — South-western France, in and near
the Pyrenees.

Diagnosis. — Like Microtus arvalis arvalis but larger (hind
foot, 15 '8 to 16*6; condylobasal length of fully adult skulls,
about 25' 5 mm.), and shghtly more yellowish.

Colour. — The colour so nearly resembles that of the typical
race as to require no detailed description. In general there
appears to be a tendency toward more conspicuous suffusion of
sides and underparts with buff.

Shull and teeth. — Apart from their greater size the skull and
teeth resemble those of M. arvalis arvalis. In some specimens,
however, the brain-case is unusually short, its outline almost
circular.

Measurements. — External measurements of type (adult female) :
head and body, 1 1 5 ; tail, 32 ; hind foot, 1 6 • 6 ; ear, 12. Average
and extremes of four adult individuals from the type locality :
head and body, 106-2 (102-115) ; tail, 31-7 (30-34); hind foot,
16*1 (15* 8-1 6 • 6) ; ear, 11*2 (10-1 2). For cranial measurements
see Table, p. 689.

Specimens examined. — Eleven, from the following localities in south-
western Prance: Port6, Pyrenees-Orientales, 1; Pic du Midi, Hautes-
Pyren^es, 1 skull (Lataste) ; Biarritz, Basses-Pyr^ndes, 8 ; Pyrenees, no
exact locality, 1.

?. Porte, Pyr^nees-Orientales, 0. Thomas (p). 8.9.1.70.
France. [A. Robert.)

2 c5, 4 ?. Biarritz, Basses-Pyrenees. J. F. Davison (c & p). 6. 6. 4. 23-28.

(6. 6. 4. 26. Type of subspecies.)

1. Pyrenees. Baron E. de S^lys- 45. 7. 5. 6.

Longchamps (p).

1. Tomes Collection. 7. 1. 1. 137.

Microtus arvalis duplicatus Rorig and Borner.

1905. \_Arvicola arvalis] forma duplicata Eorig and Borner, Arbeiten aus
der Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirt-
schaft, v, Heft ii, pi. v.

Type locality. — Rossitten, Ostpreussen, Germany.

Geographical distribution. — Shores of the Baltic in north-
eastern Germany. Limits of range not known.

Diagnosis. — Like Microtus arvalis arvalis but attaining a
larger size (hind foot, 17 to 18 '6 mm. ; condylobasal length of
skull in fully adult individuals about 25 to 25 * 5 mm.) ; colour

MICROTUS 687

paler, less buft'y and less unifonii ; skull larger and heavier, the
brain-case deeper than in the typical form.

Colour. — Upper parts paler than in unfaded specimens of
true arvalis, the tips of hairs of underf ur a dull cream-buff against
which the black tips of long hairs are rather noticeably con-
trasted, producing an evident effect of grizzling seldom approached
in the typical form. Underparts a light grey (about Ridgway's
No. 10), dulled by the appearance at surface of slaty under-colour,
and occasionally (much less often than in true arvalis) washed
with light buff". Feet hair-brown occasionally with a buffy tinge.
Tail obscurely bicolor, dark brownish above, dull white below.

Shill and teeth. — The skull differs from that of 31. arvalis
arvalis in its general greater size and more robust form, the
latter peculiarity often more noticeable in the rostrum than
elsewhere ; brain-case essentially like that of true arvalis when
viewed from above, the lateral ridges well developed in fully
adult individuals, its depth decidedly greater than in the typical
form ; auditory bull?e large and well inflated. Teeth showing no
special peculiarities.

Measurements. — Average and extremes of ten adults from the
neighbourhood of Tenkitten, Ostpreussen, Germany : head and
body, 111-3(105-120); tail, 35-5(31-41); hind foot, 17-5 (17-
18-6). For cranial measurements see Table, p. 689.

Specimens examined. — Sixty, from the following localities on or near
the Baltic coast of East Prussia : near Konigsherg, 1 (U.S.N.M.) ; Ten-
kitten, 40 (U.S.N.M.); Frische Haff, 2 (U.S.N.M.); Sauglienen, 4
(U.S.N.M.); Legehnen, 4 (U.S.N.M.); Lochstedt, 4 (U.S.N.M.); Baltic
coast, 5 (U.S.N.M.).

BemarJcs. — This race is readily distinguishable from the other
European forms of Microtus arvalis by its large size, grizzled
greyish colour, and robust, strongly angled skull. So far as at
present known it is confined to the coast i-egion of extreme
eastern Germany.

Microtus arvalis levis Miller.

1908. Microtus levis Miller, Ann. and Mag. Nat. Hist., 8th ser., j, p. 197,

February, 1908. Type in British Museum.
1910. Microtus levis Trouessart, Faune IMamm. d'Europe, p. 182.

Type locality. — Gageni, Prahova, Roumania (at foot of
Carpathians, north-west of Bucharest).

Geographical distribution. — Roumania, southern Hungary and
north-eastern Italy ; limits of range not known.

Diagnosis. — Size and external appearance essentially as in
M. arvalis duplicatus ; skull narrow and rounded, the interorbital
region not developing a distinct ridge until late in life ; brain-case
long and narrow, its length measured from interorbital constric-
tion to condyle greater than zygomatic breadth.

688

RODENTIA

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Colour. — The colour is less yellowish than in M. arvalis
nrvalls, the light element a pale ci'eam-buff, and the black tips to
the hairs slightly more conspicuous, the grizzle of upper parts
thus tending to be more evident, particularly along middle of
back. Feet dull whitish. Tail rather distinctly bicolor, dark
brown above, bufty white below. Underparts in most specimens
rather strongly suffused with pale cream-bufF.

Skull and teeth. — The skull differs from that of the other
European races of Microtus arvalis in its more elongate form,
less widely spreading zygomata, and in a peculiar general smooth-
ness and lack of angularity. Brain-case elongated, the least
distance from interorbital constriction to back of condyle decidedly
greater than zygomatic breadth, while in the related forms it is
less tlian zygomatic breadth or at most barely equal. Depth of
brain-case rather greater than in true arvalis, in this respect
resembling duplicatus. General surface of skull noticeably
smoother and less angular than in the related forms, the inter-
orbital region and sides of brain-case rarely developing evident
ridges. Auditory bulke UvSually larger and more inflated than in
true arvalis, but this character not entirely constant. Teeth with
no special peculiarities.

Measurements. — External measurements of type (adult male) :
head and body, 110 ; tail, 38 ; hind foot, 17*2; ear from meatus,
1 1 . Average and extremes of ten adults from the type locality :
head and body, 102-8 (101-110); tail, 35-5 (31-38); hind
foot, 1 6 • 6 (1 6 • 2-1 7 • 4) ; ear from meatus, 1 0 • 4 ( 1 0-1 1 • 5). For
cranial measurements see Table, p. 689.

Specimens examined. — Thirty-two, from the following localities : —
Austria-Hungary: Cepin, near Eszek, Hungary, 8.
RouMANiA : Gageni, Prahova, 17 ; Bustenari, Prahova, 1 ; near
Bucharest, 3 (Mottaz) ; Dobrudscha, 1.
Bulgaria: Rustschuk, 1 (Andersen).
Italy: Caorle, Venice, 1.

2 & 6 al. Cepin, Eszek, Hungary. Budapest Museum (e) . 94. 7. 23. 13-20.

5(^,6 9. Gageni, Prahova, Ron- Lord Lilford (p). 4. 4. 6. 53-G3.

mania. {W. Dodson.) (4. 4. 6. 55. Type of subspecies.)

6. Bustenari, Prahova. Lord Lilford (p). 4, 4. 6. 64.

(W. Dodson.)

1. Dobrudscha, Roumania. Purchased (Pruliere). 86. 4. 2. 6.

MICROTUS INCERTUS de Selys-Longchamps.

1841. Arvicola incertiis de Selys-Longchamps, Atti della Seconda Riunione

degli Scieuziati Italiani, Torino, 1840, p. .225 (near summit of

St. Gothard, Switzerland).
18G9. [Arvicola arvalis] var. fulva Fatio, Faune Vert. Suisse, i, p. 236

(near summit of Furka, Switzerland). Type in Geneva Museum.
1905. [Arvicola arvalis] var. flava Fatio, Arch. Sci. Phys. et Nat. Geneve,

4th ser., xix, p. 195, February 15, 1905 (Renaming oi fulva).
1905. [Microtus incertus] Major, Ann. and Mag. Nat. Hist., 7th ser., xv,

p. 511, May, 1905.

691

1907. Microtus arvalis inccrtus Mottaz, M6m. Soc. Zool. de France, xx,

p. 32, September, 1907.
1910. Pitymys inccrtus Trouessart, Fauna Mamm. d'Europe, p. 188.

Type localitij. — Near summit of St. Gothard Pass, Uri,
Switzerland.

Geographical distribution. — ^Mountains of Switzerland and
Tirol, from the central Alps eastward. Details of distribution
very imperfectly known.

Diagnosis. — Like Microtus arvalis, but skull tending to assume
a distinctly fossorial aspect, the occipital region depressed (ratio
of occipital depth to occij^ital width about 50*5 instead of 58),
the rostrum elongated (diastema
usually about 8 mm. in adults instead
of about 7*5 mm.), the upper incisors
projecting forward ; auditory bulhie
small and flattened.

External characters. — So far as
can be determined from skins, the
external characters are in all essen-
tials as in Microtus arvalis, though
from the peculiarities of the skull it
seems probable that the head is more
flattened and the incisors more pro-
truding ; fur tending to be longer and
less dense.

Colour. — In a series of skins the
colour is, on the average, less bufiy
above than in Microtus arvalis arvalis,
and the underparts are more fre-
quently a clear, slate-tinged grey. The colours are dull and
blended, with very slight indication of grizzling, even in
individuals with elements of the colour nearly as in M. arvalis
duplicatus.

Measurements. — Adult male and female from Andermatt,
Uri, Switzerland: head and body, 114 and 108; tail, 41 and
37; hind foot, 16 and 15-5. Adult male from Furka Pass,
Switzerland: head and body, 113; tail, 31 ; hind foot, 15-4.
Average and extremes of eight adults from Vulpera-Tarasp,
Grisons, Switzerland: head and body, 111 (104-119); tail, 36*5
(30-41); hind foot, 16-4 (15-6-17); ear from meatus, 11-7
(11-13). For cranial measurements see Table, p. 692.

Specimens examined. — Sixty-six, from the following localities : —
Switzerland: Furka Pass, 31 (B.M., U.S.N.M., Geneva and Mottaz) ;
St. Gothard Pass, 2 (B.M. and U.S.N.M.) ; Andermatt, Uri, 7 (U.S.N.M.) ;
Vulpera-Tarasp, Grisons, 16 (Rothschild); Campfer, Grisons, 7 (Roths-
child) ; Engadine, 1.

Austria-Hungary : Meran, Tirol, 1 ; Paneveggio, Tirol, 1.

Bemarlcs. — Microtus inccrtus is readily distinguishable from
M. arvalis by its flattened brain-case, small auditory bullae and

2 Y 2

Fig. 135.
Microtus incertus. Nat. size.

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MICROTUS 693

protruding vipper incisors. In immature specimens the skull is
strikingly like that of Pitymys subterraneus, but when fully adult
the interorbital region becomes narrow and ridged, so that the
resemblance to Pitymys is lost.

6, 9. Furka Pass, Switzer- Lord Lilford (p). 1. 11. 7. 13-14.
land.

2. St. Gothard Pass. Purchased. 46. 6. 15. 44, 114.

6. Engadine (Fatio). E. R. Alston (p). 79. 9. 25. 51.

S al. Meran, Tirol, Aus- Dr. V. Eric (p). 90. 1. 30. 1.

tria-Hungary.

? al. Paneveggio, Tirol. Marquis G. Doria (p). 90. 3. 5. 15.

MICROTUS ASTURIANUS Miller.

1908. ilicrofus asttirianus Miller, Ann. and Mag. Nat. Hist., 8th scr., i,

p. 198, February, 1908.
1910. Microtus asturianus Trouessart, Fauue Mamm. d'Europe, p. 181.

Ty]pe locality. — Pajares, Leon, Spain.

Geographical distribution. — Austurias, and Sierra de Guadar-
rama, Spain. Details of distribution imperfectly known.

Diagnosis. — Larger than Microtus arvalis (hind foot nearly
20 mm.), though similar in all general features. Skull massive
and deep, the dorsal profile strongly convex, the brain-case short
and broad, the zygomata widely spreading ; interorbital region
with evident ridges ; auditory bulky very large. Colour about as
in M. arvalis meridianus.

Colour. — Upper parts buffy clay-colour, rather coarsely
" lined " with black along median dorsal area, clearer and more
nearly approaching ochraceous-buff on sides ; underparts dull
grey, clear or washed with light buff; feet an indefinite bufFy
grey tinged with drab, not conspicuously different from colour of
back ; tail obscurely bicolor, buffy grey below, brownish mixed
with grey above.

Shull. — The skull is larger than in M. an-alis, its general
aspect so robust as to suggest that of a small M. orcadensis.
Zygomata widely spreading, strongly bent downward, not con-
spicuously widened at middle. Dorsal profile more strongly
convex than in any of the forms of arvalis, in this respect
resembling M. cahrerse and M. liartingi ; this peculiarity is due
chiefly to the great depth of the skull through interorbital
region. Brain-case short and broad, its general outline rather
more squarish than in M. arvalis, and suggesting that of
M. orcadensis. Auditory bullaj relatively as large as in M. hartingi,
therefore noticeably exceeding those of true arvalis, and slightly
larger than those of levis. Palate normal. Nasals strongly
cuneate, not peculiar in form, and not in any way suggesting
those of M. cabrerse, the posterior border sharply angular-
emarginate. Temporal ridges moderately developed.

Teeth. — Except for their greater size, a character noticeable

694

RODENTIA

in both incisors and molars, the teeth do not differ from those
of Microtus arvalis. First lower molar with anterior outer
re-entrant angle normally developed.

Measurements. — External measurements of type (adult
female) : head and body, 120 ; tail, 37 ; hind foot, 20 ; ear from
meatus, 14. Young adult male from La Granja, Segovia,
Spain: head and body, 100 ; tail, 38; hind foot, 19 ; ear from
meatus, 12*8. For cranial measurements see Table, p. 707.

Speciviens examined. — Seven, from the following localities in Spain: —
Pajares, Leon, 3 ; La Granja, Segovia, 4.

c5, 2 9. Pajares, Leon, Spain. 0. Thomas (p).' 8. 2. 9. 205-207.

(N. Gonzalez.) (8. 2. 9. 206. Type of species.)

2(5, 2juv. al. La Granja, Segovia. Purchased (Escalera). 8.7.30.15-18.

MICROTUS ORCADENSIS Millais.

1904. Microtus orcadensis Millais, The Zoologist, 4th ser., viii, p. 244,

July, 1904.

1905. Microtus orcadensis Major, Ann. and Mag. Nat. Hist., 7th ser., xv,

p. 324, March, 1905.
1909. Microtus orcadensis Trouessart, Faune Mamm. d'Europe, p. 177.

Type locality. — Pomona Island, South Orkney Islands,
Scotland.

Geographical distribution. — South Orkney Islands ; known at
present from Pomona, Rousay, Shapinshay and South Ronald-
shay.

Diagnosis. — Much larger than Microtus arvalis (hind foot, 18
to 20 mm., condylobasal length of skull in largest individuals
about 28 • 5 mm.) ; skull essentially like that of M. arvalis in
form, but slightly broader (ratio of zygomatic breadth to condylo-
basal length about 60 instead of about 56), the depth of brain-
case not however sensibly reduced ; teeth as in M. arvalis, the
second upper molar with no trace of postero-internal loop, the
first lower molar with well developed anterior outer re-entrant
angle ; colour much darker than in any of the other European
species with the same pattern of enamel folding, the underparts
clear ochraceous-buff.

Colour. — General effect of upper parts a rich, dark brown,
nearly approaching the mummy-brown of Ridgway. Individual
hairs slate-black at base, those of the underfur tipped (2 mm.)
with brownish ochi'aceous-buff, the longer hairs clear black but
producing a very slight effect of " lining." On sides the colour
gradually passes into the clear bright ochraceous-buff of under-
parts, this slightly obscured by the slaty bases of the hairs,
especially on chin and throat. Feet and under surface of tail
light grey tinged with ochraceous-buff. Upper surface of tail
blackish.

Shull. — Except for its larger size and relatively shorter, more

695

Fig. 136.
Microttts orcadensix. Nat. size

widely spreading zygomatic arches, the skull difters little from
that of typical Microtus arvalis. Central portion of zygoma
noticeably widened, a character ap-
preciable in individuals that are still
evidently sub-adult and giving the
bone a decidedly difterent appearance
from the slender arch of arvalis.
Interparietal with longitudinal dia-
meter greater and lateral diameter
less than in arvalis, the resulting
form more sub-quadrate than in the
smaller animal. Median ridge of
posterior termination of palate almost
invariably with evident groove along
suture. Tooth-rows relatively
smaller and less widely separated
than in M. arvalis, the distance be-
tween them the same as in the smaller
species. Auditory bulla; rather small
relatively to size of skull, their actual
measurements not much greater than
in M. arvalis.

Teeth. — In structure the teeth do not differ appreciably from
those of 31icrotiis arvalis, though the enamel pattern in general is
somewhat more compact, owing to the rela-
tively less width of re-entrant angles and
greater area of triangles. Elements of enamel
pattern exactly as in M. arvalis ; m"^ without
trace of postero-internal loop ; m-^ with an-
terior loop deeply cut on outer side by a
re-entrant angle approximately as well de-
veloped as that of inner side, a character
that shows no appreciable variation in a series
of fifty-three skulls.

Measurements. — External measurements of
type (old male), from skin : tail, 32 ± ; hind
foot, 18. Average and extremes of ten adult
males from Pomona Island : head and body,
118 (113-125); tail, 38-5 (35-44); hind foot,
18*7 (18—20); ear, 12 (11-13). Average and extremes of six
females from Pomona Island: head and body, 117 (114-125);
tail, 38 • 4 (34 • 5-42 • 5) ; hind foot, 18-6 (18-5-19); ear, 11-5
(11-12). For cranial measurements see Table, p. 699.

Microtus orcadensh
Enamel pattern. X

Specimens examined. — Eighty, from the following islands of the
South Orkney group : Rousay, 7 (B.M. and Edinburgh) ; Pomona, 64
(B.M., U.S.N.M. and Edinburgh); Shapinsay, G (Edinburgh); South
Ronaldshay, 3.

BemarJcs. — From the other European voles with similar

696

RODENTIA

enamel pattern this species is so readily distinguishable by its
large size and dark colour that it needs no special comparisons.
The nearest known relatives, apart from M. sandayensis, are
Mierotus sarnius of Guernsey and the extinct Microtus corner i
Hinton,* from the Late Pleistocene deposits of the Ightham
Fissure, Kent, and Langwith Cave, Derbyshire.

6, 1 al. Eousay Island, Orkneys. W. Cowan (c & p). 4. S. 22. 1-2.

6. Sandwick, Pomona J. G. Millais (c & p). 4.6.21.1.

Island. {Type of species.)

6(5.5 9. Loch Stenues, Pomona N. B. Kinnear (c & p). \ n' -i r,' o' i~ a'

6, 2 9. S. Ronaldshay Island. Edinburgh Museum (p). 7. 11. 16. 1-3.
6, 9. Orkneys. ■ J. G. MiUais (c & p). 5. 11. 22. 1-2.

MICROTUS SANDAYENSIS Millais.

(Synonymy under subspecies.)

Geographical disirihutioii. — North Orkney Islands, Scotland.

Diagnosis. — Like Microtus orcadensis but size slightly less
(hind foot, 17-19 mm. ; condylobasal length of largest skulls
about 27 '5 mm.), and colour not so dark, the back approaching
hair-brown rather than mummy-brown ; skull with brain-case
noticeably flattened, auditory bullae somewhat reduced in size,
and zygoma showing little if any tendency to become expanded
at middle.

Bemarhs. — For detailed descriptions see accounts of the two
subspecies.

The voles of the North Orkney Islands are distinguished
specifically from Microtus orcadensis of the southern group by
the peculiar flattened form of the brain-case. The degrees of
distinctness of the Orkney voles among themselves appear to
bear a direct relation to the depth of water separating the
islands, and therefore presumably to the length of time that the
different colonies have been isolated. The numerous specimens
of M. orcadensis examined from four islands of the southern
group show no indication of the existence of local forms ; the
depth of the channels separating these islands from each other
ranges from six to eight fathoms. Between the islands inhabited
by this species and those occupied by M. sandayensis lies a narrow
but comparatively deep strait, with seventeen to twenty fathoms
of water. Finally between Sanday and Westray with their
different though not completely segregated forms, the depth of
the water is intermediate, ten to twelve fathoms.

* Ann. and Mag. Nat. Hist., 8th ser., vi, p. 35, July, 1910. The
characters by which the fossil is distinguished from Microtus sandayensis
are less apparent than those separating the two living Orkney species.

MICROTUS 697

MiCKOTUS SANDAYENSIS SANDAYENSIS Millais.

1905. Microtus orcadensis sandayensis Millais, Mamm. Great Brit, and

Ireland, ii, p. 280.
1908. Microtus sandayensis ]\Iiller, Ann. and Mag. Nat. Hist., 8th ser., i

p. 199, February, 1908.
1910. Microtus sandayensis Trouessart, Faune Mamm. d'Europe, p. 177.

Tijpe locality. — Sanday Island, North Orkney Islands.

GeograpMcal distribution. — At present known from Sanday
Island only.

Diagnosis. — First lower molar with anterior outer re-entrant
angle obsolete, much less developed than anterior inner angle ;
upper parts greyish brown, much lighter than in M. orcadensis ;
underparts whitish grey a little washed with buffy.

Colour. — General effect of upper parts a clear, light hair-
brown, the individual hairs slate-black at base, those of underfur
tipped with a pale buffy drab, the exact shade between the
cream-buff and ecru-drab of Ridgway, the longer hairs black but
producing almost no effect of " lining." Underparts light grey
with a faint buffy tinge. Feet essentially like underparts. Tail
not distinctly bicolor, greyish white throughout but rather
strongly darkened by a sprinkling of blackish hairs along median
dorsal surface.

Skull. — The skull, as already pointed out, is readily distinguish-
able from that of M. orcadensis by its lower, more flattened brain-
case, a character equally apparent whether the skulls are viewed
from above or behind. Zygomata scarcely expanded at middle,

Fig. 138.

Microtus sandayensis (a) aud («') ;
^f. orcadensis (6). Nat. size.

their form in this respect resembling that of M. arvalis rather
than that of M. orcadensis. They are, however, as short and as
widely spreading as in the more nearly related species. Inter-
parietal as in M. orcadensis. Auditory bullae slightly smaller
than in orcadensis.

698

Teeth. — The teeth resemble those of M. arvalis and M. orca-
densis except in the form of the anterior loop of the first lower
molar. In both arvalis and orcadcnsis this loop
is deeply indented by two re-entrant angles,
one on each side, the two angles approximately
equal in depth. In M. sandayensis sandnijensis
the inner angle remains as in the related
species, but the oiiter, more posterior angle is
much more shallow than the inner. In some
specimens it is scarcely indicated, while in all
of the twelve skulls that I have examined it
is sufficiently difterent from the ordinary type
to permit of certain identification of the animal
by this character alone.

Measurements. — External measurements of
type (immature male measured on dried skin) :
head and body, 86; tail, 19 ; hind foot, 17 "4.
Adult male and two adult females from the type locality : head
and body, 117, 110 and 111; tail, 37-5, 38-5 and 35; hind
foot, 19, 18 and 18; ear from meatus, 11, 10'5 and 11.
For cranial measurements see Table opposite.

Specimens examined. — Twenty, all from Sanday Island, North Orkneys
(B.M., U.S.N.M. and Edinburgh).

J. G. Millais (c & p). 5. 11. 22. 3.

{Type of species.)
N. B. Kinnear (c & p). 6. 11. 18. 7-8.
W. R. Ogilvie - Grant 11,11.2.1-8.
(c & p).

Fig. 139.

Microtvs sandayensis
sandayensis. X 5.

9. Sanday Island, Orkneys.

2 9.
4(J49

Sanday.
Sanday.

MiCROTUS SANDAYENSIS WESTR^ Miller.

1908. Microtus sandayensis ivestrx Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 199, February, 1908.
1910. Microtus sandayensis westrx Trouessart, Faune ]\Iamm. d'Europe,

p. 178.

Tyjje locality. — Westray Island, North Orkney Islands.

Geographical distribution. — At present known from Westray
Island only.

Diagnosis. — First lower molar with anterior outer re-entrant
angle occasionally as deep as in 31. orcadcnsis ; colour not so pale
as in the Sanday vole, the underparts strongly washed with
yellowish brown.

Colour. — General eftect above a dark hair-brown approaching
bister, the arrangement of colour as in sandayensis, but light tips
to hairs of underfur more nearly a dull ochraceous-buff, and dark
shading from longer hairs more noticeable. Underparts light
ochraceous-bufi', nearly as in M. orcadcnsis, but colour not so rich
and clouding from slaty bases of hairs more evident. Feet and
tail as in sandayensis, but sprinkling of dark hairs on upper
surface of tail more conspicuous.

699

Observations.

Ridges scarcely visible.

Old.

Fully adult.

Old. Ridges joined.

Ridges nearly joined.

Ridges scarcely visible.

Immature.

Fully adult.

Old.

Fully adult.

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M. orcadensis.
Orkney Islands : Rousay Island

Pomona Island

Shapiushay .
South Ronaldshay

700 EODENTIA

Skull. — The skull so exactly resembles that of Micrvtns sati-
dayensis sandaycnsis as to require no detailed description. Even
in a very old individual (Kinnear No. 321) the zygoma shows no
indication of the central expansion characteristic of M. orcadensis.

Teeth. — The anterior loop of the lirst lower molar is inter-
mediate in form between that of orcadensis and true sandayensis,
though apparently nearer the latter. The outer re-entrant angle
tends to be shallow, so that its depth is usually much less than
that of inner angle, but the degree of tliis flattening varies so
much that specimens cannot be identified by this character alone,
while in at least one specimen (No. 321 Kinnear) the conditions
are exactly as in M. orcadensis.

Measurements. — External measurements of type (adult female) :
head and body, 108 ; tail, 33 ; hind foot, 17 ; ear from meatus, 12.
Adult male and female from the type locality : head and body,
115 and 108 ; tail, 42 and 38 ; hind foot, 18 and 17 ; ear from
meatus, 11 and 11. For cranial measurements see Table,
p. 699.

Specimens examined. — Seven, all from Westray Island, North Orkneys
(B.M. and Edinburgh).

6. Westray Island, Orkneys. N. B. Kinnear (c & p). 8. 1. 2. 1.

{Type of subspecies.)

MICROTUS SARNIUS Miller.

1909. Microtus sarnius Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

p. 420, May, 1909.

1910. Microtus sarnius Trouessart, Faune Mamm. d'Europe, p. 178.

Type locality. — St. Martins, Guernsey, Channel Islands.

Geographical distribution. — ^Island of Guernsey.

Diagnosis. — Size about as in Microtus orcadensis or slightly
less (hind foot about 18*5 mm., condylobasal length of skull
about 28 mm.) ; skull narrower than that of orcadensis, its
appearance in old individuals essentially as in the larger races of
M agrestis ; colour agrestis-like, the underparts sharply con-
trasted pale grey.

Colour. — The colour in summer in practically identical with
that of Microtus agrestis agrestis, though perhaps in most specimens
somewhat less dark and reddish. Underparts a light grey
(about the grey No. 9 of Ridgway) strongly contrasted with
wood-brown of sides, the belly sometimes with a faint buffy
tinge, the slate-grey appearing irregularly at surface, especially
in abraded skins.

Shull. — In fully adult skulls the size and general appearance
is essentially as in the large forms of Microtus agrestis, though
the interorbital region is both longer and wider, the brain-case
is more depressed posteriorly, and the interparietal is noticeably
more quadrate in form, its antero-posterior diameter distinctly
more than half transverse diameter, its lateral extremities

MICROTOS 701

squarely truncate. In younger specimens (temporal ridges
evident, but not joined) the brain-case is broader and the general
form more nearly approaches that of M. orcadensis at the same
stage, though the zygomata are longer and less abruptly
spreading, and the auditory bullae are smaller.

Teeth. — In all respects except for their slightly smaller size
the teeth resemble those of Microtus orcadensis. First lower
molar with well developed antero-external re-entrant angle.

Measurements. — External measurements of type (adult male) :
head and body, 118; tail, 42; hind foot, 18'5; ear from
meatus, 12. Adult male and female from the type locality:
head and body, 115 and 114-5 ; tail, 44 and 34 ; hind foot, 18*4
and 17 '5 ; ear from meatus, 9*5. For cranial measurements see
Table, p. 707.

Specimens examined. — Fourteen, all from the island of Guernsey.

Bemarks. — Misled by the extreme narrowness of the very
aged type skull, I at first supposed this animal to be a member of
the agrestis group with aberrant dentition. Further material
shows that it is related to the voles of the Orkney Islands and to
the extinct Microtus corneri of the British mainland, the three
living species and their fossil relative apparently belonging to an
older fauna than that now inhabiting Great Britain and the
mainland of Europe.*

i 6,19. St. Martins, Guernsey. 0. Thonaas (p). 8. 9. 2. 24-28.
{R. H. Bunting.) (8. 9. 2. 27. Type of species.)

MICROTUS CABRERA Thomas.

1906. Microtus cabrerx Thomas, Ann. and Mag. Nat. Hist., 7th ser., xvii,

p. 576, June, 1906.
1910. Microtus cabrerai Trouessart, Faune Mamm. d'Europe, p. 181.

Tyjje locality. — Rascafria, south side of Sierra de Guadarrama,
Province of Madrid, Spain.

Geographical distribution. — Probably throughout the moun-
tainous region of central Spain, but at present only known with
certainty from the type locality.

Diagnosis. — A large member of the arvalis group, equal in
size to M. orcadensis (hind foot, 22 mm. ; condylobasal length of
skull, 27 mm.). Readily distinguishable from all its allies by the
strongly convex dorsal profile of the skull, deep, rounded brain-
case, slightly tapering nasals and (in young adult skulls at least)
the conspicuous longitudinal furrow in interorbital region.
Externally the type is peculiar in the unusual length and
conspicuousness of the longer hairs above, those of the sides and
rump with noticeable pale tips or sub-terminal annulations.

Colour. — Hairs of upper parts slate-colour at base, those of

* See Hinton, Ann. and Mag. Nat. Hist., 8th ser., vi, p. 30, July, 1910.

702

the underfur a dull, buff'y wood-brown (2 mm.) at tip, the longer
hairs black, those of sides and rump with conspicuous buffy tips
or sub-terminal areas about 4 mm. in length. In both specimens
the long hairs are more conspicuous than in any other European
Microtus that I have examined. On sides the buffy wood-brown
is nearly clear and distinctly more yellow than the cream-buft' of
Ridgway, but on back it is coarsely and conspicuously "lined"
with black, the general effect of the two colours approaching
olive or bister. Underparts yellowish cream-buff irregularly
clouded by the slaty bases of hairs. Feet a dull, light, brownish

buffy. Tail obscurely bicolor, pale
buffy below, dull brownish above.

iSJcuU. — The skull differs from that
of all other known European species
of 3Ilcrotus in its very highly arched
brain-case, which gives the dorsal
profile a uniform convexity that is ex-
ceedingly characteristic. Interorbital
region, in the two skulls examined,
rather broad, with distinct median
longitudinal concavity and low but
evident lateral ridges.* Nasals
slightly tapering, their width pos-
teriorly about three-fourths that
anteriorly, their posterior termination
square, slightly exceeded by nasal
branch of premaxillary. In both of
the skulls examined the length of
nasal decidedly surpasses that of
diastema, a condition not observed
in other European species. Palatal foramina large, their width
relatively to that of rostrum greater than in
M. arvalis. Auditory bullae not peculiar, their
size relatively about as in M. arvalis.

Teetli. — The teeth are strictly of the M.
arvalis type. Terminal loop of both m-^ and
«ii shortened as compared with the other
European members of the arvalis group, so
that in the former the third inner re-entrant
angle is very shallow, and the extreme pos-
terior portion of the tooth is a nearly straight
loop, almost as wide as long, and evenly con-
stricted from both sides proximally, while in fig. 141.
the latter the outer re-entrant angle is prac- Microtus cahrerx. x 5.
tically absent, this peculiarity being carried
to a greater extreme than in M. sandayensis sandayensis, the
only other European Microtus in which it is known to occur

* The ridges may eventually unite ; but the form of the interorbital
region in the young adult is different from that in any of the other
European species at the same stage.

Fig. 140.
Microtus cabrerie. Kat. size.

MICROTUS 703

as a normal character. Upper incisors short, directed downward
and slightly backward, concealed by nasals when skull is viewed
from above.

Measurements. — External measurements of type : head and
body, 107 ; tail, 34 ; hind foot, 22 ; ear from meatus, 12. For
cranial measurements see Table, p. 707.

Specimens examined. — Two, the type, from uear Rascafria, iu the Sierra
de Guadarrama, Madrid, Spain, and a second specimen without definite
locality.

£. Rascafria, Sierra de Guadarrama, M. de la Escalera (c). G. 11. 4. 9.

Spain. {Type of species )

sk. Spain. Purchased (Parzudaki). 53. 12. G. 31.

MICROTUS DENTATUS Miller.

1910. Microtus cahrene Cabrera, Asoc. Espafi. Progr. Cien., Congr. Zara-

gosa, 1908, p. 49, June, 1910 (part).
1910. Microtus dentatus Miller, Ann. and Mag. Nat. Hist., 8th ser., vi,

p. 459, November, 1910. Type iu Madrid Museum.

Tyi)e locality. — Molinicos, Sierra de Segura, Albacete, Spain.

Geographical distribution. — Known only from the Sierra de
Segura.

Diagnosis. — Similar to Microtus cabrerse of the Sierra de
Guadarrama, but larger, the skull about 30 mm. in condylobasal
length ; teeth excessively heavy, the molars larger than in any
other known European Microtus, the length of maxillary tooth-
row 8 mm. ; jh, with a completely closed triangle on outer side.

Colour. — Like that of 31. cabrerse, but noticeably less buff,
the ground colour of back and sides not so yellow as the cream-
buff" of Ridgway, the general effect a peculiar buffy grey much
paler than the olive or bister of cabrerse ; underparts light grey
scarcely tinged with buff" ; no evident line of demarcation along
sides.

Sltull. — The only known skull is imperfect, lacking the auditory
bulhe and base of brain-case. It is that of an individual slightly
older than the type of Microtus cabrerse, but of about the same
age as the fragmentary skull of No. 55. 12. 6. 31. Size conspicu-
ously greater than in either of these ; general form the same,
but dorsal profile less concave than in the type, especially over
posterior half of brain-case ; rostrum much heavier than in
M. cabrerse, the incisive foramina fully Ih times as wide though
of about the same length • relative lengths of nasal and diastema,
and form and position of posterior border of nasals as in
M. cabrerse ; interorbital region longer and narrower, but lateral
ridges and median groove not peculiar. Mandible as in M.
cabrerse, the base of articular process scarcely thickened by
upward extension of incisor root, the dental foramen lying almost
at extreme posterior edge.

Teeth.^In general the teeth are like those of Microtus cabrerse,

704

EODENTIA

but are immediately distinguishable by the unusually large size
of the molars. Posterior loop of m^ and anterior loop of vi^ rather
short ; most of the triancrles showing an exaggeration of the

Fig. 142.
Microtus dentatuf!

X 5.

tendency to flattening characteristic of caltrerse. ; vu_ with a large,
completely closed triangle on outer side.

Measurements. — External measurements of type : head and
body, 125; tail, 40; hind foot (dry), 22. For cranial measure-
ments see Table, p. 707.

Specimen examined. — The type (Lladrid).

MICROTUS HARTINGI Barrett-Hamilton.

1903. Microtus [Microtus) liartingi Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7tli ser., xi, p. 307, March, 1903. Type in British Museum.
1910. Microtus liartingi Trouessart, Faune Mamm. d'Europe, p. 174.

Type locality. — Larissa, Thessaly, Greece.

Geograpldcal distribution. — Known only from the vicinity of
Larissa.

Diagnosis. — Size large, slightly exceeding that of Microtus
orcadensis and M. cahreree (hind foot about 18, condylobasal
length of skull about 29 mm.) ; tail relatively short ; colour more
yellowish than in any of the European forms of 31. arvalis ; skull
with rather deep brain-case and slightly convex dorsal profile ;
teeth as in M. arvalis, but triangles narrower relatively to their
width, and area of dentine spaces reduced as compared with
thickness of enamel.

Colour. — Back and sides a light dull buff, perhaps nearest the
pinkish buff of Ridgway, clear on flanks and thighs, but else-
where grizzled by the blackish tips of the longer hairs, these
most numerous over posterior half of back ; underparts, feet, and
tail bufiy ^vhite, the dorsal surface of tail slightly more yellow

MICROTUS

705

(about concolor with flanks), the belly irregularly clouded by

the slaty bases of the hairs.

Skull. — The skull, though heavier and more massive than in

any of the European members of the M. arvalis, has much the

same general outline as in typical arvalis. The brain-case is,

however, relatively deeper anteri-
orly, so that the dorsal profile is

convex throughout (though much

less so than in the Spanish 31. cah-

rerse). Interorbital region with

slight median longitudinal groove

and low lateral ridges which do not

tend to become united in any of

the four skulls examined. Nasals

strongly tapering posteriorly as in

31. arvalis, their termination a little

emarginate and slightly exceeded

by nasal branches of pre-maxil-

laries. Palatal foramina narrow, as

in M. arvalis. Auditory bullae not

peculiar in form, their size relatively

somewhat greater than in 31. arvalis.
Teeth. — While preserving the

typical arvalis pattern, the teeth

are peculiar in a certain disjointed

appearance of the prisms, due to

the great depth of the re-entrant angles. The triangles are

rather narrower in proportion to their width than in the other
members of the arvalis group, and the dentine
spaces have the appearance of being reduced
in proportion to the thickness of the enamel,
a character difficult to define but readily
appreciable to the eye. Except for these
general peculiarities the molars closely agree
with those of M. arvalis. First lower molar
with normal anterior loop, the outer re-
entrant angle well developed and approxi-
mately as deep as inner angle. Posterior
upper molar with the same elements as in
31. arvalis.
FIG. 144. 3Ieasurements. — External measurements of

Mkruius hartiniji. X 10. type (adult male): head and body, 107; tail,
26 ; hind foot, 18. A second adult male from

the type locality: head and body, 115; tail, 27; hind foot, J 8^;

ear from meatus, 10. For cranial measurements see Table, p. 707.
Specimens examincd.—'Eight, all from the type locality.

Bemarks. — Though very difierent from the other European
species of 3Iicrotus, this animal is nearly related to M. guentheri

2 z

Fig. 143.
.Uicrotus hartingi. Nat. size.

700 RODENTIA

Danford and Alston, of Asia Minor. It differs merely in its
slightly paler, more yellowish colour, somewhat more robust
skull, and more inflated auditory bullte.

6, 2 al. Larissa, Thessaly, J. E. Harting (p). 93. 4. 5. 1-3.

Greece. (93. 4. 5. 1. Type of species.)

6 al. Larissa, Thessaly. P. H. Mavrogordato 93. 3. 28. 1.

(C & 1').

6, 9, 2 juv. al. Thessaly. Dr. F. Loefflcr (c & p). 92. 12. 7. 1-4.

MICROTUS ANGULARIS Miller.

1908. Microtus ancjularis Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 198, February, 1908. Type in British Museum.
1910. Microtus angularis Trouessart, Faune Mamm. d' Europe, p. 182.

Type locality. — Transylvania (probably near Hatszeg, Hun-
yad, Hungary).

Geograjjhical distribution. — Known only from the type locality.

Diagnosis. — A large member of the Microtus arvalis group,
about equal to the Grecian M. hartingi in size, but with much
longer tail ; skull with unusually deep rostrum and abruptly
sloping nasals.

Colour. — After about thirty years immersion in alcohol the
type appears to be essentially like Microtus arvalis arvalis in
colour.

Shill and teeth. — The skull differs from that of all other

known European species of Microtus in the very deep, relatively

short rostrum (depth at back of nasal decidedly greater than

distance between front of zygoma and anterior extremity of

nasal) and the unusually conspicuous angle

r^Cr /TN. (about 34° instead of about 18° to 22° as in

r^K\ X^07 N ^' ^^^^^'^) -^- hartingi and M. orcadensis)

K^^-XxxcDSiiff *^^u ^^ which the nasals slope downward.

t/ Auditory bullae relatively larger than in

Fig. 145. jf arvalis, but not so much inflated as

Nat. size. m M. hartingi. Jjorni or brani-case not

known. Teeth with pattern of enamel

folding as in Microtus arvalis, but all of the triangles, especially

those in lower molars, with transverse diameter noticeably

increased relatively to antero-posterior diameter, and area of

dentine spaces reduced.

Measurements. — ^Type (adult male, in spirit) : head and body,
115 ; tail, 41 ; hind foot, 18 ; ear from meatus, 12. For cranial
measurements see Table opposite.

Specimen examined. — The type.

6 al. Transylvania. C. G. Danford (c & p). 80. 10. 28. 2.

{Type of species.)

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MICROTUS RATTICEPS Keyserling and Blasius.

184]. Arvicola ratticeps Keyserliug and Blasius, Mem. pr&entcs a I'Acad.
Imp. des Sci. Nat. de St. P6tersbourg, iv, livr. 3, p. 333, 1841
(date on title-page of completed volume, 1845). (Wclikii-Ustjug,
north-central Russia.)

1841. Arvicola arcnicola de Se'lys-Longchamps, Bull, de I'Acad. Royale
des Sci. des Arts et Belles-Lettres de Bruxelles, viii, pt. 2, p. 236
(Lisse, near Leiden, Holland. See Jentink, Notes from the
Leiden Museum, xxix, p. 263, February 29, 1908).

1844. Arvicola mcdius Nilsson, Ofversigt af Kongl. Vetenskaps-Akademiens
Forhandlingar, Stockholm, i, p. 34, March 20, 1844 (Lapland, and
the mountains about the Gudbrandsdal, Norway).

1857. Arvicola ratticcps Blasius, Siiugethiere Deutschlands, p. 3G5.

1887. Microtus ratticeps Lataste, Ann. Mus. Civ. Stor. Nat. Geneva,
2ud ser., iv, p. 265, January, 1887.

1899. Arv[icola'\ (Microtus) ratticcps var. stimmincji Nehring, Sitz.-Ber.
Gesellsch. Naturforsch. Freunde, Berlin, p. 58 (near Brandenburg,
Germany). See Rorig, Arbeiten aus der Kais. Biol. Anstalt fiir
Land- und Forstwirtschaft, vii, p. 471, 1909.

1910. Microtus ratticcps, M. ratticeps stimmingi, and M. arcnicola Troues-
sart, Faune Mamm. d'Europe, p. 179.

Type locality. — ^Welikii-Ustjug, on the Dwina River, north-
centi'al Russia.

Geographical distribution. — Northern portions of continental
Europe and Asia from northern Russia and the mountains of
Scandinavia eastward into Siberia, south to Holland,* northern
Germany and northern Hungary.

Diagnosis. — ^izG large (hind foot, 19-21 mm. ; condylobasal
length of skull in largest individuals, 30 mm.) ; second upper
molar without postero-internal triangle ; first lower molar with
only three re-entrant angles on outer side ; skull long and slender
(ratio of zygomatic breadth to condylobasal length about 50), the
intertemporal region developing moderate ridge with age ; length
of brain-case measured from interorbital constriction to condyle
greater than zygomatic breadth.

External characters. — Essentially as in Microtus agrcstis, but
thumb usually with somewhat better developed nail, and sole
with sixth tubercle relatively smaller.

Colour. — In general the colour is not essentially different
from that of Microtus agrcstis, though it shows a tendency to be
somewhat darker. Upper parts a coarse mixture of wood-brown
and black, the sides occasionally with a slight tinge of russet.
Underparts abruptly-defined pale grey tinged with cream-buff"
and frequently darkened by slaty under-colour. Feet light
drab or dull whitish. Tail rather sharply bicolor, dark brown
above, whitish below. In a skin from Western Hungary
(B.M. 94. 3. 1. 64) there is a noticeable dark dorsal area extend-
ing from forehead to base of tail, rather sharply defined clear

* Not known to occur in Denmark [see Winge, Danmarks Fauna,
Pattodyr, p. 78, 1908).

709

blackish brown on nape, elsewhere a mere darkening of normal
colour.

Skull. — The skull attains a greater length than that of
Microtus agrestis, but the zygomatic breadth and breadth of
brain-case scarcely if at all exceed the corresponding dimensions
in the smaller animal, so that the general form is conspicuouly
less robust. Dorsal, ventral and occipital profiles essentially
as in 31. agrestis, except that there is usually a slight concavity
in interorbital region, and the contrast between occipital and
rostral depths is less pronounced. Brain-case long and narrow,
slightly oval in outline, the moderately developed postorbital
processes not sufficiently prominent to cause the anterior portion
to appear scjuare ; upper surface rather strongly arched laterally,
not flattened as in 31. agrestis ; lateral ridges moderately
developed ; interpai'ietal relatively
small, its area distinctly less than
half that of parietal, its general
outline almost evenly biconvex except
for a slight median projection on
anterior border ; occiput when viewed
from behind essentially as in M.
agrestis, but more rounded above ;
floor of brain-case without special
peculiarities, but region of basal
suture depressed rather than ele-
vated (as in 31. agrestis) when skull
is held upside down ; auditory bulla
somewhat larger and more inflated
than in 31. agrestis, but not peculiar
in form. Interorljital region narrow
as in 31. agrestis, but more elongated.
Zygomata standing out from sides
of skull noticeably less than in 31.
agrestis and 31. arvalis ; median
expansion well defined, rather more
abrupt than usual ; infraorbital foramen wide, its lower border
less extended downward behind root of incisor than in 3Iicrotus
agrestis and 31. arvalis. Rostrum with no special peculiarities ;
nasals terminating posteriorly on level with nasal branches of
premaxillaries a little in front of middle of zygomatic root ;
incisive foramina shorter than in 31. agrestis, strongly contracted
posteriorly, terminating a little in front of level of alveolus of
m^ Bony palate with no special peculiarities, but lateral pits
rather small, and median sloping ridge broad and flat, somewhat
as in the sub-genus Chionomys. Mandible as in 31. agrestis,
but more slender.

TfeiA.— Incisors essentially as in 3I{croius agrestis, but front
face of upper teeth less oblicjuely rounded off" at outer side.
Size of molars relatively to that of skull about as in 3Iicrotus

Fia. 146.

Microtus ratticeps. Nat. size.

710

RODENTIA

Fig. 147.
Mierotas ratticeps.
Enamel pattern.

X5.

agrestis or perhaps a trifle less. Enamel pattern agreeing with
that of 31. arvalis in the absence of all trace of a postero-intei-nal
loop in m'^, as well as in the characters of m^, m„ and iHo, common
to nearly all the European members of the sub-genus ; there is
evei'ywhere, however, less contrast in size and depth between
tlie inner and outer angles and triangles. Third upper molar
frequently of the more complicated type described under
M. agrestis, the posterior loop with its inner limb isolated as a
definite, well-developed, second inner closed triangle, and the
projection on its outer side tending to assume the form of a
small but almost closed third outer triangle.
First lower molar highly characteristic and
readily distinguishable from that of all the
other European members of the sub-genus ;
posterior loop, three inner and two outer closed
triangles as in M. agrestis, except for the less
contrast in size between the elements of outer
and inner side ; anterior loop so short that it
bears no re-enti'ant angle on outer side, while
that of inner side is reduced to a shallow,
inconspicuous concavity, some trace of which
is, however, apparently always present ; angle
subtending terminal loop on outer side broad
and rather shallow ; antero-internal triangle
usually communicating with anterior loop, this
communication when broadly open frequently converting the two
elements into a secondary terminal loop of irregularly crescentic
form not unlike some of the variants of posterior loop of m^.
While differing conspicuously from that of the other European
species of true 3Iicrotus this tooth is essentially like that of the
members of the sub-genus Chionomys. Isolated specimens may
usually be distinguished by the less contrast between size of inner
and outer triangles, and by the absence of any conspicuously
projecting angle at outer base of anterior loop ; in certain forms
of Chionomys, however, the latter peculiarity is almost exactly
rejDroduced.

Measurements. — Average and extremes of nine adults from
Lappmark, Sweden : hind foot (dry), 19*9 (19-4-21). Adultfrom
Csallokoz-Somorja, Pressburg, Hungary : hind foot, 20*8. Adult
female from Texel Island, Holland : head and body, 130 ; tail, 62 ;
hind foot, 20 ; ear, 13. For cranial measurements see Table opposite.

Specimens examined. — Twenty-eight, from the following localities : —

Norway. — Gausdal, 4; Hjerkin, Kristiausamt, 1; Dovre Fjeld, 1
(U.S.N.M.).

Sweden. — Medelpad, W. Norrland, 1 ; Karesuando, Norrbotten, 3 ;
Lappmark, 5 (B.M. and U.S.N.M. ; near Stockholm, 1 (U.S.N.M.).

Finland. — Muonionniska, Uleaborg, 5 (B.M. and U.S.N.M.).

Germany. — Colpin, Brandenburg, 1 ; near city of Brandenburg, 1 (Berlin
Agric. ; coty]3e of stivimingi Nehring).

Austria-Hungary. — Csallokoz-Somorja , Pressburg, Hungary, 1.

711

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EODENTIA

Holland. — Lisse, near Leideu, 4 (Leiden ; cotypes of arenicola de
S61ys-Longcbamps) ; Texel Island, 1.

3 c5. Gausdal, Norway. Christiania Museum (e). 93.3.1.11,13.

6. Gausdal. Miller Collection. 7. 7. 7. 4536.

9. Hjerkin, Kristiansamt. Miller Collection. 7. 7. 7. 4535.

1. Medelpad, W. Norrland, Prof. Sundevall (p). 4'J. 11. 1. 21.

Sweden.

1. Tornea, Lappmark, Swe- Prof. Sundevall (p). 45. 10. 25. 10.

den.

1. Lapland. Prof. Sundevall (p). 49. 11. 1. 9.

2. Lapland. Tomes Collection. 7. 1. 1. 141-142.
1. Lapland. Jliller Collection. 7. 7. 7. 3250.

1. Muonionniska, Uleaborg, Stockholm Museum (e). 90. 8. 1. 7.

Finland.
6. Texel Island. Holland. J. L. Bonhote (c & p). 8. 10. 26. 3.
1 al. Col pin, Brandenburg, C. Gadow (c & p). 81.10.7.1.

Germany.

2. Csall ok oz-Somor j a, Budapest Museum (e). 94.3.1.04-5

Pressburg, Hungary.

Sub-Genus CHIONOMYS Miller.

1857. Paludicola Blasius, Saugetbiere Deutscblands, p. 334 (part). Not of

Wagler, 1830.
1867. Praticola Fatio, Les Campaguols du Bassin du Leman, p. 34 (part).

Not of Swainson, 1837.
1896. Microtus Miller, Nortb American Fauna, No. 12, p. 62, July 23, 1896

(part).
1908. Chionoviys Miller, Ann. and Llag. Nat. Hist., 8th ser., i, p. 97,

January, 1908.

r«/pe species. — Arvicola nivalis Martins.

Geographical distribution. — Mediteri-aneau region from the
Pyrenees to the Caucasus and Asia Minor, mostly in or near
mountains.

Characters. — In general like the sub-genus Microtus, but third
upper molar with only two re-entrant angles on each side, as in
Arvicola and some forms of Pitijmys ; skull with broad, rather
flat, smooth brain-case, and wide interorbital region, the temporal
ridges low and inconspicuous ; posterior termination of palate
essentially as in true Microtus, but with elements usually less
well-defined.

Remarks. — The sub genus Chionomys is a very natural group,
whose characters, while for the most part agreeing with those
of true Microtus, show a distinct tendency toward Arvicola and
Pitymys. Seven forms are now known, all but two of which
occur in Europe. One of these inhabits the lowlands of southern
France, but all of the others are characteristic mountain animals.
The general appearance of the members of this group is highly
characteristic as compared with that of the other European
species of Microtus. They are medium-sized, rather long-tailed
voles with a peculiar slaty greyish general colour, and full, soft
fur, the tail often mostly or entirely whitish.

MICROTUS

13

KEY TO THE EUROPEAN MEMBERS OF THE SUB-GENUS
CHIONOMYS.

Hind foot in adults less than 20 mm. ; back clear
light grey or with very slight brownish tinge ;

tail always white throughout M. lebrunii, p. 718.

Back clear light grey ; auditory bullpe small and
flattened (Neighbourhood of Nimes, Gard,

France) ^f- I- lehrmui, p. 719.

Back with slight brownish tinge ; auditory bulla;
large and strongly inflated (normal) (Basses-

Alpes) M. I. leucurus, p. 722.

Hind foot in adults 20 mm. or more ; back decidedly

brownish; tail often not entirely white.

Posterior termination of palate with median ridge

sharply defined, its width less than that of

deep lateral pit ; back rather heavily clouded

with blackish ; tail usually dark above (Tran-

sylvanian Alps) ill. ulimts, p. 723.

Posterior termination of palate with median ridge
usually not sharply defined, its width always
at least equal to that of shallow lateral pit ;
back slightly or not clouded with blackish ;

tail not usually dark above M. nivalis, p. 713.

Anterior loop of first lower molar with postero-
external salient angle usually narrow and
sharply pointed, never obsolete (Alps and

Apennines) M. n. nivalis, p. 716.

Anterior loop of first lower molar with postero-
external salient angle usually broad and
rounded, sometimes obsolete (Pyrenees) M. n. aquitanim, p. 717.

MICROTUS NIVALIS Martins.

(Synonymy under subspecies.)

GeocjrapMcal distribution. — Pyrenees, Alps (except extreme
south-western portion : Basses- Alpes and Var), Apennines and
Tirol.

Diagnosis. — Size rather large (head and body about 130 mm.
in adults, hind foot, 20-22 mm., condylobasal length of skull,
28-30-4 mm.) ; tail about half as long as head and body ; colour
of upper parts in adults grey, strongly suffused with bister, but
never much clouded with black ; tail whitish, usually tinged with
brown above, though never sharply bicolor throughout ; general
characters of skull as in the sub-genus ; mesopterygoid fossa
narrow, its width anteriorly never more than equal to distance
between edge of fossa and alveolus of m^, its outer borders
vertical ; posterior border of palate w-ith median ridge usually
flattened and ill-defined, its width at least equal to that of
lateral pit.

External characters.— General appearance more murine than
in the other European species of Microtus, owing to the long
tail and large ears rather than to any special modification in
form of body. Head relatively longer and more pointed than

714

in M. agrestis. Ear rathei' large, appearing distinctly above
surrounding fur and extending nearly or quite to eye when laid
forward ; meatal lobe well developed, high, its margin evenly
rounded. Muzzle essentially as in 31. agrestis. Feet as in
M. agrestis, but posterior plantar tubercle larger than those at
base of digits. Tail about half as long as head and body, the
hairs not entirely concealing the ambulations, of which there are
about eighteen to the centimeter at middle ; pencil inconspicuous.
Fur full, soft and loose. Mamma? : p 2 — 2, « 2 — 2 = 8.

Colour. — -Upper parts smoke-grey, strongly washed with bister
on back, and usually tinged with pale buff along sides ; under-
parts dull white, irregularly clouded by the slaty under-colour ;
feet and tail whitish, the tail usually (in about two-thirds of the
skins examined) tinged with brown above though never sharply
bicolor.

SJciill. — The skull is larger than that of Microtus agrestis
agrestis, its general form broader and more depressed, the dorsal
sui'face much less marked by ridges for muscle attachment, and
the auditory bullte more developed. Dorsal iDrofile nearly flat from

lachrymal region to lambda, or
with a faint concavity in inter-
orbital region and equally slight
convexity over middle of brain-
case* ; nasals sloping forward at
an evident though slight angle
(about 15^). Occiput sufficiently
oblique for the condyles to be just
visible when skull is viewed from
above. Ventral profile with no
special peculiarities ; contrast be-
tween occipital depth and that
of rostrum evident but less pro-
nounced than in the European
members of the sub-genus Micro-
tus. Brain-case large relatively to
general size of skull, its outline
when viewed from above broadly
ovate, wider anteriorly than pos-
teriorly ; postorbital processes small and inconspicuous, producing
no very noticeable break in contour of brain- case ; ridges along
outer portion of parietals scarcely visible except in extreme old
age, and never becoming conspicvious ; lambdal crest represented
by extreme outer portions only, these much less developed than in
M. agrestis ; interparietal relatively smaller than in M. agrestis,
its area scarcely more than half that of parietal, its form not
peculiar (antero-posterior diameter, exclusive of median projection,
slightly less than half transverse diameter). Depth of occiput
when viewed from behind barely more than half height, the
* Somewhat exaggerated in fig. 148.

Fig. 148.
Microtus nivalis. Nat. size.

M1CR0TU9

715

brain-case scarcely rising above occipital level ; paroccipital pro-
cesses rather short, their tips projecting only a little beyond
level of lower lip of foramen magnum, the ridges extending
upward from their bases evident though not very high. Floor
of brain-case nearly smooth, the median ridge of basioccipital
slightly developed ; width of basioccipital along anterior suture
contained about 21 times in median length ; auditory bulke large
but somewhat flattened, otherwise without special peculiarities.
Interorbital region rather wide (usually wider than rostrum) and
short, the upper surface flattened and with low, inconspicuous
temporal ridges which may come together very late in life, but
which are too low to form an evident median ridge ; inter-
lachrymal region always essentially smooth. Zygomata not very
widely or abruptly spreading, the contrast between zygomatic
breadth and breadth of brain-case much less than in Microtus
agresfis ; median expansion evident but not very wide ; anteorbital
foramen rather wide and low, its form diagnostic as compared
with that of M. agrestis. Rostrum slender, relatively smaller
than in M. agrestis and with less contrast in depth between
proximal and distal regions, the least depth behind nasals
distinctly greater than width in same region ; nasals less narrowed
posteriorly than in M. agrestis, their hinder border usually
truncate and scarcely extending behind level of anterior border
of zygomatic root ; nasal branches of premaxillaries usually
extending about to level of middle of zygomatic root ; incisive
foramina about as in M. agrestis, but extending backward
scarcely to level of m^. Palate narrow, its structure normal, but
all the surface sculpture lacking in relief, the sloping portion of
median ridge wider than small, shallow lateral pit ; mesopterj-goid
space nearly parallel sided, its anterior border rounded, its width
in front scarcely equal to distance from margin of fossa to
alveolus of iti^ ; plates forming outer border of space nearly
vertical ; hamulars straight ; ectopterygoid plate well developed,
the ectopterygoid pit large and deep, though slightly more
encroached on than is the case in 31. agrestis. Mandible
essentially as in M. agrestis but more slender, coronoid process
lower, and angular process longer.

Teeth. — Incisors essentially as in Microtus agrestis but
relatively a little less robust ; upper teeth projecting slightly
forward, so that their front surface is just visible when skull is
viewed from above. Molars* relatively smaller than in the
European species of true 3Iicrotus, but the general character of
their enamel folding without special peculiarity other than a
slight tendency toward roundness in outline of closed triangles of
maxillary teeth, particularly the triangles of inner side. The
pattern in m^, iv? and m.^^ shows no peculiarities worthy of special
note ; m^ without postero-internal loop. Third lower molar with
re-entrant angles on oviter side tending to be deeper than usual,
* Fig. 149, p. 718.

716

RODENTIA

SO that the second loop is often nearly or completely divided into
two closed triangles. Third upper molar witli anterior loop and
three closed triangles of the usual form, but posterior loop short,
simple and without trace of re-entrant angle on inner side
(though in some instances a slight widening of the extreme tip
causes a certain degree of concaveness), its form much like that
assumed by extreme terminal portion of loop in M. agrestis when
second inner triangle has been cut oft'; the tooth never has more
than three well developed salient and two re-entrant angles on
each side. First lower molar essentially as in Microtus ratticeps,
the outer side with three, the inner side with four re-entrant
angles, the anterior loop short, simple, without trace of re-entrant
angles, its general outline crescentic (or when more elongate
almost arrow-head shaped) when outer basal portion is well
developed, or unsymmetrical when this is short or obsolete ; first
inner triangle rarely communicating with anterior loop ; outer
side normally with four salient and three re-entrant angles, inner
side with five salient and four re-entrant angles.

Bemarks. — Microtus nivalis is a strictly alpine animal, never
inhabiting plains or true lowlands and only descending into
valleys when special local conditions, such as damp, shaded cliffs
or cool talus-slopes,* furnish it with the low summer temperature
which it requires. Two local forms are distinguishable, one
confined to the Pyrenees, the other to the region of the Alps.

Microtus nivalis nivalis Martins.

1842. Arvicola nivalis IMartins, Revue Zoologique, p. 331 (Faulhorn, Bern,

Switzerland).

1843. HyjyiidieiLS alpinus Wagner, Schrebor's Siiugthiere, Sux^pl., iii, p. 57G

(Andermatt, Uri, Switzerland).
1845. Arvicola nivicola Schinz, Synops. Mamm., ii, p. 236 (Highest Swiss

Alps ; probably near Andermatt) .
1853. Hypuclivus iictropliilus Wagner, Miinch. Gelehrt. Anzeigen, No. 38,

p. 307, ]\Iarch 28, 1853 (Oberstdorf, near Southofen, Allgiiu,

Bavaria, Germany).
1857. Arvicola nivalis Blasius, Saugethiere Deutschlands, p. 359.
1884. \_Microtus'] nivalis Lataste, Actes Soc. Linn., Bordeaux, xxxviii,

p. 23 (p. 15 of reprint).
1908. Microtus nivalis nivalis Miller, Ann. and Mag. Nat. Hist., 8th ser.,

I, p. 99, January, 1908.
1910. Microtus (Chionomys) nivalis Troucssart, Faune Mamm. d'Europe,

p. 183.

Tijpe locality. — Faulhorn, Bern, Switzerland.

Geograpliical distribution. — Alps (except south-eastern portion),
Tirol and northei*n Apennines.

Diagnosis. — Anterior loop of first lower molar tending to
assume an arrow-head-like outline, owing partly to the general

* For a study of the local conditions in a typical talus-slope see Miller,
Science, N.S., viii, pp. 615-618, November 4, 1898.

MICROTUS 717

narrowness of the loop, but more especially to the form of the
postero-external salient angle, which is usually narrow and
sharjjly pointed, and rarely, if ever, except in immature
individuals, obsolete.

Measurements. — External measurements of adult male from
Chamonix, Haute-Savoie, France : head and body, 139 ; tail, 71 ;
hind foot, 20. Two adult males from the Furka Pass,
Switzerland: head and body, 127 and 133; tail, 63 and 68;
hind foot, 19 "6 and 19. Two adult females from the same
locality: head and body, 126 and 131; tail, 72 and 69; hind
foot, 20 and 20 ; ear, 15. Adult male from Santis, Appenzell,
Switzerland: head and body, 126; tail, 70; hind foot, 20;
ear, 15. Adult male from Valasco, Valle del Gesso, Cuneo,
Italy: head and body, 132; tail, 74; hind foot, 21; ear, 17.
Adult female from Monte Cimone, Modena, Italy : head and
body, 120; tail, 63; hind foot, 20. For cranial measurements
see Table, p. 720.

Specimens examined. — Sixty-two, from the following localities : —

Feance: Chamonix, Hautc-Savoie, 2 (U.S.N.M.).

Switzerland : Sangloz, Vaud, 2 (U.S.N.M.) ; Eusannaz, Vaud, 2
(U.S.N.M.) ; Zermatt, Valais, 3 (U.S.N.M.) ; Miirren, Berne, 2 ; Meiringen
Berne, 2 ; Purka Pass, Uri, 11 (B.M. and U.S.N.M.) ; St. Gothard, Uri, 4
Andermatt, Uri, 5 (U.S.N.M.) ; Goschenen, Uri, 6 (U.S.N.M. and Mottaz)
Santis, Appenzell, 12 (U.S.N.M.); Meglis Alp, Appenzell, 2 (U.S.N.M.).
Hochmiittli, Murgsee region, St. Galleu, 2 (U.S.N.M.) ; Campfer, Grisons, 1
(Rothschild) ; no exact locality, 2.

Germany : Oberstdorf, near Sonthofen, Allgau, Bavaria, 2.

Italy : Valasco, VaUe del Gesso, Cuneo, 1 (Genoa) ; Monte Cimone,
Modena, 1.

2. Miirren, Berne, Switzerland. W. Gurtner (c & p). 92. 10. 5. 10-11.

2. Meiringen, Berne. Tomes Collection. 7. 1. 1. 143-144.

7 6, 9. Furka Pass, Uri. {E. H. O. Thomas (p). 2. 8. 4. 4:3-47.

ZolUkofer.) 4. 4. 5. 54-5G.

1. St. Gothard, Uri. Baron E. de Sulys- 45. 7. 5. 3.

Longchamps (p).

2. St. Gothard, Uri. Purchased (Brandt). 45. 11. 1. 6-7.
1. St. Gothard, Uri. Stockholm Museum 46. 6. 2. 67.

(E).

6. Switzerland. (Moscldcr.) E. R. Alston (p). 79. 9. 25. 50.

1. Switzerland. Purchased (Parreys). 46.6.15.58.

? al. Monte Cimone, Modena, Modena Museum (p). 3. 2. 22. 1.
Italy. {Dr. Major.)

MiCROTUS NIVALIS AQUITANIUS Miller.

1908. Micwlus nivalis aquitanius Miller, Ann. and Mag. Nat. Hist.,
8th sor., I, p. 99, January, 1908. Type in British Museum.

1910. Microtus [Chionomys) nivalis aquitanius Trouessart, Faunc Mamm.
d'Europe, p. 184.

Type locality. — Near I'Hospitalet, Ariege, France. Altitude
about 4800 feet.

Geographical distribution. — Pyrenees ; at present known from
the French side only.

718

RODENTIA

Diagnosis. — Similar to Microtus nivalis nivalis, but anterior
loop of first lower molar broadly crescentic in outline, the postero-
external salient angle broad and rounded, occasionally obsolete.

Enamel pattern of Microtus nivalis nivalis (a) and
il. nivalis aquitanius (b). X 5.

Measurements. — Type (young-adult male) : head and body. 111;
tail, 59 ; hind foot, 21 ; ear, 16. Two adult males from the type
locality: head and body, 124 and 124; tail, 61 and 64; hind
foot, 20 • 4 and 21 ; ear, 17*8 and 19 • 4. Adult male from Pic-du-
Midi, Hautes-Pyrenees : head and body, 129 ; tail, 71 ; hind
foot, 21 ; ear, 17. For cranial measurements see Table, p. 721.

Specimens examined. — Fourteen, from the following localities on the
French side of the Pyrenees : Porte, Pyr6n6es-Orientales, 2 ; I'Hospitalet,
Ariege, 5 ; Bareges, Hautes-Pyrenees, 1 ; Pic-du-Midi, Hautes-Pyrenees, 6.

2?.

3 6.

Port6, Pyr^n6es - Orientales,

France.
L'Hospitalet, Ariege.

3 (5, 3 9. Pic-du-Midi, Hautes-Pyrenees.
(.4. Robert.)

G. S. Miller (c). 8. 8. 4. 240-241.

G. S. Miller (c). 8. 8. 4. 237-239.
(8. 8. 4. 238. Tyiie of subspecies.)
0. Thomas (p). 8. 9. 1. 64-69.

MICROTUS LEBRUNII Crespon.

(Synonymy under subspecies.)

Geographical distribution. — Central and south-eastern France
from Puy-de-D6me to the Departments of Gard, Var, and Basses-
Alpes.

Diagnosis. — Like Microtus nivalis, but smaller (hind foot in
adults less than 20 mm. ; condylobasal length of skull less than
28 mm.), and paler, the tail always pure white throughout, and
the back a light grey without conspicuous brownish suffusion.

Skull. — Except for its smaller size, the skull does not difler
tangibly from that of M. nivalis.

Teeth. — As compared with those of M. nivalis of the same
age, the teeth differ in their slightly smaller size, but I can find

MICROTUS 719

no constant character in the enamel pattern by which they may
be distinguished. In the specimens examined the terminal loop
of vP is invariably short and broad, not showing any tendency to
the elongation occasionally occurring in 31. nivalis.

Bemarks. — Microtus lebrunii is represented by two geographic
races, one of which is remarkable, in the sub-genus Cliionomys, for
its occurrence at a distance from mountains and in a region of
unusually high summer temperature.

Microtus lebrunii lebkunii Crespon.

1844. Alr-vicolci] lebrunii Crespon, Faune M^ridionalc, i, p. 77.

1857. Arvicola nivalis b. Arvicola leucurus Blasius, Siiugethicre Deutsch-

lands, p. 359 (part).
1908. Microtus lebrunii lebrunii Miller, Ann. and Mag. Nat. Hist., 8th ser.,

I, p. 101, January, 1908.
1910. Microtus {Cliionomys) lebruni Trouessart, Faune IMamm. d'Europe,

p. 184.

Type locality. — Neighbourhood of Nimes, Gard, France.
Altitude, 180 m.

Geographical distribution. — ^Known with certainty from the
type locality only, but probably extending through the Ceveinies
and the mountains of Auvergne.

Diagnosis. — Back clear, very pale smoke-grey, without evident
wash of wood-brown ; skull with auditory bullae small and
flattened.

Colour. — Upper parts a very pale smoke-grey with a faint
bluish tinge, particularly on neck and between ears, the back
faintly " lined " with black, the sides and rump nearly clear ;
underparts buffy white, not sharply defined, the slate-black under
colour appearing irregularly at surface and producing a bluish
tinge ; sides of muzzle dull white ; feet and tail white
throughout.

Skull. — Auditory bullse decidedly reduced in size as compared
with those of M. nivalis, so that the lower edge of bulla scarcely
extends beyond line of cutting edge of molars.

Measurements. — Three adult males from the type locality :
head and body, 117, 121 and 122; tail, 56, 55 and 66; hind
foot, 18-6, 19-6 and 18-8 ; ear, 13, 14-2 and 14. For cranial
measurements see Table, p. 721.

Specimens examined. — Six, all from the neighbourhood of Nimcs (B.M.,
Mottaz and Nimes) ; no exact locality, 2.

Remarks. — This animal presents the anomaly of a member of
the nivalis group completely adapted to life in the hot dry plains
of south-central France, a region where its presence can in no
way be explained as due to special local conditions. A mounted
specimen in the Paris Museum collected in Auvergne (probably
near Clermont-Ferrand) by Lecoq, and presented in 1854, is

720

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722

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probably referable to Microhis lehrunii lehrunii, though in its
present faded and dust-stained condition exact identification is
impossible. The skull is so imperfect that it serves only to show
that the animal was not adult.

3 6. Nimes, Gard, France. O. Thomas (p). ' 8. 8. 10. 101-103.

(C. Mottaz.) {Topotyjycs.)

2. S. France. Tomes Collection. 7. 1. 1. 14G-147.

MiCROTUS LEBRUNii LEUcuRus Gerbe.

1852. Arvicola leucurus Gerbe, Eevue Zoologique, 2nd ser., iv, p. 2G0,

June, 1852.
1857. Arvicola nivalis b. Arvicola leucurus Blasius, Saugethiere Deutscb-

lands, p. 359 (part).
1908. Microtus lehrunii leuciirus Miller, Ann. and Mag. Nat. Hist., 8tb ser.,

I, p. 102, January, 1908.
1910. Microtus [Chionomys) lebruni leucurus Trouessart, Faune Mamm.

d'Europe, p. 184.

Ty2Je locality. — Barcelonnette, Basses-Alpes, France. Altitude
about 1300 mm.

Geographical distribution. — South-western Alps (Departments
of Basses-Alpes and Var).

Diagnosis. — Upper parts pale smoke-grey, slightly but evi-
dently washed with wood-brown ; skull with auditory bullae large
and well inflated, essentially as in M. nivalis.

Colour. — The colour resembles that of M. lehrunii lehrunii,
except that the back is sufiiciently suffused with bufFy or pale
wood-brown to destroy the clear grey, almost hoary appearance
characteristic of the typical, lowland race. This sufiusion is
usually more noticeable on rump than on sides, so much so that
in one specimen (No. 7. 1. 1. 145, probably a paratype) the
rump is a clear dull buff.

Skull. — The skull is distinguishable from that of 31. lehrunii
lehrunii by its large, strongly inflated, normally developed auditory
bull*. As in M. nivalis the lower edge of the bulla extends
decidedly beyond line of cutting edge of molars.

3Ieasuremcnts. — Adult female from St. Paul, near Barce-
lonnette, Basses-Alpes, France: head and body, 120; tail, 68;
hind foot, 19 ; ear, 14" 8. For cranial measurements see Table,
p. 721.

Specimens examined. — Twelve, all from the vicinity of Barcelonnette
(B.M, and Mottaz) ; also several old specimens without definite locality,
probably referable to the same race (B.M. and U.S.N.M.).

(J, 9. Barcelonnette, Basses-Alpes, O. Thomas (p.) 8. 8. 10. 104-105

France. (C. Mottaz.) (Topotypes.)

3. Basses-Alpes. ■ Purchased (Parzu- 52. 5. 27. 47-49.

daki).
1. S. France. Tomes Collection. 7. 1. 1. 145.

ARVICOLA

723

MICROTUS ULPIUS Miller.

1908. Microtiis ulpius Miller, Ann. and Mag. Nat. Hist., Sth ser., i, p. 100,

January, 1908. Type in British Museum.
1910. Microtns (CJdonomys) ulpius Trouessart, Fauue Mamm. d'Europe,

p. 184.

Type locality. — Hatszeg, Hunyad, Austria-Hungary. Altitude,
2000 feet.

Geographical clistrihuiion. — Known only from the vicinity
of the type locality, but probably occurring throughout the
Transylvanian Alps and perhaps in the true Carpathians
also.

Diagnosis. — Similar to Microfus nivalis, but colour darker and
tail usually brown above (often distinctly bicolor throughout) ;
posterior border of palate with median ridge sharply defined, its
width less than that of deep lateral pit.

Colour. — Though in general resembling that of Microtus nivalis
the colour is distinctly darker, the back strongly tinged with
wood-brown and conspicuously clouded with black ; tail usually
bicolor, often sharply and conspicuously so throughout.

Skull and teeth. — Except for the narrow, sharply defined
median ridge and deep lateral pits the palate is as in M. nivalis.
The other parts of the skull show no peculiarities. Teeth as in
Microtus nivalis, the anterior loop of ni^ usually similar to that of
M. n. aquitanius.

Measurements. — External measui-ements of type (adult female) :
head and body, 131 ; tail, 58 ; hind foot, 20 ; ear, 17. A second
adult female from the type locality : head and body, 124 ; tail, 68 ;
hind foot, 22 ; ear, 17. For cranial measurements see Table,
p. 721.

Specimens examined. — Eighteen, all from the neighbourhood of Hatszeg.

G. Danford (c). 3. 2. 2. 45-4S
3. 11. 8. 35-4
(8. 2. 2. 48. TyjK of species.)

6 6,19. Hatszeg, Hunyad, Transyl- C. G. Danford (c). 3. 2. 2. 45-49.
vauia, Austria-Hungary. 3. 11. 8. 35-42

Genus ARVICOLA Lacepede.

1799. Arvicola Lac6pede, Tableau des divisions, sous divisions, ordres et

genres des mammiferes, p. 10 [amijhibius).
1836. Hemiotomys de S61ys-Longchamps, Essai monographique sur les

Campagnols des environs de Li6ge, p. 7 (sub-genus) part.
1857. Paludicola Blasius, Saugethiere Deutsehlands, p. 333 (sub-genus)

part. Not Paludicola Wagler, 1830.
1867. Ochetomys, Fitzinger, Sitzungsb. Math. Naturwiss. CI. K. Akad.
Wissensch., Wien, lyi, p. 47 (based primarily on the water-rats of

Europe).
1867. Praticola Fatio, Les Campagnols du Bassin du Leman, p. 36

(sub-genus) part. Not Praticola Swainson, 1837.
1883. Arvicola Lataste, Le Naturaliste, ii, p. 349 (sub-genus).

3 A 2

724 RODENTIA

189G. Arvicola Miller, North Amer. Fauna, no. 12, p. GO, July 23, 189G

(sub-genus).
1908. Arvicola Miller, Ann. and Mag. Nat. Hist. 8tli ser., i., p. 195,

February, 1908 (genus).

Tt/2)e species. — 3Ihs ampliihius Linntwus.

Geographical distribution. — Paljearctic region from Great
Britain eastward into Asia, and fi'om the Meditei-ranean coast
north to northern Scotland and the shores of the Arctic Ocean.

Characters. — Like Microtus, but plantar tubercles only five
(a very rudimentary sixth rarel}'^ present) ; third upper molar as
in Chionomys ; external form modified for aquatic or subterranean
life.

Remarks. — Though not sharply differentiated from 3Iicrotus,
this group is so natural that it may conveniently be treated as a
genus. Its range does not extend beyond the Pala?arctic region.
About fifteen forms are known, eleven of which occur in western
Europe. Some of these are strictly aqviatic in their mode of life,
others fossorial, while in two {A. terresiris and A. scherman
scherman) the habits appear to vai-y locally or with season.

KEY TO THE EUROPEAN FORMS OF ARVICOLA.

Size large, hind foot more than 30 mm., coudylobasal

length of fully adult skulls usually more than

40 mm. ; skull not fossorial in form, the rostrum

and occiput tending to be squarely truncate, the

upper incisors slightly projecting; roots of nii

and vi.;, forming evident protuberances on lower

surface of mandible in old individuals ; habits

strictly aquatic [ampliihius group).
Nasals at widest region conspicuously narrower

than rostrum (Great Britain) A. ampliihiiis, p. 725.

Condylobasal length of skull usually 42 to
44 mm. ; colour dark, but melanism infre-
quent (England) A. a. awphihius,T^.l?>0.

Coudylobasal length of skull usually 40 to
42 mm. ; colour very dark, melanism frequent

(Scotland) A. a. reta, p. 732.

Nasals at widest region nearly as wide as rostrum

(Spain and southern France) A. sapidus, p. 732.

General colour rather light, the sides and face
clear yellowish brown (Spain, except As-
turias; lowlands of southern France) A. s. saiJichis, p. 733.

General colour dark (as in amphibins), the sides
and face strongly " lined " with black

(Pyrenees and Asturias) A.s. tenehricus, p. 735.

Size medium or small, hind foot usually less than

30 mm., condylobasal length of fully adult

skulls usually less than 40 mm. ; skull slightly

or conspicuously fossorial in form, the rostrum

and occiput tending to be obliquely truncate, the

upper incisors noticeably projecting ; roots of

«i, and vi„ never forming protuberances on lower

surface of mandible ; habits aquatic or terres-
trial [terresiris group).

AUVICOLA 725

Interparietal teudiug to bo somewhat ligulate in
outline ; skull representing the extreme of

fossorial adaptation A. scherman, p. 744.

Colour dark brown, the tail usually l)lackish

throughout ; habits both aquatic and fossorial

(Northern central Europe from southern

' Germany to the Baltic and from Belgium

eastward) A. s. scherman, p, 745.

Colour light yellowish brown, the tail usually

bicolor or buffy; habits mole-like, strictly

fossorial.

Anterior loop of m^ short and wide ; auditory

bullae usually somewhat flattened (Alps

and their immediate neighbourhood) A. s. cxUus, p. 746.

Anterior loop of »«, long and narrow ; auditory
bullae evenly inflated (Pyrenees and their

immediate neighbourhood) A.s. monticola, p. 749.

Interparietal tending to be quadrate in outline ;
skull less extremely fossorial in form.
Molars relatively weak ; underparts with con-
spicuous rusty wash which usually spreads
noticeably on sides of head (Norway and

Sweden) A. tcrrcstris, p. 738.

Molars relatively heavy ; underparts with at

most an ochraceous wash which rarely

spreads noticeably on sides of head.

General colour strongly yellowish, the back

with little if any clouding of black (Central

Italy) A. 7)msig)iani,]). 744.

General colour dark brown, the back with
noticeable clouding of black.
Underparts slaty, washed with yellowish
brown (Italian Switzerland and north-
ern Italy) A. italicus, p. 740.

Underparts slaty, washed with whitish

(Bosnia and Roumania) A. illyricus, p. 741.

ARVICOLA AMPHIBIUS Linnseus.

(Synonymy under subspecies.)

Geo(jra^)hiral distribution. — Great Biutain.

Diagnosis. — Size large (head and body about 200 mm., tail
about 110 mm., hind foot usually 30 to 34 mm., condylobasal
length of fully adult skulls, 40 to 44*6 mm.) ; tail somewhat
more than half as long as head and body ; colour above dark
brown, blackening along back, the sides not decidedly yellowish,
the cheeks not contrasted with surrounding parts ; skull not
fossorial in form, the rostrum and occiput tending to be squarely
truncate (vertically), the upper incisors not unusually pro-
jecting ; nasals at widest region conspicuously narrower than
rostrum ; roots of wt^ and m., forming evident pi'otuberances on
lower surface of mandible in old individuals ; habits strictly
aquatic, never mole-like.

External characters. — Genei'al form heavier and more robust
than in the house rat, the tail relatively shorter and head

726 EODENTIA

noticeably more rounded. Upper incisors slightly projecting
when mouth is closed. Ear well developed but low, scarcely
appearing above sui'face of fur, and extending barely more than
half-way to eye when laid forward, its outline sub-circular ;
meatal valve well developed, bluntly triangular, naked ; both
surfaces of ear thinly clothed with rather long hairs. Eye small,
its position half-way between muzzle and ear. Muzzle pad small,
naked, with no special peculiarities, its lower border continuous
with naked median groove of upper lip. Front foot broad and
robust, though not specially enlarged ; inner digit rudimentary,
smaller than the smallest palmar tubercle, its dorsal surface
covered by the closely appressed nail ; fifth extending to base of
third and fourth, second about to middle of third ; third and
fourth digits sub-equal and longest, the third slightly exceeding
fourth ; scales on under surface of digits well defined, three to
five in number ; claws well developed, slender, slightly curved,
their length al:)Out equal to that of first phalanx of the respective
digits. Palmar tubercles five, all well developed, the two posterior
sub-equal and largest, that lying between bases of third and
fourth digits smallest ; surface of palm between tubercles
irregularly wrinkled. Hind foot large, slightly but evidently
" feathered " for swimming ; first digit extending scarcely to
base of second, fifth to just beyond middle of fourth ; second,
third and fourth digits sub-equal, the third slightly the longest
and second slightly the shortest of the three ; scales on under
surface of digits well defined, three to four number ; claws slightly
larger and more robust than those of fore foot. Sole naked
throughout, except for a sprinkling of fine hairs on posterior
third which usually produces a slight pubescence. Plantar
tubercles five, welt developed, fully as large as pads at base of
corresponding claws, occupying about one-half surface of region
in which they lie, the skin between them finely granular like
that of posterior portion of sole except for a noticeable smooth
area indicating position of sixth tubercle ; outline of tubercles at
base of first and fifth digits sub-circular, that of the others
naiTowly ovate ; postero-internal tubercle slightly larger than
any of the others, that at base of first digit smallest. Tail thick
at base, tapering noticeably toward tip ; when laid forward it
extends about to shoulders or slightly beyond ; annulations
evident, though somewhat irregular, about 15 to the centimeter
at middle of tail ; hairs numerous, about 4 to 5 mm. in length,
nearly concealing the annulations and forming a slight pencil.
Mamma? : p 2 - 2, i 2 - 2 = 8.

Colour. — Upper parts ranging from broccoli-brown to mars-
brown or darker, in most specimens rather hea\'ily overlaid wdth
black, though the latter is usually not in excess of the ground-
colour ; median dorsal region darkest, sides and face lightest,
though without noticeable contrast, the sides of body usually
somewhat " lined " with black, the sides of head to, and including

ARVICOLA 727

ears, usually tinged with odiraceous-])uff or light raw-umber,
though not sufficiently to produce any marked contrast with
surrounding parts ; chest and belly a varying mixture of
ochraceous-l)utf and the slate-grey under colour, the two
becoming more intimately blended on throat and fading to a
light drab grey ; feet ranging from hair-brown to ecru-drab,
sometimes with a blackish shade ; tail blackish throughout, the
underside sprinkled with greyish hairs.

Skull. — The skull of Arvicola ampJiibius is lai'ge and massive,
becoming conspicuously ridged and angular in fully adult

Fig. 150.
Arvicola amphibius. Nat. size.

individuals. In general form and proportions it does not differ
notably from the skull of Microtus agrestis, except that the brain-
case tends to be shorter and wider, the dorsal profile is more
evenly convex, without any sudden angular bending down of
nasals at interlachrymal region, and the upper incisors project
sufficiently forward so that most of their anterior surface is
visible when skull is viewed from above. Brain-case somewhat
longer than wide, the well-developed postorbital processes
squaring its anterior margin and imparting the general outline
of a parallelogram ; lateral ridges strongly developed in adults
and marking ofT a conspicuous flat median surface widest at

728 RODENTIA

middle, narrowing a little posteriorly to width of interparietal,
and more noticeably in front, finally disappearing in region
between postorbital processes, where the ridges come together ;
each ridge with a distinct angle at middle and usually with
another less evident angle about half-way between middle and
point of juncture ; interparietal large, somewhat variable in
form, usually ligulate with obliquely truncate outer extremities
and slight anterior median projection, but sometimes almost a
perfect parallelogram about twice as wide as long ; lambdoid
crest high and conspicuous at sides, but reduced to a mere
angular edge along posterior border of interparietal ; truncation
of occiput somewhat oblique, slightly more so than in MIcrotus
agrestis, the entire occipital condyle usually, though not always,
visible when skull is viewed from above ; outline of occiput
viewed from behind elliptical, flattened at the middle, slightly
more than half as high as wide ; j^aroccipital processes large,
widely spreading, their extremities not applied to bulla?, their
bases continued upward as well-developed ridges joining lamb-
doid crest near outer extremities of interparietal ; foramen
magnum higher than wide, its upper border narrowly rounded or
forming a distinctly pointed arch ; base of brain-case with no
special features, the basioccipital with ill-defined median longi-
tudinal ridge, the width of the bone along anterior border
contained about 2^ times in its median length ; auditory bulla?
rather large (about 9 mm. in antero-posterior diameter), hnt not
highly inflated, their ventral border not extending to level of
cutting surface of molars. Interorbital region deeply constricted
posteriorly, where in old individuals the width is less than half
that in lachrymal region ; ridges early uniting to form an
evident median crest which, however, does not extend forward
beyond the deeply constricted region. Zygomata heavy, rather
gradually spreading, the greatest zygomatic breath at glenoid
level ; vertical expansion evident but not abrupt. Rostrum of
the same relative size as in MIcrotus agrestis, but somewhat more
slender anteriorly ; nasals distinctly smaller relatively to rostrum
than in M. agrestis, their anterior termination falling decidedly
short of level of gnathion, their greatest combined breadth
noticeably less than that of rostrum in same region ; posterior
border of nasals squarely truncate at level of about middle of
zygomatic root, ascending branches of premaxillaries reaching
somewhat further back ; incisive foramina relatively shorter
than in Microtus agrestis, their posterior border falling short of
alveolar level by 1 mm. or more, their anterior extremity lying
decidedly nearer maxillo-premaxillary suture than to posterior
surface of incisor. Bony palate with no special peculiarities as
compared with that of Microtus agrestis ; lateral pits large, but
rather shallow ; median ridge tending to be low and not sharply
defined ; mesopterygoid space with anterior border varying from
squarely truncate to pointedly arched ; posterior half of ptery-

729

golds noticeably diverging ; ectopterygold pit large and deep, not
encroached on by bulla. The mandible does not differ materially
from that of Microtus agrestis except in its conspicuously greater
size, but the articular process is more abruptly bent inward at
level of incisor root, and angular process is more widened
posteriorly.

Teeth. — Upper incisor with shaft less curved than that of
Microtus agrestis, its outline from tip to base less nearly a semi-
circle ; root at anterior margin of anteorbital foramen, where it
forms a slight though evident swelling. Cross-section of shaft
essentially as in M. agrestis, but antero-posterior diameter
relatively greater, so that inner border is slightly longer than
anterior border ; distribution of enamel as in the smaller animal.
Lower incisor with root extending well into base of articular
process, on the outer side of which it
usually forms a slight protuberance ; cross
section, like that of upper tooth, with
antero-posterior diameter relatively greater
than in M. agrestis, the enamel-covered
anterior surface thus relatively narrower.
Molars large and heavily built ; m.^ form-
ing a conspicuous capsule on inner side
of incisor, the roots of m^ and m., appear-
ing as distinct protuberances on lower
surface of mandible. Enamel pattern
well defined, all the elements distinct, the
angles sharp and definite. Details of
pattern much as in Microtus nivalis ; m^
with anterior transverse loop and four
alternating, essentially equal closed tri-
angles, each side with three salient and
two re-entrant angles ; m'^ with anterior
transverse loop and three alternating closed

triangles, the outer side with three salient and two re-entrant
angles, the inner side with two salient and two re-entrant
angles ; m^ with anterior transverse loop, a larger inner and
smaller outer closed triangle, and a short longitudinal loop,
from the antero-external base of which a small third closed
triangle is sometimes cut off; each side of tooth with three salient
and three re-entrant angles, the last two salients occasionally ill-
developed in individuals with reduced terminal loop ; lUi with
same elements as in Microtus nivalis, but first and second
triangles normally opening into each other and into short, sub-
terete anterior loop, of which in some specimens they appear to
be mere angular basal appendages ; true closed triangles only
three, one outer and two inner ; outer side of tooth with four
salient and three re-entrant angles, inner side with four and a
rudimentary fifth (anterior) salient and four re-entrant angles ;
m^ with four closed, sub-equal triangles (the antero-internal and

Arvicola amphihius.
Enamel pattern. X 5.

730

RODENTIA

antero-external often communicating) and a posterior transverse
loop ; each side with three salient and three re-entrant angles ;
7H.3 with the same elements as m., but smaller and less well-
defined, both first and second pair of triangles tending to open
into each other, though the two pairs are normally separated.

Hemarlcs. — So far as is at j^resent known Arvicola amjMbius
is confined to Great Britain. Among the continental species it
finds its nearest relative in A. sajiidus of Spain and southern
France. Two local races appear to be represented among the
specimens examined, though their status is not well defined.

AiivicoLA AMPiiiBlus AMPHiBius Linmeus.

1758. [Mus] ampliibius Liunseus, Syst. Nat., i, lOfch ed., p. 61 (England :
based on the Mus 7najor aqiiaticus of Ray, Quadr., p. 217).

1816. '} Arvicola aquatica Leach, Syst. Catal. Spec. Indig. ]\Iamm. and

Birds, Brit. Mus., p. 7 (nomen nudum : " Water Camx^agnol ").

1817. Lcnimus aquaticus F. Cuvier, Diet, des Sci. Nat., vi, p. 306, part

(substitute for amithibius).
1842. Arvicola amcricana Gray, Ann. and Mag. Nat. Hist., x, p. 226 (half-
grown individuals supposed to have been taken in South America).

Go-types in British Museum.
1845. ? [Arvicola amphihiics] sub-var. nigricans de S61ys-Longchamps, Atti

della sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 322

(nomen nudum).
1857. Arvicola amphibius a. Blasius, Siiugethiere Deutschlands, p. 344

(part).
1910. Arvicola ampliibius avivliibizis Miller, Proc. Biol. Soc. Washington,

XXIII, p. 19, March 23, 1910.
1910. Arvicola amphibius Trouessart, Faune Mamm. d' Europe, p. ix.

Tyjje locality. — England.

Gco<jrap1iical distribution. — England and southern Scotland ;
exact northern limits of range not known.

Diagnosis. — Size maximum for the species (hind foot in
adults usually 32 to 35 mm., condylobasal length of skull, 42 mm.
or more) ; colour moderately dark, the black rarely in excess of
brown on upper parts ; melanism infrequent.

Measurements. — Adult male from Diss, Norfolk : head and
body, 206 ; tail, 139 ; hind foot, 35. Adult female from Cheadle,
Staffordshire: head and body, 185; tail, 118; hind foot, 33.
Adult female from Shalford, Surrey : head and body, 201 ;
tail, 117; hind foot, 33; ear, 17. Adult male from Garrett
Park Lake, Eaidsfield, Surrey: head and body, 193; tail, 128;
hind foot, 35 ; ear, 16. Adult male from Minster, I. of Thanet,
Kent: head and body, 168; tail, 120; hind foot, 32; ear, 18.
Adult from Horsham, Sussex : head and body, 186 ; tail, 130 ;
hind foot, 33. For cranial measurements see Table, p. 736.

Specimens examined. — Seventy-two, from the following localities : —
Scotland : Blackwood, Lanarkshire, 3 (Edinburgh) ; Stockbriggs,
Lanarkshire, 1 ; Bargsly, Kirkcudbright, 1 ; Stirling, 1.

ARVICOLA

731

England : Alnwick, Northumberland, 1 ; Wylam, Northumberland, 1 ;
Hull, Yorkshire, 4; Cheadle, Staffordshire, 1; Diss, Norfolk,!; Ashton,
Oundle, Northamptonshire, 2 ; Waterbeach, Cambridgeshire, 1 ; Grafton-
bury, Hereford, 7; Great Grimsby, Lincolnshire, 2; Kingsbury, Middlesex,
1 ; Hampton, ]\Iiddlesex, 1 ; Chertsey, Surrey, 1 ; Garrett Park Lake,
Earlsfield, Surrey, 6 ; Shalford, Surrey, 9 ; Surbiton, Surrey, 1 ; New
Forest, Hampshire, 9 ; Eversley, Hampshire, 3 ; Aldershot, Hampshire, 3 ;
Clevedon, Somerset, 2 ; Blandford, Dorsetshire, 2 ; Cullompton, Devon-
shire, 2; Northlew, Devonshire, 1; Horsham, Sussex, 1; Minster, I. of
Thanet, Kent, 1 ; Yalding, Kent, 1 ; no exact locality (supposed to be from
South America), 2 (co-types of americana Gray).

9. Stockbriggs, Lanark-
shire, Scotland.

(Taylor.)
9. Stirling. [Broion.)
6. Alnwick, Northumber-
land, England.
6. Wylam, Northumber-
land.
2 (5, 2 ?. Hull, Yorkshire. [Gapt.
Hmne.)
Oundle, Northampton-
shire. [Cox.)
Graftonbury, Hereford-
shire.
Great Grimsby, Lincoln-
shire.
Kingsbury, Middlesex.

{J. E. Harting.)
Hampton, Middlesex.

(W. Dodson.)
Chertsey, Surrey.

2 6.

3 6.
6,9.

6.
6.
?.
6.
3 6,69

Earlsfield, Surrey.
Shalford, Surrey.

1 (albino) Surbiton, Surrey.

6, 9 juv., Aldershot, Hampshire.

<5al.

i 6, 5 9. New Forest, Hampshire.

E. R. Alston (p).

B. R. Alston (p).
W. E. de Winton

(c & p).
W. Burden (p).

G. Barrett-Hamilton

Hon. N. C. Roths-
child (p).

W. E. de Winton
(c & p).

G. H. Caton Haigli
(c & p).

E. R.Alston (c&p).

G. Barrett-Hamilton

Gordon Dagleish
(c & p).

C. H. B. Grant
(c & p).

W. R. Ogilvie-Grant

(c & p).
H. Buller (c & p).
R. I. Pocock (c & p).

Miller Collection.

2 c5, 1 9. Eversley, Hampshire. Miller Collection.

Clevedon, Somerset.
Blandford, Dorset.

Cullompton, Devonshire.

R. I. Pocock (c & p).

J. C. Mansel Pley-
dell (c&p).

G. C. Shortridgo
(c & p).

Purchased (War-
wick). [Co-types
Gray).

79. 9, 25. 46.

79. 9. 25. 49.
11. 1. 3. 406.

11. 1. 3. 294.

■ 11. 1. 2. 54-57.

6. 8. 18. 1-2.

11. 1. 3. 297-299.

11. 1. 8. 295-296.

79. 9. 25. 45.

11. 1. 2. 58.

4. 10. 13. 1.

11. 1. 3. 293.

6. 9. 20. 1-9.

I. 1. 14. 1.
88. 7. 14. 1-3.

7. 7. 7. 2925-
2929, 2931 -
2934.

7. 7. 7. 2930,
2935-293C.

II. 1. 3. 291.
11. 1. 3. 292.

11. 1. 3. 290.

42. 4. 12. 6-7.
of A. americana

732

Arvicola amphihius hkta Miller.

1832. Arvicola ater Macgillivray, Mem. Wenierian Soc. Nat. Hist, vi,
p. 429. Not Hypudmis terrestris ^ atcr Billberg, 1827.

1910. Arvicola amphihi^is reta IMiller, Proc. Biol. Soc. Washington, xxiii,
p. 19, March 23, 1910 (substitute for ater).

1910. Arvicola ampliibius rcta Trouessart, Faunc Mamm. d'Europo, p. x.

Tjipc locality. — Aberdeen, Scotland.

Geographical distribution. — Scotland, except southern portion ;
limits of distribution not known.

Diagnosis. — Size less than in A. amphihius amphihius (hind
foot usually 30 to 32 mm., condylobasal length of skull usually
less than 42 mm.) ; normal colour darker than in the typical
race, the black usually in excess on upper parts ; melanism
frequent.

Measurements. — Adult male from Glenfeshire Forest, Inverness :
head and body, 190; tail, 125; hind foot, 32; ear, 16. Adult
female from Windygates, Fife: head and body, 194-5 ; tail, 109 ;
hind foot, 30-5; ear, 14*5. For cranial measurements see
Table, p. 736.

Specimens examined. — Twelve, from the following localities in Scot-
land : — Elgin, 1 ; Monar Forest, Ross, 1 ; Crieff, Perthshire, 1 ; Glenfeshire
Forest, Inverness, 1 ; Cortachy, Forfar, 3 (B.M. and Wilson) ; Windygates,
Fife, 5 (B.M. and Edinburgh).

6.

Llanbryde, Elginshire.
Scotland.

W. Taylor (c & p).

11. 1. 3. 407.

6.

Monar Forest, Ross-

Maj.-Gen. C. E. Luard

92. 8. 31. 1.

shire.

(c & P).

6.

Kingussie, Inverness-
shire.

G. A. Cooper (c. & p).

5. 5. 12. 1.

9 al.

Crieff, Perthshire.

W. R. Oglivie-Grant
(c & p).

88. 5. 30. 2.

S, 9.

Windygates, Fife-
shire.

N. B. Kinuear (c & p).

6. 11. 18. 5-6.

c?, ?.

Cortachy, Forfarshire.

E. A. Wilson (c & p).

11. 1. 3. 408-409.

ARVICOLA SAPIDUS Miller.
(Synonymy under subspecies.)

Geographical distribution. — Iberian Peninsula and southern
France east nearly to the Italian border ; northern limit of
range not known.*

Diagnosis. — Like Arvicola amphibius in general size and in
form of skull ; nasal bones much widened anteriorly, their
greatest combined breadth nearly equal to that of rostrum ;
habits strictly aquatic, never mole-like.

Bemarlcs. — This species appears to be more nearly related to

* An animal of this type occurs at least as far north as Paris (Lataste,
Actes Soc. Linn. Bordeaux, xxxviii, p. 37, 1884), but I have had no
opportunity to examine specimens.

ARVICOLA 733

the large Arvicola ampldhius of Great Britain than to any of the
geographically less distant members of the genus. At present,
however, no specimens have been examined from regions in which
intergradation with true Arvicola scherman might be expected to

Fig. 152.
Arvicola sapidus. Nat. size.

take place. In the Pyrenees and their immediate neighbourhood
a form of sajndus occurs in the same region as a strictly terrestrial
form of scherman, but the two animals are completely segregated
by their different modes of life. Two races of Arvicola sapidus
are now known.

Arvicola sapidus sapidus Miller.

1908. Arvicola sapidus Miller, Ann. and Mag. Nat. Hist., 8th ser., i, p. 195,

February, 1908. Type in British Museum.
1910. Arvicola sapidus sapidus Miller, Proc. Biol. Soc. Washington, xxiii,

p. 20, March 23, 1910.
1910. Arvicola sapidus Trouessarfc, Faune Mamm. d'Europe, p. x.

Tyjje locality. — Santo Domingo de Silos, Burgos, Spain.

Geographical distribution. — Essentially the entire Iberian
Peninsula.

Diagnosis. — General colour not so dark as in A. ampMhlns
amphibius, the sides and face a clear yellowish brown without
noticeable sprinkling of blackish hairs.

Colour. — Upper parts a yellowish brown varying between the
ochraceous-buff and clay-colour of Ridgway, and not infrequently
tinged with russet, the back and crown sufficiently sprinkled with
black-tipped hairs to produce an evident effect of " lining," the
sides and cheeks nearly clear ; underparts light ochraceous-buff
clouded to a variable degree by the slaty grey under-colour ; feet
drab-grey ; tail brownish, lighter below than above.

Measurements. —-External measurements of type (adult female) :
head and body, 187 ; tail, 123 ; hind foot, 34 ; ear, 18. Average

734

and extremes of seven adults from the type locality : head and
body, 197 (187-205); tail, 126-5 (122-130); hind foot, 34-4
(34-36); ear, 18*4 (18-19). Avei-age and extremes of six
adults from Lerida, Spain: head and body, 203 '5 (195-210);
tail, 122-6 (115-130); hind foot, 33-8 (33-35). Average and
extremes of ten adults from Panticosa, Huesca, Spain : head and
body, 210 (200-218); tail, 125-6 (119-133); hind foot, 34-2
(33-35). For cranial measurements see Table, p. 737.

Specimens examined. — One hundred and twenty-four, from the
following localities : —

Spain : L6rida, 13 ; Panticosa, Huesca, 32 ; Jaca, Huesca, 6 ; Arre-
chavaleta, Vitoria, 1; Pajdres, Leon, 6; La Corufia, 2 (not typical,
approaching tenehricus) ; Silos, Burgos, 20 ; vicinity of Madrid, 1
(U.S.N.M.) ; Villalba, Madrid, 1 ; Barracas, Castellon, 12 ; Silla, Valencia, 1 ;
Venta del Baul, Granada, 1 ; Seville, 7 ; near Tarifa, Cadiz, 1 ; Audalucia, 5.

France : Foret de Bouconne, Gers, 1 (not typical) ; Leguevin, Haute-
Garonne, 6 (not tyx^ical, approaching tenehricus) ; St. Gilles, Gard, 1 (not
typical, approaching tenehricus) ; Valescure, Var, 5 (not typical) ; Var, no
exact locality, 2 (not typical).

BemarJiS. — Throughout Spain the characters of Arvicola sajndus
sapidus are very constant ; only in the extreme north-west does
there appear to be any tendency toward the darker A. s. tene-
hricus. The French specimens from the neighbourhood of
Toulouse and from the marshes near the mouth of the Rhone
agree with true sapidus in the general light yellowish hue of the
back and sides, though showing a distinct approach to tenehricus
in the noticeable sprinkling of black hairs on cheeks.

L6rida, Spain. {N. Gon- O. Thomas (p). 8. 2. 9. 192-201.

zalez.)
Panticosa, Huesca, 0. Thomas (p). 8.2.9.172-183.

Spain. [N. Gonzalez.)
Jaca, Huesca. {N. Gon- O. Thomas (p). 8. 2. 9. 184-187.

zalez.)
Arrechavaleta, Vitoria. 0. Thomas (p). 8. 2. 9. 203.

{N. Gonzalez.)
Pajares, Leon. {N. Gon- 0. Thomas (p). 8. 2. 9. 188-191.

zalez.)
Silos, Burgos.

4 c5, 3 (J juv.

2 9, 9 juv.

8 (5, 4 9.

3<?, 9.

6.

2 9,2 9.

2 J, 5 9.

9.

1 6,6 9.

6.

4 9, 2 9.
Juv. al.

6.

1.

3 al, 2.

6.

Burgos.

G. S. Miller (c). 8. 8. 4. 113-119.

(8. 8. 4. 115. Type of species.)
S. & N. Gonzalez 8. 7. 7. 25.

0. Thomas (p).

Barracas, Castellon.

[N. Gonzalez.)
Silla, Valencia. {N. Gon- 0. Thomas (p

zalez.)
Seville. (Dr. Ruiz.)
Seville.

8. 2. 9. lGO-171.
8. 2. 9. 202.

Villalba, Madrid.

{N. Gonzalez.)
Tarifa, Cadiz.

Andalucia.

Foret de Bouconne,Gers.
{A. Robert.)

Lord Lilford (p). 95. 3. 3. 23-28.

Dr. V. L. Seoane 94. 3. 19. 3.

(c & p).

0. Thomas (p). 8. 2. 9. 204.

Col. L. H. Irby 94. 4. 27. 1.

(c & p).

Lord Lilford (p). 94. 9. 2G. 1-5.

0. Thomas (p). 8. 9. 1. 76.

ARVICOLA 735

6. Legueviu, Haute- O. Thomas (p). 8.9.1.77.

GaroDiie. {A. Robert.)

?. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4. 2G1.

2 <^, juv. Valescure, Var, France. G. S. Miller (c). 8. 8. 4. 257-2G0.

6 2. Var. Purchased (Par- 52. 5. 27. 44-45.

zudaki).

Arvicola sapidus tenebricus Miller.

1884. Microtus musiniani Lataste, Actes Soc. Linn. Bordeaux xxxviii,

p. 37 (part). Not Arvicola musignani de S^lys-Longchamps.
1908. Arvicola tenebricus Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 196, February, 1908. Type in British Museum.
1910. Arvicola sapidus tenebricus Miller, Proc. Biol. Soc. Washington,

XXIII, p. 20, IMarch 23, 1910.
1910. Arvicola sapidus tenebricus Trouessart, Faune Mamm. d'Europc,

p. X.

Type locality. — Vicinity of Biarritz, Basses-Pyrenees, France.

Geographical distribution. — Pyrenees and Atlantic coast region
of south-western France, north to the Garonne ; northern limit
of range not known ; two specimens from La Coruiia, Spain,
appear to be referable to this race rather than to true sapidus.

Diagnosis. — Colour darker than in A. sapidus sapidus (essen-
tially like that of A. amphihius amphibius), the sides and face
conspicuously sprinkled with black-tipped hairs.

Colour. — Upper parts a dull greyish buff, so heavily overlaid
with black that the general effect is usually not far from a rather
light grizzled bister on back and a greyish wood-brown on sides,
the cheeks and sides noticeably sprinkled with black-tipped hairs ;
underparts slaty grey, washed with light ochraceous-buff on chest
and belly ; feet hair-brown ; tail blackish above, greyish below,
seldom distinctly bicolor.

Measurem.ents. — External measurements of type (adult male) :
head and body, 193 ; tail, 112 ; hind foot, 34 ; ear, 17. Average
and extremes of five adults from the type locality : head and body,
197-4(193-202); tail, 118-5 (112-124) ; hind foot, 34-4 (33-36) ;
ear, 17-2 (17-18). For cranial measurements see Table, p. 737.

Specimens examined. — Twenty-seven, from the following localities : —
France : — Port6, Pyren6es-0rientales, 2 ; I'Hospitalet, Ari^ge, 2 ; Luchon,
Haute-Garonne, 2 ; Montr^jeau, Haute-Garonne, 5 (U.S.N.M.) ; Biarritz,
Basses-Pyr6n^es, 7 ; Cadillac, Gironde, G (U.S.N.M.); Barsac, Giroude, 1
(Lataste).

Spain : — Coruna, Galicia, 2.

2 6. Port6, Pyrenees - Orien- G. S. Miller (c). 8. 8. 4. 254-255.

6. L'Hosp'italet, Ariego. G. S. Miller (c). 8. 8. 4. 25G.

6 juv. L'Hospitalet, Ariege. O. Thomas (p). 8. 9. 1. 71.

(A. Robert).
5 (5, 2 9. Biarritz, Basses-Pyrenees. J. F. Davison (p). G. 1. 21. 5-6.

6. G. 4. 10-14.
(6. 1. 21. 5. Type of subspecies.)
2. Coruna, Galicia, Spain. Dr. V. L. Seoauo 93. 12. 15. 1.

(p). 95. G. 25. 1.

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738 RODENTIA

ARVICOLA TERRESTRIS LiiinJBUS.

1758. [Mils'] tcrrcstris Linntcus, Syst. Nat. i, 10th ccL, p. 61 (Sweden).

1771. [Mils] imludosiis Linnaeus, Mantissa Plautarum, pt. 2, p. 522
(Sweden).

1827. [Htj])uclrus tcrrcstris] j3 atcr Billberg, Synopsis Faunae Scandinaviae,
p. 4 (Gottland, Sweden).

1827. [llyimdcvus imludosus] & littoralis Billberg, Synopsis Faunse Scandi-
naviae, p. 5 (Smaland, Sweden).

1827. [Hypuchmis paludosus] y aquaticus Billberg, Synopsis Faunre Scandi-
naviae, p. 5 (South Sweden).

1858. Arvicola ampliibms a. Blasius, Siiugethiere Deutschlands, p. 344
(part).

1910. Arvicola tcrrcstris Miller, Proc. Biol. Soc., Washington, xxiii, p. 20,
March 23, 1910.

1910. Arvicola tcrrcstris Trouessart, Faune Mamm. d'Europe, p. x.

Type locality. — XJpsala, Sweden.

Geographical distrihution. — Scandinavian Peninsula^ eastward
into Finland ; limits of range not known.

Diagnosis. — Size less than in Arvicola ampliibins, but general
proportions not peculiar (head and body about 175 mm., tail
about 100 mm., hind foot usually 28 to 31 mm., condylobasal
length of adult skulls, 36 to 39 mm.) ; colour dark, essentially as
in A. amphihitis, but cheeks usually more yellowish than surround-
ing parts ; skull slightly but evidently fossorial in form, the
rostrum and occiput tending to be obliquely truncate, the upper
incisors noticeably projecting forward ; interparietal tending to
be sub-quadrate in outline ; teeth rather heavy, but roots of m^
and m.2 never forming protuberances on lower surface of mandible ;
habits both aquatic and mole-like.

External characters. — Aside from the somewhat less robust
treneral form correlated with the animal's smaller size there
appear to be no tangible external characters to distinguish
Arvicola terrestris from A. amphihius.

Colour. — Upper parts varying from broccoli-brown to mars-
brown, heavily overlaid with black, the latter usually though
not always in excess and producing a general effect not far from
seal-brown, especially along median dorsal region ; underparts a
distinctly rusty ochraceous-buff, in some specimens almost tawny,
everywhere dulled by the slaty under-colour, and fading on throat
to an indefinite yellowish grey ; cheeks and sides of head,
frequently to and including ears, tinged with rusty like that of
underparts, the suffusion usually producing a decided contrast
with surrounding parts ; feet varying from hair-brown to a deep
blackish brown ; tail blackish throughout, the under surface often
sprinkled with greyish hairs.

Skull. — The skull is not so large as that of Arvicola amphihius
or A. sapidus. In form it differs from the skulls of the strictly
aquatic members of the genus in a decided tendency to assume a

ARA'ICOLA 739

fossorial structure. Tliis is shown by the more pronounced
forward projection of the upper incisors, and more particularly
by the shortening and widening of the brain-case in general and
the slightly greater obliquity of the occipital region. Flattened

Fig. 153.
Arvicola terre.stris. Nat. size.

area between lateral ridges on brain-case tending to assume a
diamond-shaj)ed outline rather than that of an irregular paralello-
gram. Antero-posterior diameter of interparietal usually more
than half transverse diameter, the outer margins of the bone
almost squarely truncate.

Teeth. — Except for their slightly smaller size the teeth
exactly resemble those of Arvieola ampMbius.

Measurements. — Adult female from Holme, Mandal, Norway :
head and body, 167 ; tail, 94) ; hind foot, 29. Adult male from
Christiania, Norway : tail, 105 ; hind foot, 29. Adult from
Vefsen, Nordland, Norway : hind foot, 31 ; ear, 16. Adult
female from Jem tland, Sweden: head and body, 187; tail, 104;
hind foot, 30. Two adults from near Stockholm, Sweden : hind
foot (dry), 27 "6 and 29*4. For cranial measurements see Table,
p. 742.

Specimens examined. — Thirty-eight, from the following localities : —
NoBWAY : — Vefsen, Nordlaud, 1 ; Elverum, Hedemarken, 1 (U.S.N.M.) ;

Christiania, 1 (U.S.N.M.); Asker, Christiania, 1 (U.S.N.M.); Holme,

Mandal, 15.

Sweden :—Jemtland, 7; Upsala, 2 (B.M. and U.S.N.M.); Upland, 2

3 B 2

740 RODENTIA

(U.S.N.M.) ; near Stockholm, 6 (B.M., U.S.N.M. and Stockholm) ;
Aspvik, 1 ; Drottningholm, 1 (U.S.N.M.).
Finland : — Drumso, 1.

Bemarlis. — Arvicola terrestris is readily distinguishable from
A. aniphihiiis by its smaller size, brighter coloured cheeks, and by
the slightly fossorial modification of the skull. In habits it is
less strictly aquatic than the British animal.

1. Vefsen, Nordland, Nor- E. G. B. Meade 5. 7. 1. 3.

way. Waldo (c & p).

6 c5, 9, Holme, Maudal. R. J. Cuninghamc 8. 8. 9. 19-29.

(c & p).

6. Upsala, Sweden. Lord Lilford (p). 0. 5. 15. 1.
{E. KoUhoff.)

2. Near Stockholm. Prof. Sundevall (p). 49. 11. 1, 12-13.
1 al. Aspvik. Stockholm Museum 90. 8. 1. 14.

(E).

? Drumso, Finland. Dr. Schulman(c&p). 1. 6. 9. 3.

ARVICOLA ITALICUS Savi.

1839. Arvicola ampliibius var. italica Savi, Nuovo Giorn. de' Letterati,

Pisa, xxxvii. No. 102, p. 202 (p. 5 of separate), February 1839 (for

date see de Selys-Longchamps, Micrommalogie, p. 93). Vicinity

of Pisa, Italy.
1839. Arvicola j^ertinax Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvii,

No. 102, p. 203 (p. 6 of separate), February, 1839 (MS. synonym

of italica).
1845. ? 'iArvicola amphihius var. viinor De Selys-Longchamps, Atti dclla

sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 322

(nomen nudum).
1910. Arvicola italicus Miller, Proc. Biol. Soc. Washington, xxiii, p. 20,

March 23, 1910.
910. Arvicola italicus Trouessart, Faune Mamm. d'Europe, p. x.

Type locality. — Vicinity of Pisa, Italy.

Geographical distribiition. — Italian Sv^'itzerland and northern
Italy, south at least to the vicinity of Pisa. Details of distribu-
tion not known.

Diagnosis. — Similar to Arvicola terrestris, but teeth not so
heavy and colour not so dark, the underparts washed with
yellowish brown instead of rusty, the cheeks not contrasting
noticeably with surrounding parts.

Colour. — ^Elements of colour of upper parts essentially as in
Arvicola terrestris, but broccoli-brown much in excess of black,
the latter producing a slight effect of grizzling or " lining," but
never sufficiently dominant to make the general colour approach
seal-brown ; sides light, bufFy, slightly grizzled wood-brown,
becoming a little more yellowish on cheeks. Underparts light
slaty grey washed with buffy on chest and belly. Feet a light
hair-brown, sometimes tinged with drab. Tail obscurely bicolor,
dark brown above, greyish below.

ARVICOLA 741

Skull mid teeth. — The skull resembles that of Arvicola terrestrh
in size and general appearance, but the brain-case is deeper and
noticeably longer in proportion to its breadth, the general outline
thus approaching that of the large numbers of the group.
Auditory bullae slightly larger than in the Scandinavian species,
but not peculiar in form. Teeth as in A. terrestris, but slightly
larger.

Measurements. — ^Average and extremes of five adults from
Lugano, Ticino, Switzerland: head and body, 180*6 (174-191);
tail, 87-8 (84-90); hind foot, 27-7 (27-29). Adult male and
female from Locarno, Ticino, Switzerland : head and body,
179 and 171; tail, 102 and 98; hind foot, 29 and 27-5; ear,
15 "5 and IG. Adult female from Ceresole d'Alba, Turin, Italy :
head and body, 163; tail, 90; hind foot, 29. For cranial
measurements see Table, p. 743.

Specimens examined. — Twenty-nine, from the following localities : — •
Switzerland: Lugano, Ticino, 19 (U.S.N.M. and Mottaz) ; Locarno,

Ticino, 4.
Italy: Casaleone, Verona, 1 (U.S.N.M.); Modena, 3; Ceresole d'Alba,

Turin, 1 (Turin) ; no exact locality, 1.

2 c?, 2 9. Locarno, Ticino, Swit- 0. Thomas (c & p). 2. 7. 1. 2-5.
zerland.

?,2juv.al. Modena, Italy. Marquis G. Doria (p). 90.3.5.1-3.

1. Italy. Purchased (Brandt). 46. 1. 9. 20.

ARVICOLA ILLYRICUS Barrett-Hamilton.

1899. Microtus musignani illyricus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th sor., iii, ]}. 225, March, 1899. Type in British Museum.
1910. Arvicola illyricus Miller, Proc. Biol. Soc. Washington, xviii, p. 21,

March 23, 1910.
1910. Arvicola illyricus Trouessart, Paune Mamm. d'Europc, p. x.

Type locality. — Bosnia, no exact locality.

Geographical distribution. — Bosnia. Limits of range not •
known.

Diagnosis. — Similar to Arvicola italicus, but underparts with
a decided whitish wash.

Colour. — The colour is in all respects similar to that of
Arvicola italicus, except that the chest and belly are a whitish
grey, with only the faintest tinge of buff.

SJcidl and teeth. — -The imperfect skull of the type shows
nothing to distinguish it from that of Arvicola italicus.

Measurements. — -Type: hind foot, 30; ear, 15. For cranial
measurements see Table, p. 743.

Specimen examined. — The type.

Hemarlis. — The Bosnian water vole is so similar to Arvicola
italicus that the two animals will not improbably prove to be

742

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identical. A specimen from Breba, Roumania, in some respects
resembles the type of ArvicoJa ilhjricas, but the small size of the
hind foot (25 mm.) renders its identification uncertain.

6. Bosnia. Dr. Ploericke (c). 94. 1. 5. 1.

(Type of species.)
6. Breba, Roumania. (W. Dodson.) Lord Lilford (p). 4. 4. G. GG.

ARVICOLA MUSIGNANI de 8elys-Longchamps.

1839. Ai-vicola musignani de S61ys-Longchamps, Revue Zoologique, p. 8,

January, 1839 (Rome, Italy).
1839. Arvicola destructor Savi, Nuovo Gioru. de' Letterati, Pisa, xxxvii,

No. 102, p. 204 (p. 7 of separate), February, 1839. (Maremma

Grossetaua, Tuscany, Italy). For date of publication see de Selys-

Longchamps, Micromammalogie, p. 93.
1845. ? [Arvicola vnisignani'] var. fuliginosus de Selys-Longchamps, Atti

della sesta Riuniono degli Scienziati Italiani, Torino, 1844, p. 322

(nomen nudum).
1857. Arvicola amphihius b. Blasius, Siiugethiere Deutschlands, p. 344.
1910. Arvicola musignani Miller, Proc. Biol. Soc. Washington, xxiii,

p. 21, March 23, 1910.
1910. Arvicola musignanoi Trouessart, Faune, Mamm. d'Europe, p. x.

Type locality. — ^ Vicinity of Rome, Italy.

GeograjjJiical distribution. — Central Italy ; at present known
from the west coast only.

Diagnosis. — Size and general characters as in Arvicola it aliens,
but colour pale and yellowish, like that of A. sapidus sapidus.

Colour. — Entire animal a light yellowish wood-brown, faintly
grizzled with black on median dorsal region and clouded by the
slate-gi-ey underfur on throat and belly ; cheeks faintly more
yellowish than sides of body. Feet light drab. Tail obscurely
bicolor, blackish brown above, yellowish brown below.

Shull and teeth. — The skull and teeth agree with those of
A. italicus, except that the molars appear to be usually somewhat
smaller.

Measurements. — ^Two adult males from Arsoli, Rome, Italy :
head and body, 190 and 201 ; tail, 94 and 95 ; hind foot, 29 'G
and 29; ear, 15 and 15. For cranial measurements see Table,
p. 743.

Specimens examined. — Seven, from the following localities in Italy :
Arsoli, Rome, 6 ; Velletri, Rome, 1 (Mottaz).

6. Arsoli, Rome, Italy. G. Barrett-Hamilton (p). 11. 1. 2. 37.

ARVICOLA SCHERMAN Shaw.

(Synonymy under subspecies.)

Geographical distribution. — West-central continental Europe,
from the Pyrenees and Alps to the Baltic ; eastern limits of
ranse not known.

ARVICOLA 745

D!a</)ws!s. — Essentially as in Armcola ferresfris, but more
modified for fossorial life, some of the races having become
completely terrestrial ; palmar and plantar tubercles reduced,
occupying less than half surface of region in which they occur ;
skull distinctly fossorial in form, the incisors strongly protruding ;
interparietal tending to be narrow and ligulate in outline.

Skull and teeth. — The skull does not difter essentially from
that of Arvicola terresfris, but its fossorial characters are more
accentuated, especially in the terrestrial forms. Teeth without
special peculiarities.

Beviarks. — It may eventually be shown that some form of
Arvicola scherman intergrades with A. terrestris, but so far as the
material examined is concerned the two animals, though nearly
I'elated, are distinct. Three races are now known, one of them
partly acquatic in its habits, the two others strictly terrestrial.

Arvicola scherman scherman Shaw.

1779. ? Sl-palax] minor Lesko, Anfangsgrunde der Naturgosch., i, p. 168

(Germany).
1801. Mm scherman Shaw, Gen. Zool., u,-'pt. i, p. 75 (Strassburg,

Germany).
1801. M[us] aviph{ibius] alhus Beohstein, Gemeinn. Naturgesch. Deutsch-

lauds, r, 2d ed., p. 985 (Thiiringen, Germany).
1801. M[us\amph[ibius\ carras Bechsteiu, Gemeinn. Naturgesch. Deutscli-

lands, I, 2d ed., p. 985 (Thiiringen, Germany).
1804. Mus terrestris Hermann, Observ. Zool., p. 59 (not of Linnseus, 1758).
1804. [Mus] scherviaus Hermann, Observ. Zool., p. 59. Alternative for

terrestris ; proposed but rejected on the ground that the animal

was the same as that of Linnaeus (Strassburg, Germany).
1822. Arvicola argcnioratensis Desmarest, Mammalogie, pt. ii, p. 281

(Strassburg, Germany).
1829. Lemmus arvalis yS buffonii Fischer, Synops. Mamm., p. 293 (near

Berlin, Germany?)." Based on Brants's account (Het Geschlact

der Muizen, p. 372) of two specimens in the Berlin Museum

supposed to be identical with the dark variety of " Le Campagnol "

described by Buffon, Hist. Nat. des Anim. vii, p. 372.
1839. Arvicola terrestris de S61ys-Longchamps, Etudes de Micromamm.,

p. 97, pis. I and ii, fig. 6.
1858. Arvicola amphibius, c. Airvicola} terrestris auct., Blasius, Siiuge-

thiere Deutschlands, p. 355 (part).
1910. Arvicola scherman scherman Miller, Proc. Biol. Soc. Washington,

XXIII, p. 21, March 28, 1910.
1910. Arvicola scherman Trouessart, Faune Mamm. d'Europe, p. x.

Type locality. — Strassburg, Germany.

Geographical distribution.- — Continental Europe from the
Baltic south into central France and southern Germany ; limits
of range imperfectly known.

Diagnosis. — Sole wrinkled but not conspicuously granular ;
palmar and plantar tubercles relatively smaller than in Arvicola
terrestris, though not so much reduced as in the strictly terrestrial
forms ; length of hind foot about 26 • 5 mm. ; condylobasal length

746 , RODENTIA

of fully adult skulls, 35*6 to 36*4 mm. ; colour of upper parts a
dark brown, usually much clouded with black ; tail usually dark
brown throughout. Habits partly terrestrial and partly aquatic.

External characters. — General form essentially as in Arvicola
terresiris. Palmar and plantar tubercles distinctly reduced as
compared with those of the Scandinavian animal ; sole wrinkled
near heel but not distinctly granular, and without evident
pubescence.

Colour. — ^Upper parts a dark brown, usually approaching the
prout-brown of Ridgway along median dorsal area from muzzle
to base of tail, and mars-brown or broccoli-brown on sides. In
some specimens there is a general suffusion of light wood-brown,
and the sides may even show a faint tinge of dull buffy grey.
Cheeks concolor with sides or nearly so, seldom if ever forming
any noticeable contrast with surrounding parts. Chest and belly
slaty grey variously washed with dull ochraceous-buff, the throat
usually about the slate-grey of Ridgway. Feet varying from
hair-brown to blackish. Tail blackish throughout or sj)rinkled
with whitish hairs below, never distinctly bicolor.

Measurements. — Adult female from Brunswick, Germany :
head and body, 171; tail, 102; hind foot, 27. Three adults
from Nuremberg, Germany: hind foot, 26, 26 '6 and 27. For
cranial measurements see Table, p. 750.

Specimens examined. — Forty-three, from the following localities : —

Belgium : Waremme, Liege, 3 (U.S.N.M.).

France : La Fert6-Milon, Aisne, 1 (Lataste).

Germany: Stolpin, Pomerania, 1; Brunswick, 3 (U.S.N.M. and
Merriam) ; Ingelheim, Rheinhessen, 7 ; Heidelberg, Baden, 1 (U.S.N.M) ;
Lotzen, Saxony, 1 (U.S.N.M.) ; Ummerstadt, Thiiringen, 1 ; Strass, near
Burgheim, Bavaria, 5 ; Marxheim, near Monheim, Bavaria, 1 ; Nuremberg,
Bavaria, 6 (U.S.N.M.) ; South Germany (no exact locality), 2; Strassburg,
Alsace, 1 ; Niesky, Silesia, 2.

Austria-Hungary : Hainspach, Bohemia, 4 (U.S.N.M.) ; Zuberec, 4.

1 al. Stolpin, Pomerania. Dr.H. Gadow (c & p). 82. 7. 31. 1.

4 (^, 3 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 52-58.

9. Strassburg, Alsace. 0. Thomas (p). 8. 8. 10. 100.

(C. Mottaz.)
6, juv. Niesky, Silesia. (W.Baer.) Dr. E. Hamilton (p). 97. 12. 4. 31-32
4. Zuberet', Hungary. Budapest Museum (e). 94. 3. 1. 61-64.

Arvicola scherman exitus Miller.

1839. Arvicola tcrfestris Savi, Nuov. Giorn. de' Letterati, Pisa, xxxvii,

p. 300 (p. 3 of separate). Not Mus terrestris Linnaeus. (Geneva,

Switzerland.)
1845. ? [Arvicola terrestris'] var. niger De S^lys-Longchamps, Atti della

sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 321

(Lausanne, Switzerland). Nomen nudum.
1845. ? [Arvicola terrestris'] var. castancus De S^lys-Longchamps, Atti

della sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 321

(Lausanne, Switzerland). Nomen nudum.
1857. Arvicola ampliibius c. A[rvicola^ terrestris Auct., Blasius, Siiugethiere

Deutschlands, p. 355 (part).

\ ARVICOLA 747

1910. Arvicola schcrman exitus Miller, Proc. Biol. Soc. Washington, xxiii,
p. 21, March 23, 1910 (St. Gallen, Switzerland). Type in British
Museum.

1910. Arvicola scherman exitus Trouessart, Fauna Mamm. d'Europe,p. x.

1911. Arvicola scherman exilis Lydekker, Zool. Record, xlvii (1910),

Mamm., p. 54 (Accidental renaming of exihis).

Type locality. — St. Gallen, Switzerland.

Geographical distribution. — Alps (not known from the Italian
side) at moderate altitudes, and immediately adjoining lowlands
of Switzerland and France ; eastward into Tirol ; northward
into the Vosges Mountains ; limits of range not known.

Diagnosis. — ^Sole nearly smooth ; palmar and plantar tubercles
much reduced, occupying distinctly less than half area of region
in which they occur ; length of hind foot, 22 to 25 mm. ; condy-
lobasal length of fully adult skulls, 33 to 35 mm. ; colour of
upper parts a light yellowish brown, usually without much black
clouding ; tail buffy throughout or evidently bicolor (never
uniformly blackish) ; auditory bulljB not highly inflated, their
surface often irregularly flattened ; anterior loop of m^ short and
wide. Habits strictly terrestrial, mole-like.

External characters. — Compared with Arvicola amjihihius this
animal shows the following peculiarities in external form. Upper
incisors more conspicuously protruding from mouth, and lower
teeth also much exposed, the lower lip apparently too short to
cover them. Front foot with anterior tubercles much more
reduced as compared with posterior pair, the three together
scarcely half as large as postero-external pad. (In A. amphihius
the three anterior tubercles together decidedly exceed bvilk of
postero-external pad.) Hind foot showing only a slight tendency
to " feathering." Sole with posterior portion nearly smooth.
Plantar tubercles occupying decidedly less than half surface of
area in which they occur, their size noticeably less than corre-
sponding pads at base of claws ; that at base of first digit
scarcely larger than a dust shot ; that at base of fifth toe about
three times as large, and sub-equal to the two other anterior
pads ; postero-internal tubercle larger and more elongate than
any of the others, but less noticeably so than in A. amphihius.
An exceedingly rudimentary sixth tubercle sometimes present.
Tail with annulations narrower than in A. amphihius, about 20
to the centimeter at middle.

Colour. — Upper parts varying from a light broccoli-brown to
ochraceous-bufF, the face, crown and median dorsal area faintly
darkened by blackish hair-tips, these usually most noticeable in
lumbar region ; sides in brightest specimens clear ochraceous-
buff, in others more nearly a greyish cream-bufF; underparts a
paler, less slaty grey than in 31. scherman scherrimn, the throat
nearly the grey No. 9 of Ridgway ; chest and belly washed with
cream-bufF; feet ecru-drab, sometimes with a buffy tinge ; tail

748

RODENTIA

whitish throughout or sprinkled witli blackish above, sometimes
rather distinctly bieolor.

Skull and teeth. — The skull is slightly smaller than that of
Arvicola scherman scherman (condylobasal length in fully adult
specimens seldom if ever exceeding 35 mm.), and the fossorial
characteristics, particularly the sliallowness of the rostnnn as

Fig. 154.
Arvicola scherman exitus. Nat. size.

compared with brain-case, are carried to the extreme. Teeth as
in the typical form, but upper incisors even more strongly
projecting.

Measurements. — ^External measurements of type (adult female) :
head and body, 138 ; tail, 64 ; hind foot, 24 ; ear, 13. Average
and extremes of eight adults from the type locality : head and
body, 146-6 (135-165); tail, 62-7 (56-70); hind foot, 23-7
(23-25). Adult male and female from Vitznau, Lucerne,
Switzerland: head and body, 131 and 133; tail, 67 and 61;
hind foot, 24 • 5 and 23 • 5 ; ear, 13 and 12. Average and extremes
of six adults from Les Plans, Vaud : hind foot, 23 '3 (23-25).
Average and extremes of five adults from Cranves-Sales, Haute-
Savoie, France : head and body, 138 (135-140) ; tail, 54 (52-57) ;
hind foot, 22-5 (22-23); ear, 11-8 (11-12). For cranial
measurements see Table, p. 750.

Specimens examined. — Eighty-two, from the following localities: —
Prance : Gerbamont, near Vagney, Vosges, 6 (Lataste) ; Cranves-
Sales, Haute-Savoie, 8 ; Lucinges, Haute-Savoie, 4 ; Montauban, Haute-
Savoie, 2 ; Scientriers, Haute-Savoie, 1 (Mottaz) ; Etupes, Doubs, 1
(Mottaz; not typical).

ARVICOLA

749

Switzerland : Geneva, 9 (B.M. and ]Mottaz) ; Nyon, Vaud, 1 (Mottaz) ;
Chesi^res, Vaud, 2 (Mottaz) ; St. Cerguos, Vaud, 1 (Mottaz) ; Lausanne,
Vaud, 2 (U.S.N.M.) ; Les Plans, Vaud, 10 (U.S.N.M.) ; Meiringon, Berne,
6 (U.S.N.M.) ; Vitznau, Luceruo, 2 ; St. Gallen, 1.3 (B.M. and U.S.N.M.) ;
Mels, S. Gallen, 4 (U.S.N.M.); Ziiberwangen, St. Gallen, 2 (B.M. and
U.S.N.M.) ; Roggweil, Thurgau, 3 (U.S.N.M.) ; Fribourg, 1 ; no exact
locality, 3.

Austria-Hungary: Mittelberg, Voraiiberg, 1.

Bemarks. — This form is immediately distinguishable from
A. scherman scherman by its smaller size, yellower colour, and
more complete adaptation to underground life. Its habits so
far as known are strictly terrestrial and mole-like.

2(5. Cranves-Sales, Haute-Savoie, A. Robert (c & p). 5.4.9.6-7.
France.

3 6,3 9. Cranves-Sales, Haute-Savoic. 0. Thomas (p). 5. 11. 18. 18-23.

1200 m. (A. Bobert.)

2. Montauban, Haute-Savoio. A. Robert (c & p). 97. 1. 9. 5-6.

1. Geneva, Switzerland. Baron de Selys- 55. 12. 24. 358.

Longcbamps (p).

6, 9. Vitznau, Lucerne. O. Thomas (c & p). 5. 8. 3. 19-20.

6. Ziiberwangen, St. Gallen. 0. Thomas (p). 2. 8. 4. 48.

(E. H. ZoUikofer.)

4 t5, 4 9. St. Gallen. {E. H. ZoUikofer.) 0. Thomas (p). 10. 8. 16. 1-8.

(10. 8. 16. 8. Type of subspecies.)
1. Fribourg. L. 0. Galliard (p). 75. 9. 20. 1.

1. Switzerland. Baron de Selys- 45. 7. 5. 5.

Longcbamps (p).

2. Switzerland. [Muscliler.) E. R. Alston (p). 79. 9. 25. 47-48.
1. Mittelberg, Vorarlberg, Aus- O. Thomas (p). 8. 11. 30. 17.

tria-Hungary. {Dr. Major.)

Arvicola scherman monticola de Selys-Longchamps.

1838. Arvicola monticola de S61ys-Longchamps, Revue Zoologique, p. 249.

1839. Arvicola monticola de S61ys-Longchamps, Etudes do Micromamm.,

p. 92, pis. I and ii, fig. 3.
1858. Arvicola amphibius, c. A[rvicola] tcrrcstris auct., Blasius, Saugethiere

Deutschlands, p. 355 (part).
1910. Arvicola scherman mo7iticola Miller, Proc. Biol. Soc. Washington,

xsiii, p. 22, March 23, 1910.
1910. Arvicola scherman monticola Trouessart, Faune Mamm. d'Europe,

p. X.

Type locality. — 'St.^Bertrand de Comminge, Hautes-Pyrenees,
France.

Geographical distribution. — Pyrenees and their immediate
neighbourhood (known at present from the French side only) ;
Puy-de-Dorae ?

Diagnosis. — Like Arvicola scherman exitus, but auditory bullae
usually larger and more evenly inflated, and first lower molar
with anterior loop longer and narrower than in the Alpine form.
Habits strictly terrestrial, mole-Uke,

750

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Measurements. — Avei'age and extremes of four adults from
Caterille, Haute - Garonne, France: head and body, 147 "2
(145-150); tail, 60-8 (59-67); hind foot, 24-3 (24-25); ear,
13 (12-14). Adult male from Luchon, Haute-Garonne, France:
head and body, 150; tail, 58 ; hind foot, 25; ear, 12. Average
and extremes of fovu' adults from Biarritz, Basses-Pyrenees,
France: head and body, 144-7 (143-147^; tail, 63-7 (62-66);
hind foot, 23-5 (23-24); ear, 12-2 (12-13). For cranial
measurements see Table, p. 751.

Specimens examined. —Twenty, from the following localities in south-
western France : Caterille, Haute-Garonne, 4 (U.S.N.M.) ; Luchon, Haute-
Garonne, 3 (U.S.N.M.) ; Biarritz, Basses-Pyrenees, 11 ; Pyrenees, no exact
locality (probably topotypes), 4 (B.M. and U.S.N.M.).

Hemarlis. — The Pyrenean form of Arvicola scherman, though
resembling the Alpine A. scherman exitus in colour, external
characters and general form of skull, differs rather constantly
in the greater inflation of the auditory bullse and in the outline
of the anterior loop of m^.

Two mounted specimens in the Paris Museum, collected in
the neighbourhood of Mont-Dore, Puy-de-.D6me, France (Nos.
1447 and 1510), while undoubtedly of the exitus type, are not, in
their present condition, determinable with certainty. It is not
impossible that they may be referable to the present race.

3 6, 5 9. Biarritz, Basses-Pyrenees, J. F. Davison (c & p). 6. 6. 4. 15-22.
France.
1. Pyrenees. Baron E. de Selys- 45. 7. 5. 4.

Longchamps (p).
1. Pyrenees. Purchased (Frank). 45. 5. 23. 7.

1. Pyrenees. Stockholm Museum (e). 46. 6. 2. 66.

Genus PITYMYS McMurtrie.

1830. Psammomys Le Conte, Ann. Lye. Nat. Hist., New York, iti, p. 132

(P. pinetorum Le Conte). Not of Cretzschmar, 1828.

1831. Pitymys IMcMurtrie, Cuvier's Animal Kingdom, American edition, i,

p. 434 {Psammomys pinctorum Le Conte).
1831. Ammomys Bonaparte, Saggio Distrib. Metod. Anim. Vert., p. 20,

footnote {Psammomys pinetorum Le Conte).
1836. Pinemys Lesson, Hist. Nat. des Mammif. et Ois. d6couv. depuis

1788, Compl. Oeuvres de BuiTon, v, p. 436 {Psatiimomys pinetorum

Le Conte).
1857. Microtis Blasius, Saugethiere Deutschlands, p. 387 (sub-genus). Not

Microtus Schrank, 1798.

1867. Terricola Fatio, Les Campagnols du Bassin du Leman, p. 36 (sub-
genus to contain Arvicola subterraneus de Selys-Longchamps, and
A. savii de Selys-Longchamps). Not of Fleming, 1828.

1877. Micrurus Major, Atti della Soc. Toscana di Sci. Nat., iii, p. 126 (sub-
genus to contain Arvicola nehroclensis Mina-Palumbo).

1887. Pitymys Lataste, Ann. Mus. Civ. Stor. Nat. Genova, 2d ser., iv,
p. 266 (subgenus).

PITYMYS 753

189G. Pitymijs Miller, North American Fauna, No. 12, p. 58, July 23, 1896

(sub-genus).
1907. Pitymys Mottaz, Bull. Soc. Zool. de France, xx, p. 27, September

1907 (genus).

Ty2)e species. — Psammomys pinetorum Le Conte.

Geograpjliical disfrilmtion. — Central and southern continental
Europe, eastward into Asia Minor ; eastern and south-eastern
United States ; southern Mexico.

Characters. — Like Microius but sole with five tubercles, and
mammae only four ; external form slightly or considerably modified
for underground life ; skull in most species showing some trace
of fossorial modification, this character often conspicuous ; enamel
pattern variable, but first inner and first outer triangles of m,
broadly communicating.

MemarTis. — The genus Pitymys is remarkable as the most
conspicuous instance of discontinuous distribution in the sub-
family. It is also the genus of European voles which presents
the greatest diversity of cranial and dental characters. All the
European members of the genus, twenty-five of which are here
recognized, may at once be distinguished from the other voles
occurring in the same region by the broad communication of the
first inner and first outer triangles of m^.

KEY TO THE EUROPEAN FORMS OF PITYMYS.

Third upper molar longer than second, its inner side

normally with three re-entrant angles subterraneus group.

Dorsal profile of skull flat or faintly convex from

nasals backward ; brain-case much depressed,

its depth including auditory bullse about 65

per cent of occipital breadth.

Third inner re-entrant angle of m^ shallow

(Roumania) P. dachis, p. 760.

Third inner re-entrant angle of m^ deep.

Back strongly suffused with buff y ; feet whitish
in noticeable contrast with back (Basses-

Alpes) P. drucntius, p. 762.

Back without noticeable buffy suffusion ; feet
dusky, not noticeably contrasting with

back P.S2ibterrancus,j}. 755.

Brain-case moderately flattened ; colour

dark (Belgium to Transylvania) P.s. subterranfuf,-p. 7 58.

Brain-case much flattened ; colour less

dark (Puy-de-D6me region, France) P. s. capncimi.t, p. 760

Dorsal profile of skull obviously convex from
nasals backward ; brain-case slightly de-
pressed, its depth including auditory bullse
about 70 per cent of occipital breadth.
Condylobasal length of largest skulls about
23 mm. ; dorsal profile of skull very slightly
convex (Neighbourhood of Zermatt, Swit-
zerland) P.faiioi, p. 763.

Condylobasal length of largest skulls about
25 mm. ; dorsal profile of skull noticeably

convex (Northern Italy, &c.) P. multiplex, p. 764.

3 c

754

EODENTIA

Third upper molar not longer than second, its inner
side normally with two re-entrant angles.
Outer triangles of vi^ well developed, with three
definite enamel sides and a distinct central
osteodentins area normally isolated or nearly
so from that of inner triangle and posterior
loop ; middle outer salient angle as long as
anterior and posterior salient angles (third
inner re-entrant angle present in rare indi-
vidual cases) savii group.

Brain-case much depressed, its dorsal profile
nearly flat from nasal to back of interparietal

(Pyrenees) P. planiceps, p. 772.

Brain-case not much depressed, its dorsal profile
evidently convex.
Interparietal irregularly lozenge-shaped, its
lateral extremities acute, scarcely or not
in contact with temporals; first and
second triangles of vi^ broadly communi-
cating P. gerbii, p. 773.

Interparietal strap-shaped, its lateral extremi-
ties truncate, broadly in contact with
temporals ; first and second triangles of
7)1^ usually isolated.
General outline of brain-case sub-quadrate,

the upper surface rather flat (Italy) ... P. savii, p. 7G8.
General outline of brain-case not evidently
sub-quadrate, the upper surface rounded
off at sides.
Mesopterygoid space rather wide (Sicily) P. nebrodensis, p. 770.
Mesopterygoid space narrow (South-
western France) P. pyrcnaicns, p. 770.

General colour above clear hair-brown
without noticeable yellowish cast

(Pyrenees) P. i). pyrenaicus, p. 771.

General colour above brown with a
noticeable yellowish cast (Plains

of south-western Prance) P.p. brunneus, p. 772.

Outer triangles of m^ not well developed, usually
opening widely into inner triangle and pos-
terior loop, the central osteodentine area
small or absent ; middle outer salient angle
noticeably shorter than anterior and posterior
salient angles, sometimes nearly obsolete
(third inner re-entrant angle never present)... ibericus group.
Upper incisors moderately long and not strongly
projecting, not a highly conspicuous feature
of skull as viewed from above.
Brain-case much flattened, the occipital depth
about one-half occipital breadth (Central

Spain) -. P. deprcssiis, p. 779.

Brain-case not much flattened, the occipital
depth noticeably more than one-half
occij)ital breadth.
Upper incisors slightly projecting (Portugal) P. lusitanictcs, -p. 776.
Upper incisors nearly vertical.

Skull narrow ; outline of brain-case dis-
tinctly elongate (North-western

Spain) P. mari.T, p. 777.

Skull broad ; outline of brain-case not
specially elongated (North-central
Spain) P. pelandonius,'p.178.

PITYMYS 755

Upper incisors unusually long, and so strongly
projecting as to be a highly conspicuous
feature of skull as viewed from above.
Condylobasal length of fully adult skulls

about 23 mm. (South-eastern France) P. provincialis,]^. 7S5.

Condylobasal length of fully adult skulls

24 to 26-5 mm.

Size large, condylobasal length of largest

skulls about 26-5 mm.; transverse

diameter of auditory bulla about 7 mm.

(Montenegro) P. thoviasi, p. 78G.

Size medium, condylobasal length of largest
skulls not more than 25 mm. ; trans-
verse diameter of auditory bulla about
6 mm. or less.
Occiput so obliquely truncate that con-
dyles are conspicuous when skull is
viewed from above ; interparietal
narrowly ligulate (South - eastern
France) P. duodecimcostatus,

Occiput not very obliquely truncate, so ?• '
that condyles are not highly con-
spicuous when skull is viewed from
above ; interparietal broadly ligulate

or somewhat lozenge-shaped P. ihericus, p. 780.

Length of upper tooth-row about 5*4

mm. (Granada, Spain) P. i. regulus, p. 784.

Length of upper tooth-row 6 to G • 4 mm.
Colour above a dark bister brown

(Dehesa de Valencia, Spain) P. i.pascuus, p. 783.

Colour above a light yellowish brown.

General colour above hair-brown

with or without a buffy cast

(Central Spain) P. i. centralis, p. 782.

General colour above a pale buffy
broccoli-brown or drab (South-
eastern Spain) P. L ibericus, p. 782.

PITYMYS SUBTERRANEUS de Selys-Longchamps,

(Synonymy under subspecies.)

Geographical distribution. — Central Europe from Belgium to
central France, eastward through Switzerland to Transylvania.
Details of distribution imperfectly known.

Diagnosis. — Size small (hind foot usually about 14*5 to
15 "5 mm., condylobasal length of largest skulls, 22 to 23*4 mm.) ;
skull slender and lightly built, depressed, the dorsal profile nearly
flat or at most only a little convex from nasals backward ; brain-
case much depressed, its depth including auditory bulke about
G5 per cent of occipital breadth ; thii-d upper molar noticeably
longer than second, normally with three closed triangles, its
posterior loop elongated and with deep inner re-entrant angle ;
general colour a clear dark brown without noticeable sufiusion of
buffy ; feet dusky, not contrasting with colour of back.

External characters. — General external form essentially as in

3 c 2

756

RODENTIA

Microius arvalis, the tail about one-third as long as head and
body, the feet slender and without special modification for
burrowing, the incisors not protruding conspicuously from mouth.
Ear small, but scarcely more so than in Microius arvaJis, its
form with no special peculiarities except that meatal lobe is
slightly less developed ; both surfaces of ear conch essentially
naked at base, thinly clothed with rather long hairs distally.
Eye relatively smaller than in Microius arvalis. Muzzle not
peculiar. Front foot as in Microius arvalis, except that the claws
are slightly larger. Hind foot slender, its general form as in
M. arvalis, the sole similarly hairy behind tubercles, the tubercles
of the same form and relative size, their only peculiarity the
complete absence of the sixth. Tail as in M. arvalis. Mammje :
4, all inguinal.

Colour.- — Upper parts usually a dark nearly uniform hair-
brown, tinged with sepia along middle of back, everywhere
inconspicuously varied by silvery reflections and blackish hair-
tips ; ground colour sometimes more nearly approaching mars-
bi'own or wood-brown, but rarely if ever with any distinct bufiy
cast ; basal portion of hairs slate-black ; underparts pale smoke-
grey, occasionally with a slight wash of broccoli-brown, the slaty
bases of the hairs everywhere showing through at surface ; feet
an indefinite drab or dusky grey, often tinged with dark hair-
brown ; tail dull whitish grey beneath, dark brown above, the
two colours sometimes though not usually well contrasted.

Skull. — The skull is smooth and lightly built, with large,
somewhat flattened brain-case, rather wide interorbital region,
and small rostrum. Dorsal profile nearly straight from near
posterior extremity of nasals to a little behind middle of parietals,
then slightly convex to posterior margin of interparietal, beyond
which the occiput curves abruptly downward ; nasals sloping
forward at an angle of about 11°; ventral profile nearly straight
from about middle of rostrum to lower border of auditory bullfe,
the general outline of skull as viewed from the side rather
narrowly cuneate. Brain-case long as compared with that portion
of the skull which lies in front of it, the distance from back
of interorbital constriction to hinder border of interparietal
decidedly greater than breadth over roots of zygomata, its out-
line rather broadly oval, its surface smooth, or at most with
slightly indicated ridges along outer sides of parietals ; larabdoid
crests slightly developed at sides ; postorbital ridges low and
indistinct, rarely if ever developing an angular projection ; '•■
interparietal narrowly ligulate, the outer extremities usually
pointed, the anterior border with median projection ; in lateral
view the brain-case appears much flattened, its depth through
auditory bullae about 65 per cent of occipital breadth ; occiput

* A character that often proves convenient in distinguishing between
imperfect skulls of this species and the smaller forms of Microtus, in which
the ridges are usually more developed and somewhat angular.

757

rounded off behind, not obliquely truncate, the occipital condyles
barely visible when skull is viewed from above ; paroccipital
processes slender and inconspicuous, rather closely applied to
surface of bullae, their tips descending below lip of foramen
magnum, the ridges extending upward from their bases slightly
developed, much obscured by the somewhat unusual inflation of
petrous portion of squamosal ; floor of brain-case nearly smooth,
but with slightly indicated median ridge ; diameter of basioccipital
along suture contained about two and one-half times in median
length ; auditory bullje moderately large, about as in Microtus
nrvnlis, though somewhat less evenly inflated, their anterior
borders encroaching more deeply into ectopterygoid pits. Inter-
orbital region short, abruptly though not deeply constricted, the
least width usually greater than that of rostrum, the dorsal
surface nearly flat, its outer edges becoming slightly angular in
old age though never developing distinct ridges. Zygomata
weak, scarcely expanded at middle,
rather abruptly though not widely
spreading, the two arches parallel
through a considerable portion of their
extent ; deflection of arches slight,
the median portion lying about at level
of middle of skull ; anteorbital foramen
relatively smaller than in Microtus
arvalis. Rostrum relatively weak, its
least depth behind incisors about equal
to width at same region ; nasals rather
noticeably narrowed behind middle,
their posterior border squarely trun-
cate at about level of middle of zygo-
matic root, the nasal branches of
premaxillaries extending a little further

backward ; incisive foramina scarcely as large as in Mkrotus
arvalis, their posterior extremity at level of front of 7h\ their
anterior extremity separated from incisors by distance equal to
about twice their greatest combined breadth. Palate relatively
somewhat wider than in Microtus arvalis, its surface less sharply
sculptured ; grooves moderately well-defined, but lateral pits
usually rather shallow, their width slightly exceeded by the
short, flattened median ridge ; pterygoids nearly parallel, the
hamulars slightly turned outward ; mesopterygoid space rounded
anteriorly ; ectopterygoid pits relatively smaller than in Microtus
arvalis. Mandible slender, with weak articular process, its
general form essentially as in 31. arvalis.

Teeth. — Upper incisors strongly curved, the roots forming
slight protuberances in infraorbital foramina, the anterior portion
nearly perpendicular, so that anterior surface is barely visible
when skull is viewed from above ; cross section of shaft obscurely
triangular, the anterior border longest, slightly concave near

Fig. 155.
Pitymys subterraneus.

758

RODENTIA

middle, the postero-external border shortest, the inner border
nearly as long as anterior ; enamel extending over entire anterior
border and outer fourth of inner border. Lower incisors slender,
their roots extending well into bases of articular processes but
not forming noticeable protuberances on outer surface of man-
dible ; cross section of shaft differing from that of upper tooth
in the more triangular general outline and
relatively shorter, strongly convex antei'ior
border. Molars so like those of Microtus
arvalis that aside from the peculiarities of
the first mandibular tooth there is nothing to
distinguish with certainty between the two
patterns. Crown of third upper molar distinctly
longer than that of second. In Pitymys suh-
terraneus there is, however, a tendency for the
re-entrant angles to be wider and tlae closed
triangles relatively smaller, while the pattern of
neMs-^^EiiameT" "*^ appears to be more subject to distortion,
pattern. X 5. First outer triangle of m^ not infrequently
opening into inner triangle, but its apex always
on level with that of anterior loop and second triangle. Second
upper molar strictly as in M. arvalis. First lower molar
like that of Microtus arvalis except that third inner re-entrant
angle fails to penetrate to enamel wall of opposite side of tooth,
thus having a broad area of communication between first inner
and first outer triangles ; no exception to this condition has
come under my observation.

BemarJi's. — Although bearing a strong superficial resemblance
to Microtus arvalis, and more particularly to the Alpine M.
incertus, this species is distinguishable externally by its rather
smaller ears, and by the number of mammae and plantar tubercles.
In colour it is usually a clearer, less yellowish brown than the
smaller forms of Microtus. Old and faded museum specimens are
often impossible to determine with certainty. The skull is
recognizable by its small size, nearly flat dorsal profile, wide
interorbital region, and by the almost perpendicular upper
incisors. Two geographical forms are known.

Pitymys subterraneus subterraneus de Selys-Longchamps.

1836. Arvicola subterraneus de Selys-Longchamps, Essai Monographique
sur les Campagnols des environs de Li6ge, p. 10 (Waremme, Li6ge,
Belgium).

1845. H'ly'puclfeus] rufcsccnte-fuscus Schinz, Synopsis Mamm., ii, p. 240
(Urserenthal, Uri, Switzerland). See Mottaz, Bull. Soc. Zool. de
France, xx, p. 27, September, 1907.

1845. Hlypudn'us'] rufofuscus Schinz, Synopsis Mamm., ii, p. 240 (synomym
of rufescente-fiiscus).

1857. Arvicola subterraneus Blasius, Saugethiere Deutschlands, p. 388.

PITYMYS

759

I'JOO. [Arvicola agrestis] fusca Fatio, Revue Suisse de Zool., vin, p. 472
(Untervats, Grisons, Switzerland). See Mottaz, Bull. Sou. Zool.
de Geneve, i, p. 159, November 15, 1908.

1907. Pitymys suhtcrrancus ]Mottaz, Mem. See. Zool. de Franco, xx, p. 27,
September, 1907.

1910. Pitymys suhtcrrancus Trouessart, Faunc Mamm. d'Europo, p. 185.

Ttjiie locality. — Waremme, Liege, Belgium.

Geographical distrihuiion. — From Belgium and northern
France eastward through Switzerland to Transylvania. Limits
of range not well understood.

Diagnosis. — Colour dark and clear ; brain-case showing no
marked tendency to become unusually broad and flat.

Measurements. — Adult male from the type locality : head and
body, 98; tail, 31; hind foot, 14-6; ear, 8. Adult male and
female from St. Cergues, Yaud, Switzerland : head and body,
105 and 101 ; tail, 31 and 32 ; hind foot, 15 and 15-4. Average
and extremes of six adults from Andermatt, Uri, Switzerland :
head and body, 97 (94-100) ; tail, 37*3 (35-39) ; hind foot, 14*8
(14-6-15). Adult male from St. Moritz, Grisons: head and
body, 94; tail, 31 '4; hind foot, 14'4; ear, 9. For cranial
measurements see Table, p. 761.

Specimens examined. — Seventy-one, from the following localities : —

Belgium : Waremme, Li6ge, 13 (B.M. and U.S.N.M.) ; no exact
locality, 1 (Lataste).

France: Near Boulogne, Pas-de-Calais, 3; Dinan, C6tes-du-Nord, 1 ;
Etupes, Doubs, 2 (B.M. and Mottaz) ; Les Lignerets, Orne, 1 (Mottaz).

Switzerland : Near St. Cergues, Vaud, 4 (U.S.N.M. and Mottaz) ;
Bioux, Vaud, 1 (Mottaz); Sengloz, Vaud, 1 (U.S.N.M.); Vidy, near
Lausanne, Vaud, 1 (Geneva) ; Les Plans, Vaud, 1 (Geneva) ; Andermatt,
Uri, 20 (U.S.N.M. and Mottaz) ; Schellenen, near Goschouen, Uri, 6
(Mottaz) ; Furka Pass, Uri, 1 (Mottaz) ; IMurgsee Region, St. Gallon, 8
(U.S.N.M. and Mottaz); St. Moritz, Grisons, 2 (Geneva); Poschiavo,
Grisons, 1 (Mottaz) ; Untervats, above Chur, Grisons, 1 (St. Gallen ; type
ot fusca Fatio).

Austria-Hungary : Hatszeg, Hunyad, Hungary, 8.

4 & 2 al. Liege, Belgium. Baron E. de Selys- 37. 1. 3. 170-171,

Longchamps (c & p). 174-175, 177-178.
4. Li^ge. {de Selys- G. R. Waterhouse (p). 44.10.4.15-18.
Longchcmips).
2 6,19. Boulogne, Pas -de- 0. Thomas (c & p). 98. 1. 9. 21-23.
Calais, France.
6. Etupes, Doubs. {C. 0. Thomas (p). 8. 8. 10. 130.

Mottaz).
1 6,19. Hatszeg, Hunyad, C. G. Danford (c). 3. 11. 8. 44-51.

Transylvania.

760

RODENTIA

PiTYMYS SUBTERRANEUS CAPUCINUS Miller.

1908. Pitymys subterraneus cajmcimcs Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 202, February, 1908.
1910. Pitymys subterraneus cainicinus Trouessart, Faune Mamm. d'Europo,

p. 185.

Type locality. — Spruce forest near " Salon du Capucin,"
Mont-Dore, Puy-de-Dome, France. Altitude about 4000 feet.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — Similar to P. subterraneus subterraneus, but colour
not so dark, and skull with brain-case broader and more
flattened.

Colour. — General hue of upper parts not so dark as in
P. subterraneus subterraneus, the exact shade near the mars-
brown of Ridgway but with an evident bufly cast ; feet a clear,
very pale smoke-grey.

Skull and teeth. — In all general features the skull agrees
with th^t of P. subterraneus subterraneus, but the brain-case is
noticeably broader and more flattened, its length to back of
interparietal about equal to breadth over base of zygomata, its
lateral rounding off much less evident, and the occipital profile
wider in proportion to its depth. Teeth as in the typical form.

Measurements. — Type (adult female) : head and body, 102 ;
tail, 33; hind foot, 15. Adult male from the type locality:
head and body, 100; hind foot, 15. For cranial measurements
see Table, p. 761.

Specimens examined. — Two, both from Mont-Dore.

?. Mont-Dore, Puy-de-D6me, France. G. S. Miller (c). 8. 8. 4. 266-267.

(8. 8. 4. 267. Type of subspecies.)

PITYMYS DACIUS Miller.

1903. Pitymys dacius ]Miller, Ann. and Mag. Nat. Hist., 8th ser., i, p. 202,

February, 1908.
1910. Pitymys dac'ms Trouessart, Faune Mamm. d'Europo, p. 188.

Type locality. — Gageni, Prahova, Roumania (at foot of Car-
pathians, north-west of Bucharest), Roumania.

Geographical distribution. — Known at present from the type
locality only.

Diagnosis. — Similar to Pitymys subterraneus but skull larger,
the nasals more strongly bent downward anteriorly, the brain-
case more depressed posteriorly ; auditory bullae larger ; posterior
vipper molar with third inner re-entrant angle very shallow.

Colour. — The colour of the type is indistinguishable from that
of typical P. subterraneus.

Skull. — In general the skull resembles that of Pitymys sub-
terraneus, but the brain-case is more depressed posteriorly and

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762

RODENTIA

the nasals are more bent downward in front, the dorsal profile
being thus made faintly convex throughout. Though even more
flattened than in P. subterraneus cajmcinus, the brain-case is as
long relatively to its breadth as in true subterraneus. Auditory
bvdlte larger than in P. subterraneus though not peculiar in form.

Teeth. — The teeth resemble those of Pitymys subterraneus,
except that the third inner re-entrant angle of m^ is so shallow
that the terminal loop is essentially straight, its main axis
directed backward and slightly outward, its inner border with a
short, blunt, projecting point somewhat behind middle.

Measurements. — Type (adult female) : head and body, 88 ;
tail, 32*5; hind foot, 14' 5; ear, 8 '5. For cranial measure-
ments see Table, p. 761.

Spccimeii examijied. ^Tho typo.

9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 65.
{W. Dodson.) (Type of species.)

PITYMYS DRUENTIUS Miller.

1852. Arvicola (Microtus) sehjsii Gerbe, Rov. et Mag. de ZooL, 2d ser., iv,
p. 159, March, 1852 (Near Barcelonnette, Basses-Alpes, France).
Not Arvicola sclysii Bonaparte, 1845.

1911. Pitymys druentius Miller, Proc. Biol. Soc, Washington, xxiv, j). 39,
February 24, 1911 (Substitute for selysii).

Tyye locality. — Terres-plaines, near Barcelonnette, Basses-
Alpes, France.

Geographical distribution. — South-western Alps ; limits of
range not known.

Diagnosis. — In general similar to Pityvi'/s subterraneus, but
colour differing in a strong bufFy suffusion of the entire pelage,
particularly noticeable on ventral surface ; feet greyish white in
evident contrast with colour of back.

Colour. — Upper parts a light wood-brown, sometimes tinged
with raw-umber, faintly grizzled with greyish and black ; on
sides the wood-brown becomes lighter and more buffy, passing
rather abruptly into colour of belly ; underparts light ochraceous-
buff irregularly darkened by the appearance at surface of slaty
under-colour, particularly on chin and throat ; feet scantily
clothed with silvery greyish white hairs, the dorsal surface
of foot strongly contrasted with back ; tail obscurely bicolor,
brownish above, whitish below.

Skull and teeth.^The skull and teeth resemble those of
Pitymys subterraneus, but the dorsal profile of the brain-case is
somewhat less flattened, and the molars are usually more robust,
particularly the posterior maxillary tooth. The terminal loop of
this tooth shows a tendency to assume the short, abruptly
rounded form characteristic of P. multiplex. Inner border of
posterior triangle of m"^ with a tendency to develop an evident

PITYMYS 763

convexity, the first stage in the formation of a postero-interual
loop. Such an incipient loop rarely if ever occurs in P. subter-
raneus, though it is common and not infrequently well developed
in P. fatioi and P. multiplex.

Measurements. — Average and extremes of six adults from the
type locality: head and body, 101 '2 (96-105); tail, 33 -3
(31-35); hind foot, 14-9 (14-4-15); ear, 9-0 (8-5-9-3). For
cranial measurements see Table, p. 766.

Sj)ccimcns examined. — Eleven, from the following localities in France :
neighbourhood of Barcelonuette, Basses-Alpcs, 9 (B.M. and U.S.N.M.) ;
Alps of department of Var, 1 (U.S.N.M.) ; no exact locality, 1.

3. Basses-Alpes, France. Purchased (Parzu- 52. 5. 27. 53-55.

daki).
6,3 9. Barcelonuette, Basses-Alpes. 0. Thomas (r). 8.8.10.126-120.

(C. Mottaz.)

PITYMYS FATIOI Mottaz.

1909. ntjjmys vmUiplex fatioi Mottaz, Bull. Soc. Zool. de Geneve, i,

p. 180, January 15, 1909. Type in Mottaz collection.

1910. Pitymys multiplex fatioi Trouessart, Faune Mamm. d'Europe, p. 189.

Type locality. — Zermatt, Valais, Switzerland.

Geographical distribution. — At present known only from the
neighbourhood of Zermatt.

Diagnosis. — Similar to Pitymys druentius but larger (hind foot,
15 to 16 mm., condylobasal length of skull about 23 mm.) ;
brain-case less deepened ; convexity of dorsal profile very slight ;
auditory bullse not specially inflated, scarcely rising to level of
cutting surface of molars.

Colour. — The colour does not differ appreciably from that of
Pitymys selysii.

Measurements. — Average and extremes of seven adults from
the type locality: head and body, 99-7 (95-104); tail, 35
(30-39) ; hind foot, 15-4 (15-16) ; ear, 9-4 (9-10). For cranial
measurements see Table, p. 766.

Specimens exam,ined. — Twenty-six, all from the neighbourhood of
Zermatt (U.S.N.M., Mottaz and Geneva).

Bemarlcs. — Though specimens of this animal were included
by Fatio under the name multiplex, the Zermatt Pitymys is,
as pointed out by Mottaz, easily distinguishable from the form
inhabiting northern Italy and the Italian slope of the Alps.
The relationships seem in fact to be much more intimate with
the western Alpine P. druentius; but more extensive material
than that now available will be required before the exact status
of the members of this group can be properly understood.

764

KODENTIA

PITYMYS MULTIPLEX Fatio.

1905. [Arvicola'] multiplex Fatio, Arch. Sci. Phys. et Nat., Geneve, 4tli ser.,

XIX, p. 193, February 15, 1905 (Lugano, Ticino, Switzerland).

1906. M[icrottis'] leponticus Thomas, Ann. and Mag. Nat. Hist., 7th ser.,

XVII, p. 419, April, 1906 (Locarno, Ticino, Switzerland). Type in

British Museum.
1908. PiUjmys multiplex Mottaz, Bull. Soc. Zool. do Geneve, i, p. 105

November 15, 1908.
1910. Pitymys multiplex Trouessart, Fauue ^Mamm. d'Europe, p. 189.

Type locality. — Lugano, Ticino, Switzerland.

Geographical distribution. — Italian Switzerland and northern
Italy, south to Florence ; limits of range not known.

Diagnosis. — Similar to Pitymys druentiris and P. fatioi but
decidedly larger (hind foot usually 16 to 17 mm., condylobasal
length of skull, 24 to 25 mm.) ; skull with dorsal profile notice-
ably convex ; auditory bullae not specially inflated.

External characters. — Ears relatively smaller than in Pitymys
snhterraneus ; feet less slender, and both palmar and plantar
tubercles sensibly reduced, their relative size approaching that
in the members of the ibericus group.

Colour. — The colour so nearly resembles that of Pitymys
subterraneus as to need no detailed description. The dark wash
along back tends to be rather extensive, and the entire upper
surface is frequently suffused with a light bister or raw-umber.
Feet usually with less dusky clouding than in P. subterraneus.

SJiull. — As compared with that of Pitymys subterraneus the
skull is throughout larger and more heavily built, though the
surface remains equally smooth, and the parietal and temporal
ridges are never more than barely indicated. Dorsal profile
slightly but evidently convex throughout. Brain-case larger
and less depressed than in P. subterraneus, the postorbital ridges
rather well developed, sometimes with a slightly indicated angular
projection, which, however, never becomes so distinct as in the
small species of Microtus. Length of brain-case to back of
interparietal about equal to vvddth over roots of zygomata.
Occiput squarely truncate posteriorly, the condyles nearly or
entirely hidden when skull is viewed from above. Auditory
bullae of the same relative size as in P. subterraneus, but with
smoother, more evenly inflated finish.

Teeth. — The teeth resemble in general those of Pitymys sub-
terraneus, but the third upper molar is not so long (though still
sensibly exceeding the middle tooth) and the posterior portion of
its terminal loop is distinctly reduced, the dentine space scarcely
as large as that in the first oviter triangle, and the incurved
enamel portion very short and abruptly hooked, thus distorting
the crescentic form of the loop as a whole. Posterior outer
triangle usually opening rather widely into terminal loop, though
sometimes completely isolated, the exact opposite to the condi-

765

Fig. 157.

Pitymys iiiultiplex.

Enamel pattevii. X 5.

; ear, 10. Adult

tion in P. suhferraneus. Second upper molar showing the
maximum of the tendency common to the three related
forms, (Iruentius, fatioi and multiplex, for the inner side of third
triangle to develop an imperfect postero-
internal loop. At least some trace of this
tendency may be detected in nearly every
specimen, though it is often present in the
tooth of one side only. In the most extreme
examples that I have seen the supplemental
loop is as large as in many specimens of
Microtus agrestis, though in none is it com-
pletely isolated from the outer triangle.

Measurements. — Adult male from Gor-
dola, near Locarno, Ticino, Switzerland
(type of lepontii'us Thomas) : head and body,
95 ; tail, 38 ; hind foot, 16 "8 ; ear, 10. Adult
female from Locarno, Ticino, Switzerland :
head and body, 103 ; tail, 38 ; hind foot, 16 -7
male from Bellinzona, Ticino, Switzerland : head and body, 100 ;
tail, 31 ; hind foot, 17. Adult male from Mirabello, Mon-
ferrato, Italy: head and body, 102; tail, 33; hind foot, 16 '5;
ear, 9. Adult female from Prato, near Genoa, Italy : head and
body, 102 ; tail, 35 ; hind foot, 16. Adult female from Borzoli,
Genoa, Italy: head and body, 105; tail, 27; hind foot, 16;
ear, 10. Adult from Vaccarezza, Italy : head and body, 110 ;
tail, 33; hind foot, 16 "5. For cranial measurements see Table,
p. 767.

Specimens examined. — Thirty-nine, from the following localities : —
Switzerland : Near Locarno, Ticino, 5 (including type of leponticus) ;
Bellinzona, Ticino, 1 ; Comano, Ticino, 4 (B.M. and Mottaz) ; near
Lugano, Ticino, 12 (B.M. and Mottaz) ; Pico INIagadino, Ticino, 1
(Ghidini).

Italy : Porlezza, Como, 3 ; Domodossola, Novara, 1 ; Ceresolo d'Alba,
Turin, 1 (Turin) ; ]\Iirabello, Monferrato, Alessandria, 1 (Turin); Vaccarezza,
6 (Genoa) ; Caorsi, Autola, 2 ; Prato, Genoa, 1 (U.S.N.M.) ; Borzoli,
Genoa, 1; Florence, 1 (U.S.N.M.).

Remarks. — This species was first distinguished by Forsyth
Major, who, however, omitted to publish its description. Using
the name leponticus intended for it by Major, Thomas in 1906
referred to it in his description of the Asia Minor P. majori, and
thereby became responsible for the name. But in the mean-
time it had been described by Fatio with a mixture of other
forms, its type locality and history being afterwards cleared up
in the paper by Mottaz above referred to.

2 6,2 9. Locarno, Ticino, Switzer- 0. Thomas (c & p). 5. 8. 2. 11-14.
land. (5. 8. 2. 11. Type of P. leponticus Thomas.)

9. Borzoli, Genoa, Italy. Marquis G. Doria (p). 8. 7. 18. 7.

766

RODENTIA

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RODENTIA

PITYMYS SAVII de Selys-Loiigchamps.

1838. Arvicola savii de S^lys-Longchamps, Revue Zoologiquc, p. 248,

October, 1838.
1845. A[rvicola'] selysii Bonaparte, Atti della sesta Eiunione degli Scien-

ziati Italian!, Milano, 1844, p. 350 (synonym of savii ; Pecchioli

cited as autliority).
1857. Arvicola savii Blasius, Siiugethiere Doutschlands, p. 394.
1908. Pitymys savii Miller, Ann. and Mag. Nat. Hist., 8th ser., i, p. 205,

February, 1908.
1910. Pitymys savii Trouessart, Faune Mamm. d'Europe, p. 187.

Type locality. — Neighbourhood of Pisa, Italy.
Geographical distrihution. — Peninsula of Italy from Piedmont
southward ; apparently occurring for the most part in the plains
and lower country.

Diagnosis. — Size and form essentially as in Pitymys subter-
raneus, but feet more robust, ear relatively smaller, and palmar
and plantar tubercles more reduced ; skull usually more heavily
built, much less depressed, the dorsal profile slightly convex, the
general appearance much as in P. multiplex, except that brain-
case is shallower, less elongated and more sub-quadrate ; third
upper molar about as long as second, normally with three closed
triangles, its posterior loop simple, without inner re-entrant
angle, its main axis directed backward and slightly outward.

Colour. — The colour resembles that of Pitymys suhterraneus
and P. multiplex, but is paler, and seldom, if ever, a clear hair-
brown. Upper parts suffused with a light indefinite buffy or
very pale wood-brown. Feet whitish grey as in P. multiplex,
none of the specimens examined showing any trace of the
dusky suffusion characteristic of P. suhterraneus.

Skull. — As compared with that
of P. suhterraneus the skull is broader,
more angular (parietal and temporal
ridges usually evident, though low)
and less depressed, with slightly
convex dorsal outline, relatively
shorter rostrum, and more squarish,
less elongated brain-case (length to
back of interparietal slightly less
than breadth over roots of zygo-
mata). The z^^gomata are shorter
and more widely projecting. Brain-
case squarely truncate posteriorly
as in the related species. The
palatal aspect of the skull shows no
special peculiarities. The skull is
smaller than that of P. multiplex,
less elongate. The rostrum is also

FIG. 158.
Pitymys savii. Nat. size.

and the brain- case
relatively shorter.

769

Pitymyn sanli.
Enamel pattern. X 5.

Teeth. — Except for the form of the third upper molar the
teeth show no pecuharities. This tooth is so shortened that the
length of its crown does not exceed that of second molar. It
consists of the usual elements,- except that the
third inner re-entrant angle is lacking and the
rather short terminal portion ends posteriorly
in a simple elliptical or slightly ovate loop, the
main axis of which is directed backward and
a little outward. Outer re-entrant angles well
developed, sub-equal, the posterior slightly the
larger, the second space usually communicating
with posterior loop as in P. multiplex, both
isolated or not from large inner triangle.
Second upper molar as in P. suhterraneiis, the
third triangle not normally showing any trace
of an incipient postero-internal l5op.

Measurements. — Adult male from Gozzano,
Novara, Italy : head and body, 99 ; tail, 27 ; hind foot, 15. Two
adult males from Ceresole d'Alba, Turin, Italy : head and body,

96 and 96; tail, 25 and 25; hind foot, 14-8 and 15-4. Two
adult males from Mirabello, Monferrato, Italy : head and body,

97 and 99; tail, 24 and 25; hind foot, 16 and 16-5; ear, 9
and 9. Two adult males from Frugarolo, Alessandria : head and
body, 102 and 105 ; tail, 24 and 28 ; hind foot, 15 and 15; ear,
9 and 9. Adult male from Parma, Italy : head and body, 99 ;
tail, 29; hind foot, 16; ear, 9. For cranial measurements see
Table, p. 774.

Specimens examined. — Ninety-nine, from the following localities : — ■
Italy : Gozzano, Novara, 1 (Genoa) ; ]\Iilan, 1 ; St. Angelo, Pavia, 3 ;
Serravalle, Piedmont, 1 ; Ceresole d'Alba, Turin, 5 (Turin) ; Frugarolo,
Alessandria, 21 (B.M., Genoa, and Mottaz) ; Mirabello, JMonferrato, Ales-
sandria, 2 (Turin) ; Caorsi, Mount Antola, 3 ; Modena, 3 ; Pisa, 1 (Mottaz) ;
Florence, 29 (U.S.N.M. and Mottaz); Urbino, Marche, 3; Benagna,
Perugia, 1 (Genoa) ; Ostia, Rome, 1 ; Chieuti, Foggia, 24 (B.M. and Genoa).

1.

Milan, Italy.

6,2 9 al.

St. Angelo, Pavia.

?al.

Serravalle, Piedmont,

2 (J, 2 ?, 2 al.

Frugarolo, Alessan-

dria.

6.

Mirabello, Alessan-

dria.

6,2 9 al.

Caorsi, Liguria.

<5, ?, ? juv al.

Modena.

2 9, juv al.

Urbino, Marche.

1.

Ostia, Rome.

2 6,9.

Chieuti, Foggia.

Baron E. de Selys-

Longchamps (p).
Marquis G. Doria (p).
Marquis G. Doria (p).
jMarquis G. Doria (p).

Dr. E. Festa (p).

Marquis G. Doria (p).
INIarquis G. Doria (p).
jMarquis G. Doria (p).
Dr. L. Sambon (c & p).

Genoa Museum (e).

45. 7. 5. 10.

90. 4. 7. 3-5.
90. 3. 5. 10.

7. 1. 9. 10-15.

8. 8. 1. 5.

90. 3. 5. 11-13.
90. 3. 5. 4-6.
90. 3. 5. 7- 9.
1. 1. 2. 10.

(8. 7. 18. 6.
18. 8. 2. 1-2.

6 D

Y70

HODEN'rlA

PITYMY9 NEBRODENSIS Milia-Paluinbo.

18G8. Arvicola iiebrodcnsis Mina-Palumbo, Ann. Agric. Sicil., XM, p. 6k
1910. Pltymys savii ncbrodcnsis Trouessart, Faune Mamm. d'Europa,
p. 187.

Tyipe locality. — Le Madonie, Sicily-.

Geogra])liical distribution. — Sicily.

Diagnosis. — Similar to Pitymys samii, >biit sfeiill wittli deeper,
•more elongate, less flattened brain-case, and more evidently
■convex dorsal profile ; mesopterygoid space relatively wider than
in P. sami.

Colour. — The colour does not differ appreciably from that of
the related species.

Shull. — The general form of the skull as viewed from above
is much as in Pitymys savii ; length of brain-case about equal to
breadth over roots of zygomata. In lateral and posterior view,
however, it is seen to be decidedly deeper, more as in ordinary
members of the ibericus group.

Teeth. — The teeth are essentially as in Pitymys savii, though
the posterior upper molar is usually not so long owing to the
excessive shortening of the posterior loop. In one specimen*
among the ten skulls examined, the first outer triangle is
opposite the inner triangle, the two together forming a single
transverse loop as in the ibericus group. The length of the outer
triangle is, however, not reduced, so that the resemblance to
ibericus is only superficial.

Measurements. — Adult male from Castelbuono, Sicily : head
and body, 100; tail, 29; hind foot, 16; ear, 10. For cranial
measurements see Table, p. 774.

Specimens examined. — Nine, all from the island of Sicily (B.M. and
U.S.N.M.).

6, 9. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 16-17.

3 6,19. Castelbuono. {A. Robert.) 0. Thomas (p). !«" q' i' 96-27'

6 al. Madonie. Marquis G. Doria (p). 90. 3. 5. 14.

PITYMYS PYRENAICUS de Selys-Longchamps.
(Synonymy under subspecies.)

Geographical distribution. — Pyrenees (known from the French
side only) and region to the north as far as the Garonne. Limits
of range not known.

Diagnosis. — Related to Pitymys savii and P. nebrodensis ;
general form of skull somewhat intermediate between that of the
Italian and Sicilian species, but -mesopterygoid fossa decidedly
narrower.

* i. No. 98. 10. 6. 16, Palermo, Sicily.

tXtYMYS • 771

Shill. — The skull has iuuch the same general outline as in
P. nehrodensis, that is, the brain-case is slightly longer than
broad and the dorsal profile is slightly convex, but the depth is
about as in P. savii and P. Diultiijlex. It is readily distinguished
from that of P. suhferraiieus by its much less flattened brain-case
and by the evidently convex dorsal profile. Palate with lateral
pits unusually shallow and ill-defined.

Teeth. — As in Piti/mys savii. In one otherwise normal
specimen* the terminal loop of the last upper molar is cut by a
deep inner re-entrant angle.

PiTYMYS PYRENAicus PYRENAicus de Selys-Longchamps.

1847. A[rvicola'] pyrenaicus de Selys-Longchamps, Revue Zoologique,

p. 305, October, 1847.
1908. Pitymys pyrenaicus pyrenaicus Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 203, February, 1908.
1910. Pitymys pyrenaiczis Trouessart, Faune Mamm. d'Europe, p. 189.

Type locality. — Bagneres de Bigorre, Hautes- Pyrenees,
Prance.

Geographical distribution. — Pyrenees (not known from the
Spanish side).

Diagnosis. — General colour above a nearly clear hair-brown
as in P. suhterraneus.

Measurements. — Adult female from I'Hospitalet, Ariege : head
and body, 104 ; tail, 34 ; hind foot, 16 ; ear, 8. Adult male and
female from Ax-les-Thermes, Ariege : head and body, 101 and
97 ; tail, 33 • 4 and 29 • 6 ; hind foot, 15 • 8 and 15 • 4. Two adult
males from Bareges, Hautes- Pyrenees : head and body, 94 and
96 ; tail, 32 and 32 ; hind foot, 15*8 and 16 ; ear, 8-6 and 8-2.
For cranial measurements see Table, p. 775.

Specimens exanmied. — Thirteen, from the following localities : —
France : L'Hospitalet, Ariege, 2 ; Ax-les-Thermes, Ariege, 3 ; Luchon,

Haute-Garonne, 1 ; Bareges, Hautes-Pyr6n6es, 5 ; Pyrenees, no exact

locality, 2.

?. L'Hospitalet, Ariege, France. G. S. Miller (c). 8. 8. 4. 272.

2 6. Ax-les-Thermes, Ariege. G. S. Miller (c). 8. 8. 4. 268-269.

6. Luchon, Haute-Garonne. 0. Thomas (p). 6. 4. 1. 78.
{A. Robert.)

2 6. Bareges, Hautes-Pyren^es. G. S. Miller (c). 8. 8. 4. 270-271.

1. Pyrenees. Baron E. de S61ys- 45. 7. 5. 6.

Longchamps (p).

1. Pyrenees. Purchased (Parzudaki). 56. 3. 12. 6.

<^. No. 8. 8. 4. 271, Bareges, Hautes-Pyren6es, Prance.

3 D 2

772

RODENTIA

PiTYMYS PYRENAICUS BRUNNEUS Miller.

1908. Pitijmys piircnaicus brunneus INIiller, Ann. and Mag. Nat. Hist.,
8th ser., i, p. 203, February, 1908. Tyi^e in British Museum.

1910. Pitymys injre7iaicus brunneus Trouessart, Faune Manim. d'Europe,
p. 189.

Tijiie locality. — Forest of Bouconne, Gers, France.

Geographical distribution. — Lowlands of south-western France
from near base of Pyrenees to the Garonne. Limits of distribu-
tion not known.

Diagnosis. — Like Pitymys pyrenaicus pyrenaiciis, but colour of
upper parts with a decided cast of buffy or pale wood-brown,
much as in P. savii.

Measurements. — Type (adult female) : head and body, 93 ;
tail, 26 ; hind foot, 15 ; ear, 8. Average and extremes of five
adults from the type locality : head and body, 96 (93-104) ; tail,
25-6 (23-29); hind foot, 15-4 (15-16); ear, 7-7 (7-9). For
cranial measurements see Table, p. 775.

Spechnens examined. — Fifteen, from the following localities in south-
western France : Forest of Bouconne, Gers, 14 ; Cadillac, Gironde, 1
(U.S.N.M.).

4 (5, 3 ?. Foret de Bouconne, Gers, 0. Thomas (p). 6. 4. 1. 79-85.

France. {A. Bobert.) (6. 4. 1. 82. T^/pe of subspecies.)

PITYMYS PLANICEPS Miller.

1908. Pitymys planiccps Miller, Ann. and ]\Iag. Nat. Hist., 8th ser., i,

p. 203, February, 1908. Type in Lataste Collection.
1910. Pitymys planiceps Trouessart, Faune Mamm. d'Europe, p. 190.

Type locality. — Bareges, Hautes-Pyrenees, France.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — Size about as in Pitymys subterraneus (condylo-
basal length of skull, 23 mm.) ; skull more flattened and depressed
than in any other known European member of the genus, the
dorsal profile nearly straight from posterior extremity of nasals
to back of interparietal ; teeth as in P. savii and P. pyrenaicus.
External characters not known.

Skull and teeth. — In general outline as viewed from above
the skull resembles that of Pitymys savii in its rather short
rostrum, broadly spreading zygomatic arches and squarish brain-
case (length to back of interparietal equal to breadth over
zygomatic roots). Laterally and posteriorly it suggests P. stib-
terraneus, but with the normal characters of the species carried
to the extreme. The dorsal profile is essentially sti'aight from
posterior extremity of nasals to back of interparietal, though
there is a faint concavity in interorbital region and a slight
convexity at middle of brain-case. Interorbital region about as

PITYMYS 773

in P. savil, the tempoi-;il ridges wide apart and barely per-
ceptible. Auditory bull;? rather larger than usual in P. mhter-
raneus and more smoothly inflated. As compared with that of
P. pyrenaicus the skull is immediately recognizable by its flat
dorsal profile and short, broad, excessively flattened brain-case.
Teeth as in Piti/mijs pyrenaicus.

Measurements. — External measurements not known. For
ci'anial measurements see Table, p. 775.

Specimen examined. — The type.

PITYMYS GERBII Gerbe.

1879. Arvicola (Microtus) gerbii Gerbe, Le Naturaliste, i, p. 51, July 1,

1879.

1880. Arvicola (Microtus) gerbei Gerbe, Bull. Soc. Zool. de France, Paris,

V, p. 49, pi. IV, figs. 1-9.
1910. Pitymys gerbei Trouessart, Faune Mamm. d'Europe, p. 186.

Type locality. — Dreneuf, Loire-Inferieure, France.

Geographical distribution. — Known only from the type locality.

Diagnosis. — In general like Pitymys savii and P. pyrenaicus,
but skull somewhat heavier, and interparietal lozenge-shaped,
narrowed at outer extremity to an acute point which usually
does not come in contact with temporal ; m^ with crown
decidedly shorter than that of m^, its first outer triangle opening
broadly into inner triangle.

E.rternal characters and colour. — Apparently about as in
P. pyrenaicus, so far as can be determined from a somewhat
faded mounted specimen.

Shdl. — The skull dift'ers from that of Pitymys pyrenaicus in
greater relative depth and breadth throughout, shortei', heavier
rostrum, shorter incisive foramina, wider interorbital region,
and in the very peculiar form of the interparietal, as already
described ; nasals conspicuously angular-emarginate posteriorly ;
auditory bullae rather large.

Teeth. — The teeth do not differ from those of Pitymys
pyrenaicus except that the third upper molar is shorter and its
first outer triangle opens broadly into inner triangle, a condition
more usual in members of the ibericus group.

Measurements. — The following measurements are given by
Gerbe : head and body, 94-95 ; tail, 27-28 ; hind foot (c. u.) 17.
For cranial measurements see Table, p. 775.

S'pecimen examined. — One from Dreneuf (paratypc) in Paris Museum.

PemarJcs. — This species is well characterized by the great
general breadth of skull, unusually wide interorbital region, and
very peculiar interparietal. The last character might be sup-
posed to be an accidental variation or abormality were it not

774

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776 RODENTIA

present in the Paris specimen* as well as in the eleven skulls
examined by Gerbe. No approach to this form of interparietal
has been seen in other members of the savii group, though it is
sometimes approximated in ibericns and its allies. The depth
of skull relatively to its length is intermediate between that of
P. pyrenaicus and P. ibericus.

PITYMYS LUSITANICUS Gerbe.

1879. Arvicola (Microtus) htsitanicus Gerbe, Rev. et Mag. de Zool., 3rd ser.,

VII, p. 44.
1905. Microtus {Pitymys) htsitanicus Major, Ann. and Mag. Nat. Hist.,

7th ser., xv, p. 512, May, 1905.
1908. Pljtyrmjs'] kisitanicus Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

"p. 204, February, 1908.
1910. Pitymys lusitanicus Trouessart, Faune Mamm. d'Europe, p. 193.

Ty^je local! t II . — Portugal.

Geograplncal distrihution. — Portugal ; limits of range un-
known.

Dicujnosis. — Size small (hind foot, 14" 5 to 15-5 mm. ; condy-
lobasal length of skull, 22*6 to 23 mm.); skull slender and
lightly built, not depressed, the dorsal profile convex through-
out ; brain-case high and rounded, its depth, including auditory
bulla, about 74 per cent of occipital breadth ; third upper molar
scarcely as long as second, normally without closed triangles, its
posterior loop short, simple, without inner re-entrant angle,
directed backward and slightly outward ; upper incisors rather
strongly projecting ; colour dark.

Colour. — The colour is essentially as in Pitymys suhterraneus,
except that the undei'parts are a less clear grey. Upper
parts a dark hair-brown tinged with bister or sepia, and very
faintly grizzled by black tips and light reflections. Underparts
a dull bufFy grey, in some specimens becoming almost a dull
ochraceous-bufi' on cheeks and on sides along boi'der of dark area ;
feet buffy grey without dusky wash ; tail obscurely bicolor, bufFy
white below, dusky mixed with buffy white above.

Skull. — The skull very closely resembles that of Pitymys
nehrodensis. Dorsal profile convex throughout, or somewhat
flattened in interorbital region. Interorbital constriction wider
than anterior portion of rostrum, flat or slightly concave laterally,
the outer edges marked by faint ridges in extreme old age, these
ridges rarely extended back across parietals. Brain-case high,
strongly rounded off at sides, its length to back of interparietal
about equal to breadth over z3'gomatic roots. Posteriorly the

* Though presented by Gerbe this specimen is not likely to be one of
those whose cranial peculiarities were mentioned by the describer of the
species. It was originally mounted with the skull inside, and the nose
and upper lips were found to be attached to the rostrum when the skull
was removed for my examination, through Dr. Trouessart's kindness, in
1908.

PITYMYS 777

brain-case is less squarely truncate than in P. nehrodensls or
P. subterraneus, so that the condyles are usually visible when
skull is viewed from above. Auditory bullte rather small, their
surface smoothly rounded. Palate with lateral pits large and
sharply defined, noticeably wider than median ridge.

Teeth.* — Though resembling in a general way those of Pitymys
savii the teeth are at once recognizable by the size and position
of the anterior outer triangle of m^. This triangle is so reduced
in size that it is smaller than the succeeding triangle, and its
point does not reach the level of a line joining the outer
extremity of the anterior loop with tip of second outer triangle.
In position the first outer triangle is opposite the large inner
triangle, the two communicating freely by their bases, and
together forming a single transverse loop varying somewhat in
form according to the completeness of fusing of the two triangles.
The second outer triangle never communicates with the loop
thus formed, but usually opens into the tei-minal loop as in
P. savii. Terminal loop short and broad, without trace of inner
re-entrant angle, its main axis directed backward and slightly
outward. The length of the third tooth is noticeably less than
that of the second, and the shortness of the first outer triangle
often causes the crown to assume a peculiar outward-bowed
general outline that is highly characteristic of the members of
the group. Second upper molar as in P. subterraneus, the third
triangle without trace of postero-internal loop. Upper incisors
slightly projecting, most of their anterior surface visible when
skull is viewed from above, but not sufficiently thrown forward
to be a conspicuous feature. Other teeth without special
peculiarities.

Measurements. — Average and extremes of seven adults from
Cintra, Portugal: head and body, 93-2 (90-98); tail, 23-4
(21-27); hind foot, 15 (14-5-15'7); ear, 8-3 (8-8-6). For
cranial measurements see Table, p. 788.

Specimens examined. — Eighteen, all from Cintra, Portugal.
16 7 ?, 2 al. Cintra, Portugal. 0. Thomas (c & i'). 98. 2. 2. 37-52.

PITYMYS MARI^ Major.

1905. Microtus {Pitymya) maritv ]\Iajor, Ann. and Mag. Nat. Hist., 7th ser.,
XV, p. 515, May, 1905. Type in British Museum.

1908. P[itymy$] mariee Miller, Ann. and Mag. Nat. Hist., 8th ser., i,p. 204,
February, 1908.

1910. Pitymys marin- Trouessart, Faune ]\Iamm. d'Europe, p. 192.

Tyj)e locality. — Villalba, Lugo, Galicia, Spain.
Geograph'eaJ ilintrihiition. — Known at present from the type
locality only.

* Essentially similar to those of P. ibericns, fig. 1G2 (p. 781).

778 RODENTIA

Diagnosis. — Similar to Pifi/)nys lusitanicus, but skull with
narrower, lower brain-case ; upper incisors so slightly projecting
as to be scarcely visible when skull is viewed from above.

Colour. — The colour is probabl}" similar to that of Pifyriujs
lusitanicus, though in the only known specimens, skinned after
immersion in alcohol, it has a peculiar, perhaps unnatural
russet cast.

Skull. — General outline of skull when viewed from above
essentially as in Pitymys subterraneus, the length of brain-case to
back of interparietal evidently more than width over zygomatic
roots. Whole skull more depressed than that of P. lusitanicus,
so that dorsal profile is much less convex, and general outline
viewed from the side is less strongly cuneate. Auditory bullse
and structure of palate as in P. lusitanicus.

Teeth. — As in Pitymys lusitanicus, but upper incisors nearly
vertical, so that part of the anterior surface of teeth is invisible
when skull is viewed from above.

Measurements. — Type (adult female) : head and body, 79 ;
tail, 30; hind foot, 13' 6; ear, 76. Not fully adult male from
the type locality: head and body, 84; tail, 25; hind foot, 14;
ear, 7*5. For cranial measurements see Table, p. 788.

Specimens examined. — Eight, all from north-western Spain.

2 6,6 al. Yillalba, Lugo, N.W. Dr. V. L. Seoane (c & p). 94. 1. 1. 16-17.
Spain. (94. 1. 1. 16. Type of species.)

PITYMYS PELANDONIUS Miller.

1908. Pitymys pelandonius Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 204, February, 1908. Type iu British JMuseum.
1910. Pitymys pelandonius Trouessart, Faune Mamm. d'Europe, p. 192.

Type locality. — Silos, Province of Burgos, Spain. Altitude
about 3000 feet.

Geographical distribution. — Not known at present elsewhere
than in the neighbourhood of the type locality.*

Diagnosis.- — Similar to Pitymys marise Major, but skull with
broader, less elongated brain-case.

Colour. — The colour of the four skins examined resembles
that of the lighter, more wood-brown specimens of P. lusitanicus.

Shull and teeth. — The skull differs from that of both Pitymys
lusitanicus and P. marise in its broader more quadrate bx"ain-case.
Posteriorly the brain-case is rather low, as in P. marise, though
much less reduced in depth than that of P. dcpressus. Teeth as
in P. marise, the upper incisors nearly vertical.

Pleasure mcnts. — Type (adult female) : head and body, 96 ;
tail, 28 ; hind foot, 14*4 ; ear, 8*4. Adult female from Castrillo

* Occurs among brush and shrubbery, avoiding the open fields
inhabited by P. ibericus.

PITYMYS 779

de la Reina, Burf^os : head and body, 92; tail, 22*4; hind
foot, 14-4; ear, 8. For cranial measurements see Table,
p. 788.

Specimens examined. — Four,- from the following localities in Spain :
Silos, Burgos, 3 (B.M. and Mottaz) ; Castrillo de la Reina, Burgos, 1.

2 9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 294-295.

(8. 8. 4. 294. Type of species.)

PITYMYS DEPRESSUS Miller.

1908. Pitymys depressus Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 204, February, 1908. Type in British Museum.
1910. Pitymys depressus Trouessart, Faune Maram. d'Europe, p. 193.

Tiji^j locality. — Rascafria, south side of Sierra de Guadarrama,
Province of Madrid, Spain.

Geographical distrihution. — Centi'al Spain ; at present known
from the region of the Sierra de Guadarrama only.

Diagnosis. — A small animal related to Pitymys marise and
P. pelandonius, but immediately recognizable by its broadened,
much flattened skull, and very small auditory bulhe.

Colour. — The colour is apparently similar to that of Pitymys
lusitanicus and P. pelandonius, but the only known specimens
have been preserved in alcohol and may not now be in normal
condition.

Skull. — The skull differs conspicuously from that of the
other known members of the ihericus group, all of which,
P. marise and P. pjelandonius not excepted, are characterized by
depth of brain-case, in the much flattened
general form, the outline, viewed from
the side, not very unlike that of P. savii.
Dorsal profile slightly but evidently and
very evenly convex, the nasals not so
abruptly sloping as usual. Rostrum
excessively shallow immediately behind
incisors, its least vertical depth barely
exceeding greatest combined breadth of
nasals. Posterior termination of nasals
straight or slightly eniarginate, exceeded
by nasal branch of premaxillary to extent
of nearly 1 mm. Interorbital region
about as wide as anterior portion of
rostrum, faintly concave laterally, the
temporal ridges evident in adults, though
very low and always remaining wide
apart. Zygomata rather noticeably expanded at middle, pro-
jecting conspicuitu.sly beyond level of sides of brain-case. Length
of brain-case to posterior edge of interparietal less than lireadth
over roots of zygomata, the general outline sub-circular ; occipital

Fig. 160.

Pitijmys depressus.

Nat. size.

780

RODENTIA

region obliquely truncate posterioi-ly, so that condyles are plainly
visible when skull is viewed from above, very low and wide in
posterior aspect ; postorbital ridges low but evident, slightly
angled ; temporal ridges continued back across frontal and
parietal to outer edge of interparietal, abruptly angled at middle
of parietal. Auditory bulhe very small and low, their greatest
diameter contained about four times in coudylobasal length of
skull (about 3 to 3h times in lusitanicus, marise, and pelandonius),
their surface rather smoothly inflated. Palate with well-defined
grooves asd rather shallow lateral pits exceeding in breadth
the flattened median ridge.

Teeth. — The teeth show no peculiarities as compared with
those of other members of the group.

Measuremenis. — Type (adult female) : head and body, 85 ;
tail, 25; hind foot, 13; ear, 8. Two adult females from the
type locality : head and body, 85 and 93 ; tail, 29 and 23 ; hind
foot, 14 and 14; ear, 7*5 and 8. For cranial measurements see
Table, p. 788.

Spcchnens examined. — Thirteen, from the following localities in Spain :
La Granja, Segovia, 6 ; Rascafria, Madrid, 6 ; Villalba, Madrid, 1.

6 al. La Granja, Segovia, Spain. M. de la Escalera (c). 8. 7. 30. 9-14.
? & 5 al. Rascafria, Madrid. M. de la Escalera (c). 6. 11. 4. 10-15.

(6. 11. 4. 15. Type of sj^ecies.)
9. ViUalba, Madrid. 0. Thomas (p). 8. 2. 9. 212.

(N. Gonzalez).

PITYMYS IBERICUS Gerbe.
(Synonymy under subspecies.)

Geograioliical distribution. — Probably the entire Iberian Penin-
sula, though at present known only from central and southern
Spain.

Diagnosis.- — Larger than Pitymys lusitanicus and its allies
(hind foot, 15 to 18-4 mm., condylobasal length of skull, 24 to
25 mm. or more), and adaptation to underground life more
complete ; skull with brain-case less evidently parallel-sided, its
outline in many individuals almost sub-orbicular ; upper incisors
long, conspicuously protruding, nearly their entire front face
visible when skull is viewed from above ; colour usually rather
pale, ranging from hair-brown to a light bufly drab. Habits
strictly subterranean.

External characters. — General form diflfering rather con-
spicuously from that of Pityvms subferraneus and Microtus arvalis
in its much more evident adaptation to underground life, the
head large, with incisors noticeably protruding from mouth,
the neck short and thick ; eai's much reduced, almost hidden in
the fur ; feet strong and robust, the hind foot shortened and
broadened, the digits and claws of both front and hind feet

781

thickened, palmar and plantar tubercles somewhat reduced in
size, crowded into a relatively smaller area than in P. siihterraneus ;
tail short, scarcely more than one-quarter as long as head
and body.

Colour. — Upper parts a peculiar drabby grey, varying
considerably in exact shade as described under the subspecies,
the sides often with a noticeable bufFy tinge ; underparts and
feet whitish in rather marked contrast with back ; tail whitish,
the upper surface usually clouded with brown.

Skull. — -The skull resembles that of Pitymys lusitanicus, but
its general size is greater, the zygomata are more abruptly
spreading, the brain-case is relatively
shorter and deeper, and the occiput
is more obliquely truncate, so that
condyles are more plainly visible
when skull is viewed from above.
Depth of rostrum immediately be-
hind incisors about equal to width
in same region. Interorbital con-
striction about as wide as anterior
portion of rostrum, its dorsal surface
convex laterally ; temporal ridges
low and indistinct, scarcely visible
except in rather old individuals,
sometimes rather closely approach-
ing each other, though without
coalescing, and seldom if ever
extending back over parietals.
Length of brain-case about equal

to or slightly less than width across zygomatic roots, the outline
when viewed from above sub-quadrate or occasionally almost
circular ; postorbital ridges low but attaining
a fair degree of development in old age,
never, however, strongly angled. Auditory
bullie well developed, smooth, their greatest
diameter contained about three and a half
times in condylobasal length of skull. Palate
showing no special peculiarities. Incisive
foramina as usual in the genus, their longi-
tudinal diameter about equal to one-half
distance "from gnathion to alveolus of first
molar.

Teeth. — The teeth of Pitymys ihericus so
exactly resemble those of P. lusitanicus as
to need no detailed description. They are,
however, larger throughout, and the upper incisors project more
strongly forward, .so that practically the entire front face is
visible when skull is viewed from above.

Fig. 101.
Pitymys ibericus. Nat. size.

FIG. 162.

Pitymys ibericus.
Enamel pattern. X

782

PiTYMYS iBERicus iBERicus Gerbe.

1854. Arvicola ibericiis Gcrbe, Revue Zoologique, 1854, p. 400. Type in

Paris Museum.
1879. Arvicola (Microtus) ibcricus Gerbe, Rev. et Mag. de Zool., 3rd ser.,

VII, p. 42.
1905. Microtus ibcricus Major, Anu. and Mag. Nat. Hist., 7th ser., xv,

p. 515, May, 1905.
1908. Pitymys ibcricus ibcricus Miller, Ann. and ]\Iag. Nat. Hist., 8th ser.,

I, p. 205, February, 1908.
1910. Pitymys ibcricus Trouessart, Faune IMamni. d'Europe, p. 190.

Tyj^e locality. — Province of Murcia, Spain.

Geographical distribution. — Coast region of south-eastern Spain
(Provinces of Alicante, Murcia and eastern portion of Granada).

Diagnosis. — Size large (hind foot, usually 17 "6 to 18-4 mm. ;
upper tooth-row, 6 to 6 "4 mm.) ; general colour very pale.

Colour. — Upper parts ranging from a pale hair-brown slightl}^
tinged with bister along middle of back and suffused with buffy
along sides to a light, buffy broccoli-brown, brightening nearly to
cream-buff on flanks and on sides of neck. Underparts usually
greyish white dulled by the slaty undercolour, the general effect
not far from the grey No. 9 of Ridgway, but occasionally washed
with buffy. Pect clear greyish white. Tail whitish with a
varying amount of dark bi-own along median dorsal region, this
brown occasionally almost absent.

Measurements. — Type (adult) : head and body, 114 (skin) ;
tail, 26; hind foot, 18. Adult female from Venta del Baul,
Granada, Spain: head and body, 107; tail, 27; hind foot, 18;
ear, 8. Three adult females from Elche, Alicante, Spain : head
and body, 101, 105 and 106; tail, 27, 27 and 29; hind foot,
17*6, 18 and 18*4. For cranial measurements see Table, p. 789.

Specimens examined. — Eleven, from the following localities in Spain :
Venta del Baul, Granada, 3 ; Murcia, no exact locality, 1 (Paris ; type) ;
Elche, Alicante, 7.

6, 2 ?. Elche, Alicante, Spain. 0. Thomas (p). 8. 2. 9. 209-211.
{N. Gonzalez.)

3 9. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 273-275.

6, ?. Venta del Baul, Granada. G. S. Miller (c). 8. 8. 4. 276-277.

Pitymys ibericus centralis Miller.

1908. Pitymys ibericus centralis Miller, Ann. and Mag. Nat. Hist., Sth ser.,

I, p. 205, February, 1908. Type in British Museum.
1910. Pitymys ibericus centralis Trouessart, Faune Mamm. d'Europe, p. 190.

Type locality. — Pastures near Silos, Province of Burgos, Spain.
Altitude about 3000 feet.

Geographical distribution. — Central Spain from the province
of Burgos south to Valencia and Seville ; exact limits of range
not known.

PITTMYS 783

Diagnosis. — Not so lai-ge as P. ibcricus ibericus (hind foot,
16 to 17*2 instead of 17 • 6 to 18*4) ; colour hair-brown usually
tinged with bufly, never with the paUid tints characteristic of
the typical race ; feet whitish in rather noticeable contrast with
back.'

Measurements. — Type (adult male) : head and body, 102 ;
tail, 24 ; hind foot, 16-8 ; ear, 8. Average and extremes of ten
adults from the type locality : head and body, 99 (95-103) ;
tail, 25-9 (24-28); hind foot, 16-6 (16-17-2); ear, 8-4 (8-9).
Adult male from Catarroja, Valencia : head and body, 100 ;
tail, 21 ; hind foot, 16 ; ear, 9. Three adults from Seville :
hind foot (dry), 16, 16 and 16 '6. For cranial measurements
see Table, p. 789.

Specimens examined. — Sixty, from the following localities in Spain :
Silos, Burgos, 27 ; near Burgos, 3 ; Catarroja, Valencia, 1 ; near Seville, 22
(B.M. and U.S.N.M.) ; Jerez de la Frontera, Cadiz, 5 (perhaps referable
to P. ibericus regulus) ; Coto Dofiana, Huelva, 2.

8 6,2 9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 278-287.

(8. 8. 4. 278. Ty2)e of subspecies.)
N. Gonzalez (c). 8. 7. 7. 45.
0. Thomas (p). 8. 2. 9. 208.

Lord Lilford (p). 95. 3. 3. 29-40.
Abel Chapman (c 0. 5. 10. 2-3.

&p).
A. Trevor Battye 7. 9. 12. 1-2.

(c & p).
Abel Chapman (c 8. 3. 26. 6.

&p).
Abel Chapman (c 8. 3, 8. 10-11.

&p).

PiTYMYS IBERICUS PAscuus Miller.

1908. Pitymys ibericus fuscus Miller, Ann. and Mag. Nat. Hist., 8th scr.,
I, p. 206, February, 1908. 'Not Arvicola agresfis fusca (= Pitymys
subterraneus) Fatio, 1900. Type in British Museum.

1911. Pitymys ibericus pascuus Miller, Proc. Biol. Soc, Washington, xxiv,
p. 39, February 24, 1911 (Substitute iov fuscus).

Type locality. — Dehesa de Valencia, Province of Valencia,
Spain.

Geographical distribution.— AT^jparently confined to the Dehesa
de Valencia.

Diagnosis. — Size as in Pitymys ibericus centralis, but colour
dark, resembhng that of P. lusitanicus and P. pelandonius.

Colour. — Upper parts a uniform bister, nearly as dark as that
of Ridgway, faintly varied by blackish hair tips, and becoming
tinged with wood-brown on sides. Underparts dull slaty grey,
washed with bufFy. Feet whitish in rather marked contrast with
body. Tail very obscurely bicolor, whitish except median dorsal
region, which is tinged to a varying degree with dark brown.

lal.

Burgos.

cJ.

Catarroja, Valencia.

{N. Gonzalez.)

6 6,4:9,2 juv.

Seville. {A. Ruiz.)

2al.

Jerez de la Frontera,

Cadiz.

2al.

Jerez de la Frontera,

Cadiz.

1.

Jerez de la Frontera,

Cadiz.

2.

Coto Dofiana, Huelva,

784 RODENTIA

Skull and teeth. — As in P. ibericus centralis.

Measurements. — Type (adult male) : head and body, 104 ;
tail, 22 ; hind foot, 17 ; ear, 10. Adult male from the type
locality: head and body, 107; tail, 27; hind foot, 17. For
cranial measurements see Table, p. 789.

Specimens examined. — Two, both from the Dehesa de Valencia.

2 6. Dehesa de Valencia, Spain. O. Thomas (p). 8. 2. 9. 213-214.
{N. Gonzalez.) (8. 2. 9. 214. Type of subspecies.)

PiTYMYS IBERICUS REGULUS Miller.

1908. Pitymys ihericus reguhis Miller, Ann. and IMag. Nat. Hist., 8th ser.,

I, p. 206, February, 190S. Tj^pe in British Museum.

1909. [Pitf/mys] ilbericjis] regnlarls Lydeliker, Zoological Record, xlv

(1908), Mamm., p. 75, December, 1909. (Accidental renaming of
regjilus.)

1910. Pityynys ihericus regulus Trouessart, Faune Mamm. d'Europe, p. 191.

Type locality. — North slope of the Alhambra hill, Granada,
Spain.

Geographical distribution. — Known at present from Granada
and Malaga, Spain.

Diagnosis. — General size rather less than in the other forms
of Pitymys ibericus (hind foot, 15 to 16 mm.); skull with small
auditory bullae ; teeth disproportionately reduced (length of tooth-
row about 5*4 mm.) ; colour essentially as in P. ibericus centralis.

Measurements. — Type (adult female) : head and body, 100 ;
tail, 26; hind foot, 16; ear, 9. Adult female from the type
locality: head and body, 97; tail, 21*4; hind foot, 15*4;
ear, 8. For cranial measurements see Table, p. 789.

Specimens examined. — Eight, from the following localities in Spain :
Granada, 7 ; Malaga, 1.

2 6, i 9. Granada, Spain. G. S. Miller (c). 8. 8. 4. 288-293.

(8. 8. 4. 290. I'ype of subspecies.)
1 al. Malaga. J. G. Haggard (p). 8. 4. 28. 1.

PITYMYS DUODECIMCOSTATUS de Selys-Longchamps.

1839. Arvicola diiodccimcostatus de S61ys-Longchamps, Revue Zoologique,

p. 8, January, 1839.
1905. Microtiis duodecimcostatus Major, Ann. and Mag. Nat. Hist., 7th ser.,

XV, p. 508.
1908. Pitymys duodecimcostatus Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 204, February, 1908.
1910. Pitymys duodecimcostatus Trouessart, Faune Mamm. d'Europe,

p. 192.

Type locality. — Geneva, Switzerland? In the original descrip-
tion it is merely stated that the type was received from Professor
Pictet de la Rive, of Geneva.'"' Later de Selys wrote of the

* " Je dois le squelette que je possede a la gen^rosite de M. le professeur
Pictet de la Rive (de Geneve)."

PITYMYS 785

animal as occurring at Geneva.* All recent attempts to redis-
cover the species in Switzerland have, however, led thus far to
no result. A specimen in the United States National Museum
labeled Geneva, Switzerland, was received in 1857 from the
late H. de Saussure.

Geographical distribution. — Mediterranean coast region of
France ; northward to western Switzerland 1 Limits of range
unknown.

Diagnosis. — Similar to Pitymys ibericns, but skull with occiput
more obliquely truncate, so that condyles are plainly visible
when skull is viewed from above ; interparietal narrowly ligulate
instead of broadly ligulate or lozenge-shaped.

Colour. — As in P. ibericns centralis.

Shi II and teeth. — Except for the peculiarities already noted
the skull so closely resembles that of Pitymys ibericus as to need
no detailed description. Teeth as in P. ibericus.

Measurements. — Four adults from the vicinity of Marseilles,
France (skins): tail, 25, 26, 26 and 27; hind foot, 16-4, 16-6,
16*6 and 16*8. For cranial measurements see Table, p. 790.

1. No locality. Purchased (Parzudaki). 52. 5. 27. 56.

PITYMYS PROVINCIALIS Miller.

1909. Pitymys provincialis Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

p. 420, May, 1909. Type in British Museum.

1910. Pitymys provincialis Trouessart, Faune Mamm. d'Europe, p. 191.

Type locality. — Saint-Gilles, Gard, France.

Geographical distribution. — At present known from a few
localities in southern France (departments of Gard and Var).

Diagnosis. — Size small (hind foot about 14' 6 mm. ; condylo-
basal length of skull, 22-6 to 23 mm. ; upper tooth-row, 5'0 to
5*2 mm.) ; skull essentially as in Pitymys duodecimcosiatus, except
for its much smaller size ; auditory bulhe very small and flat.

Colour. — Upper parts a light wood-brown, becoming paler and
more cream-buff on sides ; underparts a light grey formed by the
blending of slate-grey under colour with creamy white of hair-
tips ; feet soiled whitish ; tail whitish throughout, the upper
surface sprinkled with brown hairs. Young ecru-drab faintly
tinged with wood-brown, the underparts and feet dull greyish.

Skull arid teeth. — The skull of Pitymys provincialis is imme-
diately distinguishable from that of Pitymys duodecimcostatus,
its only near geographical ally, by its exceedingly small size, a
character in which it essentially agrees with P. lusitanicus and
the related small species of the Iberian Peninsula. From
these it differs in the conspicuously projecting upper incisors
and the very narrow interorbital region. Brain-case unusually
deep relatively to length of skull, its outline when viewed from

* Etudes de Micromammalogie, pp. 110, 111.

3 E

786 RODENTIA

above sub-orbicular, the sides strongly rounded ofi'. Occiput not
so obliquely truncate as in P. duodecimcostatus, and interparietal
less narrowly ligulate ; in both characters it more closely resembles
P. ihericus. Rostrum very slender, though perhaps not dispro-
portionately so. Auditory bullse smaller and more flattened than
in any other member of the group except P. dejiressus of central
Spain, their contrast with the large bullae of P. duodecimcostatus
particularly noticeable. Upper incisors very strongly projecting,
practically the entire front surface of the teeth visible when
skull is viewed from above. Molars showing no peculiarities of
enamel pattern, but smaller and weaker than in any other
member of the group, the small Iberian species not excepted.

Measurements. — External measurements of type (adult female) :
head and bod}^, 96; tail, 22; hind foot, 14* 6. For cranial
measurements see Table, p. 790.

Speciviens examined. — Eleven, from the following localities in France :
St. Gilles, Gard, 5 ; Province of Var, 1 ; no exact locality (" Provence "), 4
(B.M. and U.S.N.M.).

2 6,2 9. St. Gilles, Gard, G. S. Miller (c). 8. 8. 4. 262-265.

Prance. (8. 8. 4. 265. Type of species.)

1. Provence. Baron E. de S61ys-Long- 45. 7. 5. 9.

champs (p).

2. Var Purchased (Parzudaki). 52. 5. 27. 51-52.

PITYMYS THOMASI Barrett-Hamilton.

1903. Microtus (Pitymys) thomasi Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., xi, p. 306, March, 1903- Type in British Museum.
1905. Microtus [Pitymys) tJiomasi Major, Ann. and Mag. Nat. Hist.,

7th ser., xv, p. 515, May, 1905.
1910. Pitymys thomasi Trouessart, Fauna Mamm. d'Europe, p. 188.

Type locality.- — Vranici, Montenegro.

Geographical distribution. — Known only from Montenegro.

Diagnosis. — Size about the maximum for a European species
of Pitymys (hind foot about 17 "4 mm., condylobasal length of
skull about 27 mm.). Cranial and dental characters essentially
as in Pitymys ihericus. Colour dark, as in P. lusitanicus.

Colour. — Upper parts a dark hair-brown, tinged with bister
along back and with wood-brown on sides ; underparts slaty
grey washed with bufFy ; feet dull whitish ; tail obscurely bicolor,
the median dorsal area brownish, the rest nearly white.

Shdl. — Except in its larger size the skull does not differ
very appreciably from that of true Microtus ihericus. The dorsal
profile is, however, slightly more convex, owing to the greater
depth of the anterior portion of brain-case, the general outline
of the brain-case tends more to assume a nearly circular form,
and the auditory bullae appear to be relatively larger.

Teeth. — As in P. ihericus ihericus.

Measurements. — Type (adult female): head and body, 118;
tail, 22; hind foot (dry), 17-4; ear, 10. Adult male: head

PITYMYS 787

and body, 107; tail, 23; hind foot, 17; ear, 10. For cranial
measurements see Table, p. 790.

Specimens examined. — Four, from the following localities in IMonte-
negro : Beri, 1 ; Daljans, 1 ; Vranici, 2.

RemarJcs. — Among tlie members of the group to which it
belongs this species is easily recognizable by its dark colour and
large size.

6. Beri, Montenegro. {L. Fiihrer.) 0. Thomas (p). 5. 8. 4. 23.
?. Daljans. (L. Fiihrer.) O. Thomas (p). 5. 8. 4. 25.

6, 9. Vranici. (L. Fiihrer.) 0. Tliomas (p). 5. 8. 4. 24, 26.

(5. 8. 4. 26. Type of species.)

PITYMYS ATTICUS Miller.

1910. Pityviys atticus Miller, Ann. and Mag. Nat. Hist., 8th ser., vi,
p. 460, November, 1910. Type in British Museum.

Type locality. — Kephissia, near Athens, Greece.

Geographical distrihii;tion.—ls.nown only from the type locality.

Diagnosii<. — Size much smaller than in the only other known
form occurring in the Balkan Peninsula, P. ihomasi, and about
equal to that of P. dnodecimcostatus. Differs from the latter in
less projecting upper incisors and less obliquel}^ truncate occiput.

Colour. — Upper parts bufTy wood-brown very inconspicuously
lined with black, the sides a little paler and approaching a dull
ochraceous-buff behind ear, on flanks and across rump. There
is no true line of demarcation along sides, but transition to
buffy white of underparts is rather abrupt. Basal portion of
fur everywhere slate-colour, this appearing at surface throughout
underparts. Feet bufFy white in strong contrast with back.
Tail obscurely bicolor, pale cream-bufF below, buffy wood-brown
above. Muzzle tinged with hair-brown.

Skull. — The skull of the type is so injured that its exact form
cannot be seen. Apparently it most nearly resembles the skull
of Pitymys duodecimcostatus in size, but differs in the shorter,
broader and deeper I'ostrum and nasals, relatively wider inter-
orbital region, less strongly projecting incisors and less obliquely
truncate occiput. Auditory bullae flatter and less inflated than
in P. duodecimcosfatus, though of about the same diameter (not
reduced as in P. provincialise).

Teeth. — The teeth are essentially like those of Pitymys duode-
cimcostatus and P. ihomasi, but the anterior outer re-entrant
angle of m^ is more pronounced than in any of the skulls of the
two related species with which I have compared it.

Measurements. — Tyj^e (adult female) : head and body, 95 ;
tail, 25; hind foot (dry), 16; ear, 8 '2. For cranial measure-
ments see Table, p. 790.

Specimen examined. — The type.
1. Athens, Greece. {C.Mottaz.) Hon. N. C. Rothschild (p). 8.10.2.51.

(Type of species.)

3 E 2

788

RODENTIA

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791

Hub-Family MURINE.
1857. Murinas Baird, Mamm. N. Amor., p. 434 (part).

Geographical distribution. — Originally confined to the Old
World south of or barely entering the Arctic region, but
members of two genera now everywhere artificially distributed.

Characters. — Molars tuberculate, brachyodont, rooted, the
tubercles of the maxillary teeth arranged in three primary
longitudinal rows ; external form with no special modifications.

Remarhs. — The sub-family Murinse contains about fifty
described genera, five of which occur in Europe.

KEY TO THE EUROPEAN GENERA OF MURIN.^.

Mesopterygoid fossa reduced to a triangular space

scarcely extending in front of hamulars ; coronoid '

process of mandible obsolete ; back densely and

conspicuously spinous (Crete) A.coynys, p. 883.

Mesopterygoid fossa normal, extending decidedly in

front of hamulars, its sides usually parallel or

nearly so ; fur not spinous, though back sometimes

with inconspicuous flattened bristles (Distribution

general).

First and second upper molars with two tubercles on

inner side.

Upper incisor with outer side of cutting edge entire ;

first upper molar 5-rooted, its crown not so

long as that of in- and vi^ combined ; size

large (House-rats) Epimys, p. 848.

Upper incisor with outer side of cutting edge broken

by conspicuous notch and projection ; first upper

molar 3-rooted, its crown longer than that of

VI- and ?«' combined ; size small (House-mice) Mus, p. 863.

First and second upper molars with three tubercles on

inner side when unworn.

Tail not prehensile, completely haired at tip ; ear

without lobe closing meatus ; rostral portion of

skull normally developed, the length of diastema

noticeably greater than depth at front of wi'

(Field-mice) AiMilcmus, p. 791.

Tail prehensile, naked at tip above; ear with con-
spicuous lobe closing meatus ; rostral portion
of skull so shortened that length of diastema is
less than depth at front of m' (Harvest-mice)... Micromys, p. 840.

Genus APODEMUS Kaup.

1829. Apodemus Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt.,

I, p. 150 (agrariua).
1857. Mus Blasius, Siiugethiere Deutschlands, p. 309 (part).
1905. Micromys Thomas, Ann. and Mag. Nat. Hist., 7th ser., xv, p. 492,

May, 1905 (part).
1908. Apodcmus Thomas, Ann. and Mag. Nat. Hist., 8th ser., i, p. 447,

May, 1908 (part).

Type species, — 3Iiis agrarius Pallas.

Geographical distrihution. — Northern temperate portion of

792

RODENTIA

01(1 World from Ireland to Japan, south to nortlicrn India and
the Mediterranean region of Africa.

Characters. — Skull without special modifications of form, the
rostral portion well developed (diastema noticeably exceeding
depth of skull at anterior root of w}), the mesopterygoid fossa
squarish or broadly double- or single-rounded anteriorly ; first
and second upper molars with three cusps on inner side ; ear
without valve for closing meatus ; tail not prehensile ; fur
without spines.

Remarks. — Among PahTearctic Miiridse the genus Apodemus is
well characterized by the complicated pattern of enamel folding,
the non-prehensile tail, and the normal skull. A similar combi-
nation of peculiarities occurs, however, in the African Thamnomys.
About a dozen species are known, seven of which occur in
Europe.

KEY TO THE EUROPEAN FORMS OF APODEMUS.

First lamina of m^ with crescentic form much dis-
torted ; m- with no antero-extenial tubercle ;
iuterorbital region with conspicuously beaded
edges ; back with blacli median line (Eastern

Europe, west to Denmark) A. agrarius, p. 834.

First lamina of to' with crescentic form not dis-
torted ; m^ with small but distinct autero-
external tubercle ; interorbital region with
edges not beaded ; back with no black median
line.
First upper molar with four tubercles on outer
margin of crown ; alveolar width of m^ equal
to about half greatest width of palate between
tooth-rows ; size maximum for the European
members of the genus, the condylobasal length
of skull not infrequently more than 30 mm. ;

colour above huffy grey (Balkan Peninsula)... A. cpimelas, p. 794.

First upper molar with three tubercles on outer

margin of crown ; alveolar width of »i' less

than half greatest width of palate between

tooth-rows ; size never attaining maximum

for European members of the genus, the

condylobasal length of skull seldom attaining

28 mm.

Skull becoming massive and relatively angular

in old age, the brain-case eventually with

low but evident lateral ridges ; size large,

the hind foot most frequently 25 mm. in

length (23-27 mm.) ; condylobasal length

of skull in individuals with noticeably worn

teeth most frequently 26 mm. (25-29 mm.) A. flavicollis, p. 828.
Underparts usually creamy white, the pectoral
spot usually not forming complete collar
and not spreading backward over chest
(Continental Europe north of Mediter-
ranean region) A. f. flavicollis, -p. 829.

Uiiderparts usually greyish white, the pectoral
spot usually forming complete collar and
spreading backward over chest (England) A.f. wintoni, p. 831.

APODEMUS 793

Skull rcinahiiug light and smooth, even in
extreme old age, the brain-case never
developing evident lateral ridges ; size
medium or small, the hind foot most
frequently 22-23, mm. (20-25 mm.) ;* condy-
lobasal length of skull in individuals vpith
noticeably worn teeth most frequently
23-24 mm. (21-26 mm.).*
General form of skull slender, especially in
rostral portion ; naandible with corouoid
process reduced ; colour dusky above,
dull white below, the pectoral spot obso-
lete ; size maximum for this section, the
hind foot and condylobasal length of
skull sometimes exceeding 26 mm. (]j''air

Isle, Scotland) A.fridariensis, p. 825.

General form of skull normal, the coronoid

process of mandible well developed ;

colour reddish, yellowish or pallid above,

underparts whitish or washed with

brown, the pectoral spot usually well

developed ; size moderate or small.

Underparts always heavily washed with

yellowish brown ; ear shortened and

narrowed.

Condylobasal length of skull 24 to 25 mm.

(Hebrides, Scotland) ^4. Jiebridensis, p. 824.

Condylobasal length of skull 25 "6 to

27 mm. (St. Kilda Island, Scotland) A. hirtensis, p. 825.
Underparts usually whitish, occasionally
washed with yellowish brown ; ear

normal A. sylvaticus, p. 797.

Size small, hind foot most frequently

22 mm., condylobasal length of skull
most frequently 23 mm.

General colour above rich yellowish

brown usually with a decided

russet tinge (Europe north of

Mediterranean region) A. s. sylvaticus, p. 803.

General colour above light yellowish

brown without russet (Crete) A. s. creticus, p. 813.

Size medium, hind foot most frequently

23 mm., condylobasal length of skull
most frequently 25 mm.

General colour above rich yellowish

brown usually with a decided

russet tinge (Pyrenees, Asturias

and Portugal) A. s. caUipides,'p. 809.

General colour above pale yellowish

or greyish brown without russet

(Mediterranean region in general) A. s. dichrums, p. 810.

* These dimensions exceeded in the local form inhabiting Fair Isle,
Scotland.

794

EODENTIA

APODEMUS EPIMELAS Nelirinir.

1882. Mus mystaciniis Winge, Vidensk. IMeddel. fra den naturh. Foren. i
Kji>benhavu, 4th ser., iii (1881), p. 21 (Dekelia, near Athens,
Greece). Not Mus mystacinus Danford and Alston.

1902. Mus epimelas Nehring, Sitz.-Ber. Gesellsch. Naturforsch. Freunde,
Berlin, p. 95, February, 1902.

1910. Mus {Epimys) epimelas Trouessart, Faune Mamm. d'Europe, p. 144,

Type locality. — Agoriani, Parnassus, Greece.

Geographical distribution. — Balkan Peninsula, west to Cepha-
lonia, Corfu and Montenegro.

Diagnosis. — Largest European member of the genus ; adults
with hind foot, 26 to 29 mm., condylobasal length of skull, 28 to
31 mm. ; colour light greyish buff with no decided yellowish or
russet tinge, the back noticeably clouded with black ; skull and
teeth essentially as in Apodemus sylvaticus except for their
greater size, but molars relatively larger, the alveolar width of
m^ equal to about one-half width of palate, vi^ and m"^ with
minute though evident fourth tubercle at postero-external
margin of crown, and outer ledge of 7/;^ and w;.^ somewhat better
developed.

External characters. — As in Ap)odemus sylvaticus except for
such differences as are correlated with larger size. Ear large,
completely covering eye when laid forward, its form not
peculiar. Palmar and plantar tubercles apparently larger than
in A, sylvaticus (only dry specimens examined) ; sole completely
bare to heel. Tail about as long as head and body, the
annulations plainly visible, fourteen to the centimeter at middle,
the hairs rather sparse, their length equal to width of about
three rings ; pencil scarcely indicated.

Colour. — Entire upper parts cream-buff with a faint greyish
tinge, everywhere darkened by an admixture of blackish hair-
tips, these inconspicuous on sides of body but numerous enough
over middle of back to produce a noticeable dark clouding ;
cheeks, sides of neck, outer surface of front legs, and region
behind ears nearly clear cream-buff' ; underparts and inner
surface of legs buffy white, the line of demarcation along sides
sharply defined ; feet white ; ear an indefinite dark brown ; tail
sharply bicolor, the hairs on upper surface blackish, those on
lower surface white. In immature individuals the cream-buff is
replaced by smoke-grey, the black remaining essentially as in
the adults.

Skull. — In all essential respects, apart from its conspicuously
greater size, the skull agrees with that of Apjodemus sylva-
ticus. As in the smaller animal the brain-case remains smooth
above, even in extreme old age, never developing the lateral
ridges found in old individuals of A. fiavicollis. Anterior
border of incisive foramina extending relatively further for-

APODEMUS

795

ward than in A. syhat/cus, the distance from foramen to alveolus
of incisor barely more than half combined width of the
two foramina instead of nearly equal to it.

FIG. 163.
Apodemus epimelas. Nat. size.

Teeth. — Though in general resembling those of Ayodemus
syloaticus, the teeth of A. epimelas differ in several important
details. The crowns of all the cheek-teeth are perceptibly.

Fig. 164.
Apodemus epimelas. Cheek-toeth. X 10.

longer proportionately to their breadth than in the smaller
animal, but at the same time the distance between the upper
tooth-rows is so reduced that the alveolar breadth of m^ is fully

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equal to oue-lialf width of palate in same region. Pattern of
enamel folding essentially as in Apodemus sylvaticus, but
re-entrant angles of first lamina of m^ without rudimentary
supplemental loops ; postero-external border of crown of m^ and
m'^ with small though evident supplemental tubercle appearing in
worn teeth as a narrow outward-projecting loop ; outer ledge
and rudimentary tubercles of m^ and w., better developed than in
A. sylvaticus, those of the second tooth nearly as large as those of
first, a condition rarely if ever occurring in the smaller animal.

Measurements. — Adult male from Montenegro : head and
body, 150; tail, 146; hind foot, 26; ear from meatus, 20.
Adult male from northern Albania: head and body, 135;
tail, 115; hind foot, 27; ear from meatus, 19. Adult male
from Corfu, Greece: head and body, 128; tail, 134; hind
foot, 27'5; ear from meatus, 21 "7. Adult female from
Cephalonia, Greece: head and body, 131; tail, 129; hind
foot, 27*4; ear from meatus, 20 • 8. Adult male from the same
locality: tail, 120; hind foot, 29; ear from meatus, 20-8. For
cranial measurements see Table opposite.

Spccimeyis examined. — Thirty-five, from the following localities : —

MoNTENEGKO : No Gxact locality, 6.

Albania : Northern Albania, 8.

Greece : Corfu, 3 ; Cephalonia, 6 ; near Athens, 12.

Bemarhs. — While Apodemus eptmelas is very unlike the other
European members of the genus it closely resembles A. mystaciniis
Danford and Alston from Asia Minor, differing chiefly in its
somewhat larger skull and heavier teeth (see measurements in
Table opposite), and also in the slightly duller, less buffy ground
colour of upper parts.

3 <J, 3 9. Montenegro. (L.Filhrer.) O. Thomas (p). 5. 8. 4. 10-15.

i 6, 4:9. North Albania. 0. Thomas (p). 5. 8. 4. 27-34.

{L. Filhrcr.)

2 6. Corfu, Greece. [G.Mottaz.) J. I. S. Whitaker (p). 8. 10. 1. 28-29.

6, ?. Argostoli, Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 26-27.

(C. Mottaz.)

6,9. Tatoi, Athens. (C. lfo«02.) Hon. N. C. Rothschild 8.10.2.88-39.

APODEMUS SYLVATICUS Linnseus.
(Synonymy under subspecies.)

Geographical distribution. — Central and southern Europe from
Ireland eastward and from the Mediterranean coast north at
least to central Scandinavia ; northern limits of range not known.

Diagnosis. —Size small or medium ; head and body, about
95 to 105 ; tail vertebrjB, about 8 0 to 110; hind foot, 20 to 25,
most frequently 22 or 23 ; condylobasal length of skull, 22 to
26 mm. ; ear large, its height from meatus about 16 to 18 mm.:
general colour of upper parts ranging in different local forms

798

RODENTIA

from a bright russety wood-brown nearly to buff and greyish
drab ; underparts whitish, sometimes a nearly pure bluish or
creamy white, more often tinged with slaty or washed with
yellowish brown, the line of demarcation along sides always
sharply defined in adults ; a yellowish brown spot or longitudinal
streak usually present on middle of chest between fore legs, this
spot rarely spreading to form a complete collar ; teeth rather
weak, the alveolar width of m^ less than half width of palate
between tooth-rows ; m'^ with three distinct tubercles on outer
side, but neither this tooth nor m^ with well-developed
supplemental loop behind third outer tubercle (sometimes a
slight trace of this loop in m^) ; outer side of m^ and m^ with
noticeable cingulum-like ledge extending nearly entire length of
crown and bearing distinct though minute cusps ; skull remaining
light and smooth even in extreme old age, the interorbital
margins not beaded, the brain-case never developing evident
lateral ridges ; rostrum not specially elongated ; mandible with
well developed coronoid process.

External characters. — External form slender, much as in the
house mouse, but feet, eyes and ears i-elatively larger. Ear
extending slightly but evidently beyond eye when laid forward,
its general outline broadly ovate, sometimes a little flattened
above and on posterior border ; antitragus reduced to a low
inconspicuous ridge not capable of closing meatus ; surface of ear
finely pubescent. Muzzle with ill-defined naked area surrounding
nostrils ; nostril opening obliquely forward and outward, wider
anteriorly than posteriorly, its inner and lower margin thickened
and irregularly wrinkled ; a deep furrow under lower border and
another dividing nostrils in median line and continuous below
with cleft in upper lip. Feet slender, without special modifica-
tions. Thumb reduced to a mere tubercle, the upper surface of
which is covered by the rudimentary nail ; palm naked,
5-tuberculate, the tubercles well developed, distinct, the two
hindermost slightly larger than the others, but showing no
tendency to unite along median line, the surface of the palm
between the tubercles irregularly wrinkled ; under surface of
digits with five or six deep transverse furrows. Hind foot
relatively longer and more slender than in the house mouse, the
three median toes sub-equal, the outer extending almost to end of
first phalanx of fourth, the inner scarcely to base of first phalanx
of second, the under surface marked off by conspicuous cross
furrows into well defined, entire, scales or plates, of which there
are seven or eight on the longer toes. Sole naked except at
sides of heel (this hairy area sometimes extending completely
adross), its surface papillose in region among tubercles, rugose at
heel, smooth between last tubercle and wrinkled posterior
portion ; tubercles six, well developed and distinct, ovate in
outline, the four anterior largest and sub-equal, the postero-
external smallest. Tail about as long as head and body.

APODEMUS

799

rings.

somewhat quadrate in section, the annulations distinct, about
eighteen to the centimeter at middle ; hairs rather sj^arse, not
conceaHng the annulations at middle but forming a small though
evident pencil at tip, their length equal to width of 2^
Mammfe : p 1 — 1, i 2 — 2 = 6.

Colour. — For detailed description of colour see accounts of
subspecies.

Skull. — The skull is moderately large, smoothly rounded, and
without any striking characteristics. Dorsal surface straight
and gradually rising from tip of nasals to middle of frontals,
then modez'ately convex to back of interparietal, the convexity
most strongly marked over middle of parietals ; ventral profile
horizontal to back of molars, then bent rather abruptly
downward to lowest portion of bulla.
Brain-case ranging from broadly ovate
to sub-circular in outline when viewed
from above, its surface everywhere
smoothly rounded except for the faintly
angular posterior border to orbits, a
slight perpendicular ridge in mastoid
region, and in very old individuals a
barely indicated lambdoid angularity.
Interparietal rather large but narrow,
its extremities gradually tapering and
usually rather sharp-pointed. Posterior
aspect of occiput low and narrow, much
exceeded in height by the median region
of brain-case ; paroccipital processes small
and inconspicuous, close to outer surface
of condyle. Floor of brain-case with no
conspicuous features, the basioccipital

with low median ridge and shallow lateral excavations ; bulla
moderate, smoothly rounded, the large meatus with faintly
indicated tubular rim. Interorbital region somewhat abruptly
though not very narrowly constricted, flat above, the margins
rounded anteriorly, squarely angled posteriorly but never raised,
and bead-like. Zygoma slender, rather abruptly but not widely
spreading anteriorly, the median portion of the two arches
essentially parallel when viewed from above. Plate forming
outer border of infraorbital canal well developed, its anterior
border perpendicular. Rostrum slender, not elongated, the
nasals simple, narrowing gradually backward, their posterior
termination squarely truncate or bluntly pointed, usually a little
in front of posterior extremity of nasal branches of premaxillaries
and of level of front of lachrymal. Incisive foramina rather
long, parallel-sided, extending from about 1-1 '5 mm. behind
incisors to level of anterior root of j/t' ; palate nearly flat, without
special features, the portion formed by palatine bones extending
forward to middle of in^ ; meeopterygoid space narrow anteriorly.

Fir,. 105.
Apodeinns si/lvaticus.

Nat. size.

800 RODENTIA

wider posteriorly, its anterior border straight or faintly curved,
usually not extending forward quite to level of last molar ;
hamulars slightly thickened at tip, curved a little outward,
barely or not in contact with bullae ; ectopterygoid well
developed, nearly horizontal, its posterior portion spreading
considerably beyond outer alveolar line. Mandible slender, with
no special peculiarities of form ; coronoid process small but well
developed, directed noticeably backward, its extremity about on
level with condyle ; angular process long and i-ather slender, its
inner sui'face with noticeable longitudinal concavity ; dental
foramen at level of alveoli ; projection marking root of incisor
slight but evident, directly under base of coronoid process.

Teetli. — Relatively to size of skull the teeth are rather small :
alveolar breadth of 111^ distinctly less than half greatest width of
palate between tooth-rows ; alveolar length of upper tooth-row a
little less than half diastema. Upper incisor strongly curved.

Fig. 166.
Apodemus sylvatieus. Cheek-teeth, («) slightly worn ; (6) much worn. X 10.

forming nearly half of a circle, its root appearing as an evident
protuberance on side of rostrum slightly in front of lower portion
of infraorbital foramen ; shaft compressed, nearly triangular in
section, its antero-posterior diameter about double its greatest
width, the anterior and inner faces flat, joining each other at a
well marked nearly right angle, the outer edge of anterior face
curving rather broadly into outer face, the curved portion giving
this part of the section a somewhat oblique appearance ; posterior
face narrow, abruptly rounded, not sharply differentiated from
outer and inner faces. Lower incisor slightly longer and much
less curved than upper incisor, its root forming a protuberance
on outer surface of mandible under base of coronoid process and
slightly above alveolar level ; cross-section of shaft essentially
as in upper incisor, but anterior surface relatively narrower.
Upper molars M'ith tubercles arranged according to a ground-
plan of three longitudinal rows of tubercles, those of the median

APODEMUS 801

i-()\v iiuR'li ];irt((^i' than those of the ijinci- and outer rows, tlie
tubercles of the dili'erent rows partly joined by cross-ridges, ajid so
disposed as to form three transverse laminae having somewhat the
form of forwardly-bowed crescents. In each tooth this complete
pattern is only approximated ; in m'^ it is imperfect posteriorly,
in m'^ both anteriorly and posteriorly, while in m^ it is scarcely
more than recognizable. For convenience of description the
tubercles representing the complete pattern may be numbered
transversely 1 to 9, beginning with the first on inner side and
ending with the third on outer side ; those of the inner row
will therefore be 1, 4 and 7, those of the middle row 2, 5 and 8,
those of the outer row 3, 6 and 9. Crown of m^ more than one
and a half times as long as broad, its length nearly and area
fully equal to that of two succeeding teeth combined ; tubercles
of outer row slightly larger than those of inner ; anterior
crescent slightly distorted by the small size oi tl and the very
broad, shallow angle between its inner border and the outer
border oi. t2 ; a small but evident projecting supplemental loop
usually present in angle between to and t2, a similar but less
developed loop occasionally present in that between t2 and tl ;
second crescent essentially similar to first, and partly oi- com-
pletely joined to it by a narrow ridge extending between tS and
the outer base of t5 and occasionally a similar ridge between tl
and the outer base of to, the ridges variable in development
though rather constant in position ; third crescent much distorted
by the pushing forward of i9 and tl so that the three cusps of
this lamina lie in the same transverse line, and the re-entrant
angles separating them from t6 and ti respectively are much
narrower than those behind iirst lamina ; t9 is broadly connected
with tQ anteriorly, so that the two might almost be interpreted
as parts of a deeply divided t6 ; t7 is similarl}^ connected with
ti, and, though well developed, is the most reduced cusp in the
tooth ; postero-external border of crown usually with a faintly
indicated supplemental loop, this never so well developed as in
Ajwdemus epimelas. Second upper molar sub-circular in outline,
its first lamina incomplete by the absence of V2 ; i3 very small,
terete ; second and third lamintB essentially as in m^ ; postero-
external loop excessively minute, though usually present in unworn
teeth. Third upper molar similar to m? in form but scarcely half
as large. In moderately worn condition it usually has the appear-
ance of a sub-circular tooth with two re-entrant angles on inner
side extending about half way across crown and occasionally a
slight notch on outer side near middle ; when unworn it is
possible to identify tubercles 1, 4, 5 and 8, the inner re-entrant
angles corresponding to the spaces between first and second and
second and third lamina;, the notch on outer surface to the
re-entrant angle at front of th. Lower molars consisting of a
series of paired, equal, nearly distinct tubercles, each tooth
terminating posteriorly in a single median tubercle usually much

3 F

cMIJ UODEKTIA

smaller than the others. First lower molar with six paired
tubercles and a median anterior loop nearly as large as median
posterior tubercle. Second lower molar with four paired
tubercles and a small median posterior tubercle. Third molar
with two paired tubercles essentially like those of the other
teeth, but with posterior median tubercle slightly larger than
■either of the paired tubercles. The relative size of the three
teeth is indicated by the number of pairs of tubercles. Outer
margin of w;, and m.^ with a narrow ledge bearing two or three
minute terete cusps. The ledges and cusps are variable in their
degree of development ; usually they are more evident in m^
than in m,, ; in m^^ there are usually two at outer margin of
second outer main tubercle, and a third somewhat better
developed at margin of postero-external main tubercle ; in Wg the
most constant supplemental cusp is that which lies at outer
base of antero-external main tubercle ; behind this there are
usually no distinct supplemental cusps, though one or two are
occasionally present.

Bemarks. — This is the most abundant and most universally
distributed of European mammals. Except in cities, at the
extreme north, on the highest mountains, and perhaps in some
parts of the Mediterranean region,* it is probably everywhere
more numerously represented in individuals than any other
species. Adults are I'eadily distinguishable from the house
mouse by their brighter colour, larger ear and longer hind foot ;
but at all ages and in all conditions of pelage the structure of
the teeth and the absence of supplemental tubercles to all of the
pads on palm and sole differentiate it from all members of the
genus Mus. The only European mammal with which Apodemus
sylvaticus is likely to be confused is A. fiavicollis. Detailed
comparisons of the two animals will be found under the latter
species.

Throughout the greater part of its range, from Ireland to
Roumania, and from northern Scotland and central Scandinavia
to Switzerland and the plains of south-western France, Apodemus
sylvaticus is represented by its typical form, small sized and rich
in colour, the back in adults in fresh pelage usually with at
least a trace of russet. In the Pyrenees, Asturias and Portugal
occurs a larger race, which, however, retains the bright colour
of true sylvaticus. This tendency toward larger size is shown
throughout the Mediterranean region, though a small race
occurs on the island of Crete. Unlike the Pyrenean-Asturian
form the true Mediterranean races are all pale, with yellowish
or greyish tints predominating to the exclusion of russet. Along
the Mediterranean coast of France, and in northern Italy lies

* In the vicinity of Elche, Alicante, Spain, I found AiMclemus sylvaticus
much less numerous than Miis S2}icilegus, and practically confined to
ravines and river banks, while the smaller animal was abundant every-
where.

APODEMUS 803

an area of iiitergradatiou between true t<ylmflrus and tlie larger
Mediterranean form. Hpecimens from this region are indeter-
minate in characters, thougli for the most part apparently
nearest to the southern race. In addition to the race already
alluded to as inhabiting Crete several other insular forms have
become to a greater or less degree differentiated. That from
the island of Corfu appears to be slightly more grey than that
occurring on the mainland of Greece. Specimens from the
Channel Islands and Scilly Islands show a tendency to be larger
than on the neighbouring mainland. But none of these local
races is suthciently distinct to admit of characterization. On
the other hand, the Hebrides, St. Kilda, and Fair Isle, oflf the
coast of Scotland, are each inhabited by a peculiar, well-
differentiated form, sufficiently distinct to take rank as a species.
The few specimens that I have seen from the Shetland and
Orkney Islands appear to be indistinguishable from true
sylvaiiciis. It seems probable, therefore, that they have been
recently introduced.

Apodemus sylvaticus sylvaticus Linua'us.

1758. [Mus] sylvaticus Linnaeus, Syst. Nat., i, 10th ed., p. 62 (Upsala,

Sweden).
1796. Mus sylvaticus parvus Bechsteiu, Getreue Abbild. Naturhist. Gegen-

stande, i, p. 100 (Thiiringen, Germany).
1796. Mas sijlvaticus candidus Bechstein, Getreue Abbild. Naturhist.

Gegenstiinde, i, p. 100 (Thiiringen, Germany).
1796. Mus sylvaticus varius Bechstein, Getreue Abbild. Naturhist. Gegen-

stande, i, p. 100 (Thiiringen, Germany).
1796. Mus sylvaticus niger Bechstein, Getreue Abbild. Naturhist. Gegen-
stiinde, I, p. 100 (Thiiringen, Germany).
1801. Mus syl\vaticus] albus Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2d ed., p. 965 (Thiiringen, Germany).
1801. Mus syUvaticus] leucocephalus Bechstein, Gemeinn. Naturgesch.

Deutschlands, i, 2d ed., p. 966 (Thiiringen, Germany).
1827. Mus islandicus Thienemann, Reise im Norden Europas, ii, p. 153

(Iceland).
1834. Mus sylvaticus Melchior, Den Danske Stats og Norges Pattedvr,

p. 102.
1839. Mus intcrmcdius Bellamy, Nat. Hist, of South Devon, p. 330 (Devon-
port, Devonshire, England).
1857. Mus sylvaticus Blasius, Saugethiere Deutschlands, p. 322 (part).
1900. Mus sylvaticus intermedius Barrett-Hamilton, Proc. Zool. Soc,

London, p. 398.
1900. Mus sylvaticus celticus Barrett-Hamilton, Proc. Zool. Soc, London,

p. 401 (Caragh Lake, Co. Kerry, Ireland).
1910. Mus sylvaticus intermedius and M. sylvaticus celticus Trouessart,

Faune Mamm. d'Europe, pp. 149, 150, 151.
1910. [Apodenms] sylvaticus Miller, Ann. and Mag. Nat. Hist., 8th ser.,

VI, p. 460, November, 1910.

Type locality. — Upsala, Sweden.

Geograjihical distribution. — Central Europe from Ireland

3 F 2

804 RODENTIA

castwurd, ;ui(l fi'oin contral Sweden and Norway soutli to
iiortheT-ii Italy and the soiitliern I'^ranct; ; Iceland (probably
introduced).

Diagnosis. — Size small ; head and body, about 95 ; tail
vertebrae, about 90 ; hind foot, about 22 ; condylobasal length of
skull, 22 to 24 '6 mm. ; general colour of upper parts relatively
dark and rich, the sides and flanks a russety, yellowish, wood-
brown.

Colour. — Ordinary adult specimens in fresh pelage are wood-
brown above, distinctly tinged with russet on posterior half of
back, and becoming paler and more bufty on head, neck,
shoulders and sides ; everywhere there is a noticeable sprinkling
of blackish hairs, these usually producing a decided effect of
" lining," and over middle of posterior half of back a slight
though evident dark clouding ; cheeks, sides of neck, outer
surface of fore legs and a narrow, ill-defined region along hne
of demarcation, dull light buff; uuderparts and dorsal surface of
feet dull white, the throat and belly always clouded by the slate-
grey undercolour, and not infrequently with a slight or decided
buffy wash ; chest usually with a buff median area between fore
legs, this occasionally forming a complete collar or spreading
backward along median line ; ear dull brownish with no decided
markings, the edges usually lighter and often faintly silvery ;
tail bicolor, though in most specimens not conspicuously so, the
upper surface dark brown, the under surface whitish. Immature
individuals are duller than the adults, the upper parts usually
with a plumbeous tinge, the underparts more slaty, so that there
is occasionally no well marked line of demarcation along sides.
Specimens in this pelage often resemble some of the bicolored
forms of the 3Ius miisciiJus group, though they may always be
distinguished by their relatively larger eyes and ears, relatively
longer hind foot, and by the absence of supplementary tubercles
to all of the pads on sole.

Measurements. — Two adult males from Iceland : head and
body, 90 and 95; tail, 87 and 92; hind foot, 22-6 and 23-2;
ear, 18 and 17 '6. Average and extremes of five adults from
Ireland: head and body, 97 (90-103) ; tail, 83-7 (71-97) ; hind
foot, 22-3 (22-23); ear from meatus, 15-5 (15-16). Average
and extremes of ten adults from the Isle of Man : head and
body, 95 (90-103); tail, 84 (75-93); hind foot, 22-2 (21-23);
ear from meatus, 16 (15-17). Average and extremes of five
adults from Cromarty, Scotland : head and body, 95 '6 (92-101) ;
tail, 87-2 (83-91); hind foot, 22-6 (21-23); ear from meatus,
16 "8 (16-17). Average and extremes of ten adults from
Wormsley, Oxfordshire: head and body, 96 "5 (90-105); tail,
90-1 (79-99); hind foot, 22-1 (21-23). Average and extremes
of six adults from Tresco, Scilly Islands, Cornwall : head and
body, 95-5 (92-99); tail, 88 (83-96); hind foot, 22-2 (21-23):
ear from meatus, 15 5 (15-16). Average and extremes of ten

APODEMUS

805

adults from Bergen, Norway: head and body, 96 •! (89-107);
tail, 98-3 (89-107); hind foot, 22-1 (21-22 6). Adult male
and female from Upsala, Sweden : head and body, 95 and 93 ;
tail, 89 and — ; hind foot, 21 and 21. Average and extremes
of ten adults from Waremme, Liege, Belgium : head and body,
94-3 (85-99); tail, 92-3 (85-99); hind foot, 22-0 (21-23).
Average and extremes of ten adults from the Forest of Bouconne,
Gers, France: head and body, 93-1 (87-101); tail, 85 ■ 5
(78-90); hind foot, 20-5 (20-21); ear from meatus, 15-2
C 15-1 6). Average and extremes of ten adults from Brunswick,
Germany: head and body, 94-3 (91-105); tail, 87-1 (81-94);
hind foot, 21 '3 (20-22). Average and extremes of seven adults
from Gageni, Roumania : head and body, 93*0 (90-96); tail,
85-3 (81-91) ; hind foot, 21-5 (21-22); ear from meatus, 16-0
(15-5-17). Average and extremes of ten adults from Meiringen,
Bern, Switzerland: head and body, 96 "0 (93-99); tail, 93-7
(91-107) ; hind foot, 22-2 (21-23-4). Average and extremes of
seven adults from Locarno, Ticino, Switzerland : head and body,
96 (92-101); tail, 97-4 (87-103); hind foot, 22-5 (21-23-6);
ear from meatus, 16 '1 (15-17). For cranial measurements see
Table, p. 814.

Specimens examined. — Eight hundred and twenty-six, from the
following localities : —

Iceland: Near Reykjavik, 6 (U.S.N.M.).

Ireland : Ciandeboye, Down, 3 ; Co. Meath, 1 ; Woodpark, Co. Clare, 5 ;
Scariff, Galway, 3 ; Clare Island, Co. Mayo, 6 ; Inishmore, Aran Islands, 4 ;
Fermoyle, Clare, 4 ; Nenagh, Tipperary, 1 ; Cashel, Tipperary, 2 ; Ballagh-
moon, Carlow, 6; Cappagh, Wexford, 1; Duncannon, Wexford, 3;
Kilmauock, Wexford, 2 ; Caragh Lake, Kerry, 6 (including type of celticus
Barrett-Hamilton); Castlegregory, Kerry, 4; Glencar, Leitrim, 1.

Scotland : Islay, 1 ; Skye, 1 ; Duurossness, Shetland, 3 ; South Sutor,
Cromarty, 7; Black Isle, Cromarty, 4; Craigellachie, Elgin, 1; Lossie-
mouth, Elgin, 2; Lhanbride, Elgin, 2 ; DuniDhail, Elgin, 8; Tiree, 2;
Kinloch. Rannoch, Perthshire, 2 : Cortachy, Forfar, 3 (Wilson) ; Dalmeny
Park, Edinburgh, 2; Beech Hill, Haddington, 1; Stockbriggs, Lanark-
shire, 2 ; Wyseby, Dumfriesshire, 1.

England: Isle of Man, 36; Riding-Mill-on-Tyne, Northumberland, 1;
Alnwick, Northumberland, 3 ; Filey, Yorkshire, 1 ; Grimsby, Lincoln-
shire, 3 ; Scole, Norfolk, 1 ; Cheadle, Staffordshire, 3 ; Swincliffe, Shrop-
shire, 1 ; Swithland, Leicestershire, 12 ; Belvoir, Leicestershire, 2 ; Anglesey,
1 ; Aberia, Merionethshire, 2 ; Truoyn-y-peurleyu, Merionethshire, 2 ;
Penrhyndeudraeth, JMerionethshire, 1 ; Laugharne, Carmarthenshire, 4 ;
Merlins Hill, Pembroke, 2 ; Southerndown near Bridgend, Glamorgan-
shire, 10 ; Bishopstone, Herefordshire, 3 ; Graftonbury, Herefordshire,
14 ; Ledbury, Herefordshire, 1 ; Darsloy, Gloucestershire, 3 ; Clifton,
Gloucestershire, 7 ; Crippetts, Gloucestershire, 11 (Wilson) ; Wyre Forest,
Worcestershire, 1 ; Rugby, Warwickshire, 2 ; Bloxham, Oxfordshire, 2 ;
Wormsley, Oxfordshire, 18 ; Luton, Bedfordshire, 1 ; Oundle, Northampton-
shire, 1 ; Colne, Cambridgeshire, 1 ; Histon, Cambridgeshire, 1 ; Fulbourn,
Cambridgeshire, 3 ; Levesham, Suffolk, 2 (Wilson) ; Lowestoft, Suft'olk, 4 ;
Haileybury, Hertfordshire, 1 ; Tring, Hertfordshire, 1 ; Hillingdon,
■Middlesex, 1; Chiswick, ]\Iiddlesex, 1; Regent's Park, Loudon, 2; Ban-
stead, Surrey, 18; Wandsworth Common, Surrey, 4; Botchworth,
Surrey, 1; Yalding, Kent, 4; Crowl)orough, Sussex, 3; Dobbin Isbmd,
Pagham Harbour, Sussex, 1; Alum Bay, Isle of Wight, 4; Bemhridge,

806

RODENTIA

Isle of Wight, 3 ; St. Helen's, Isle of Wight, 1 ; Andover, Hampshire, 1 ;
Basingstoke, Hampshire, 3 ; New Forest, Hampshire, 16 ; Monxton,
Hampshire, 1 ; Blandford, Dorsetshire, 1 ; Colleigh, Honiton, Devonshire,
1 ; Comhmartiu, Devonshire, 1 ; Brent Knoll, Somersetshire, 1 ; Northlew,
Devonshire, 3 ; Tresco, Scilly Islands, Cornwall, 6.

Norway : Near Bergen, 18 (B.M. and U.S.N.M.) ; Eggedal, 10
(U.S.N.M.) ; Aker, Christiania, 1 ; Asker, Christiania, 6 (U.S.N.M.).

Sweden: Upsala, 9 (B.M. and U.S.N.M.).

Denmark : Hillerod, Zealand, 2.

Holland: Oosterbeek, Guelderland, 2; Noordwijk, near Leiden, 3;
Leiden, 3 (U.S.N.M.).

Belgium: Esneux, Li6ge, 4; Waremme, Li^ge, 39 (U.S.N.M.).

France: Guines, Pas-de-Calais, 2; near Boulogne, Pas-de-Calais, 4;
Trinity, Jersey, Channel Islands, 3 ; St. Martins, Guernsey, Channel
Islands. 1 ; Alderney, Channel Islands, 6 ; Sark, Channel Islands, 1 ;
Dinan, 7; Huelgoat, Finistere, 1 ; Barbizon, Seine-et-Marne, 1 ; Mt. Dore,
Puy-de-D6me, 3 ; Cadillac, Gironde, 27 (U.S.N.M.) ; Solferino, Landes, 9 ;
Poret de Bouconne, Gers, 14; Montr^jeau, 11 (U.S.N.M.); Biarritz,
Basses-Pyr6n6es, 7 ; St. Paul, near Barcelonnette, Basses-Alpes, 27 ;
Chamonix, Haute-Savoie, 6 (U.S.N.M.); Crauves-Sales, Haute-Savoie, 8 ;
Lucinges, Haute-Savoie, 6 ; Montauban, Haute-Savoie, 3 ; fitupes,
Doubs, 4 ; Manonville, Meurthe-et-Moselle, 2.

Germany : Brunswick, 34 (U.S.N.M.) ; Hausbruch, Hanover, 2 ;
Bodethal, Harz INIountains, 4 (U.S.N.M.) ; Bahrenberg, Harz Mountains, 4
(U.S.N.M.) ; Wolfshau, Riesengobirge, Silesia, 22 (U.S.N.M.) ; Euiengrund,
near Schneekoppe, Silesia, 4 (U.S.N.M.); Niesky, Silesia, 7 ; Magdeburg,
Saxony, 2; Marxheim, Bavaria, 5; Strass, near Burgheim, Bavaria, 3;
Schwarzburg, 3 ; Kalbe, 1 ; Ingelheim, Rheinhessen, 2 ; Strassburg, 6.

Austria-Hungary : Haida, Bohemia, 2 ; Hainspach, Bohemia, 4
(U.S.N.M.) ; Csallokoz-Somorja, Pressburg, 2 ; Hatszeg, Hunyad, 3 ;
Transylvania, 1 ; Eszek, Slavonia, 4.

RouMANiA : Gageni, Prahova (at base of Carpathians, north-west of
Bucharest), 7.

Sw^itzerland : Geneva, 7 (U.S.N.M.) ; St. Cergues, Vaud, 4 (U.S.N.M.) ;
Les Plans, Vaud, 3 (U.S.N.M.); Zermatt, Valais, 1 (U.S.N.M.); Meiringen,
Bern, 43 (U.S.N.M.) ; Brunig, Bern, 5 (U.S.N.M.) ; Lucerne, 3; Vitznau, 1
(U.S.N.M.) ; Goschenen, Uri, 2 (U.S.N.M.) ; Andermatt, Uri, 1 (U.S.N.M.) ;
St. Gallen, 4 (U.S.N.M.) ; Murgthal, St. Gallen, 1 (U.S.N.M.) ; Sitterwald,
St. Gallen, 5 (U.S.N.M.) ; Wildpark, St. Gallen, 1 (U.S.N.M.) ; Ziiber-
wangen, St. Gallen, 1 (U.S.N.M.); Wolfhalden, Appenzell, 5; Albulapass,
Grisons, 5 (U.S.N.M.) ; Bergiin, 'Grisons, 1 (U.S.N.M.) ; Vulpera-Tarasp,
Grisous, 5 (Rothschild) ; Paido, Ticiuo, 3 (B.M. and U.S.N.M.) ; Lugano,
Ticino, 6 (U.S.N.M.) ; Capolago, Ticino, 1 ; Locarno, Ticino, 7.

Italy: Padola, Cadore, 1 (Turin).

2 6,19. Clandeboye, Down, Ire- W. R. Ogilvie-Grant 11. 1. 3. 318-320.

land. (c & p).

1 al. Meath. Dr. G. E. Dobson (p). 80. 12. 14. 5.

3 c5, 3 9. Clare Island, Mayo. G. Barrett-Hamilton 11. 1. 2. 132-137.

(R. H. Bunting.) (p).

2 c5, 2 9. Inishmore, Aran Islands. G. Barrett-Hamilton 11. 1. 2. 138-141.
{B. H. Bunting.) (p).

3 6. Scariff, Galway. G. Barrett-Hamilton 11. 1. 2. 68-70.

(R. F. Hihhcrt.) (p).

6. Clare. {R. F. Hibbcrt.) G.Barrett-Hamilton 11.1.2.71.

(P)-
6. Nenagh, Tipperary. Capt. W. F. Smith 11. 1. 3. 321.

(c & p).
6. Cashel, Tipperary. W. E. de Winton 11. 1. 3. 322.

(c & p).
2 6,2 9. Ballaghmoon, Carlow. G. Barrett-Hamilton 11. 1. 2. 64-67.
(J. G. Symes.) (p).

Al'ODEMUS

807

6. Cappagh, Wat.orford.

{W.J. Ussher.)
2 6. Kilmanock, Wexford.

(Connell.)
9. Kerry.

2 <S, 9. Caragh Lake, Kerry.

1 al. Islay, Scotland.
6, 9. Tiree Island.

(P. Anderson.)
9. Isle of Skye.
2 6, 9. South Sutor, Cromarty.

6, 9. Black Isle, Cromarty.

2 S. Lossiemouth, Elgin.
9 al. Craigellachie, Elgin.

2 6,2 9. Dunphaii, Elgin.

6, 9. Lhanbride, Elgin.

2 6. Dalmeny Park, Edin-

burgh.
6. Stockbriggs, Lanarkshire,
2 6,9. Ramsay, Isle of Man.

3 6. Selby Bridge, Isle of Man.
2 6,2 9. Swithland, Leicester-
shire, England.

6, 9 al. Belvoir, Leicestershire.

1 al. Swiucliffe, Shropshire.

2 6. Aberia, Merionethshire,

Wales.
2 6,2 9. Laugharne, Carmarthen-
shire.

4 6. Bridgend, Glamorgan-

shire.
(5, 4 9. Graftonbury, Hereford-
shire, England.

3 al. Bishopstone, Hereford-

shire.
6. Bloxham, Oxfordshire.
2 6,2 9. Wormsley, Oxfordshire.

S 6,9. Yalding, Kent.

G. Barrett-Hamilton 11. 1. 2. 72.

G. Barrett-Hamilton 11. 1. 2. 62-03.

(P)-
Col. J. W. Yerbury 0. 3. 11. 1.

(c & p).

{Type of celticus Barr.-Ham.)
Col. J. W. Yerbury 11. 1. 3. 323-325.

(c & p).
H. Russell (c & p). 9. 9. 11. 4.
LordElphinstone(p). 8.3.16.1-2.

9.
9 6,2 9.

Chiswick, Middlesex.
New Forest, Hampshire.

3 6. Basingstoke, Hampshire.
3. Clifton, Gloucestershire.

2 c?, 9. Clifton
9. Clifton

3 6, 9. Lowestoft, Suffolk.
6. Dobbin Island, Sussex,

England.

{J. E. Harting.)
4 c?, 2 9. Scilly Islands, Cornwall,
England.
9. Brent Knoll, Somerset.
3 6, 9. Opheim, Bergen, Norway.

J. S. Elliott (c & p).
W. R. Ogilvie-Grant

(c & p).
W. R. Ogilvie-Grant

(c & p).
G. Denoon (c & p).
J. E. Harting (c & p).
W. R. Ogilvie- Grant

(c & p).
W. Taylor (c & p).
W. Evans (c & p).

E. R. Alston (c & p).
C. H.B. Grant (c&p).
C. H.B.Grant (c&p).
F.A. Butler (c&p).

F. T. Mott (c & p).
A. Smith (c&p).

G. H. Caton-Haigh
(c & p).

W. E. de Winton

(c & p).
R. I. Pocock (c&p).

W. E. de Winton

(c & p).
Rev. S. 0. Ridlcv

(c & p).
O. V. Aplin (c & p).
W. R. Ogilvie-Grant

(c & p).
W. R. Ogilvie-Grant

(c & p).
Miss Sharpe (c&p).
Miller Collection.

Miller Collection.
Miller Collection.
R. I. Pocock (c&p).
R. Powell (p).
O. Thomas (c & p).
E. R. Alston (p).

F. J. Cox (c & p).

A. B. Percival (p).
Miller Collection.

11. 1. 3. 315.
11. 1. 3. 303-305.

11. 1. 3. 301-302.

11. 1. 3. 312-313.

89. 5. 17. 1.

11. 1. 3. 306-309.

11. 1. 3. 310-311.
11. 1. 3. 316-317.

79. 9. 25. 60.
11. 1. 3. 346-348.
11. 1.3. 349-351.
11. 1. 3. 361-364.

85. 5. 5. 1-2.
74. 11. 25. 3.
11. 1. 3. 326-327.

11. 1. 3. 328-331.

11. 1. 3. 332-335.

11. 1. 3. 370-374.

81. 1. 1. 1-3.

11. 1. 3. 369.
11. 1. 3. 365-368.

11. 1. 3. 352-355.

11. 1. 3. 356.
7.7.7.2887-2890,

2891-2902.
7.7.7.2903-2905.
7.7.7.3601-3603.
11. 1. 3. 342-344.
11. 1. 3. 345.
11. 1. 3. 357-300.
79. 9. 25. 41.

11.1.3. 336-341.

3. 12. 17. 1.

7.7.7.4464-4407.

808

RODENTIA

1. Aker, Christiauia, Nor- Chrisfciauia Museum 93. 3. 1. 10.

way. (e). ,

9. Upsala, Sweden. Lord Lilford (p). 11. 1. 1. 163.

(Kolthoff.)

2 9. Hillerod, Zealand. Den- 0. Thomas (c & p). 98. G. 7. 11-12.

mark.

^,9. Oosterbeek.Guelderland, 0. Thomas (c & p). 98.2.1.16-17.

Holland.

6, 2 9. Noordwijk, Leiden, Hoi- 0. Thomas (c & p). 98. 2. 1. 18-20.

land.

6, 8 9. Esneux, Li6ge, Belgium. Lord Lilford (p). 95. 1. 1. 12-20.

(H. Gronvold.)

6, 9. Guines, Pas -de -Calais, 0. Thomas (c & p). 94. 6. 6. 14-15.
France.

2 (J, 2 9. Boulogne, Pas-de-Calais. 0. Thomas (c & p). 98. 1. 9. 12-15.

4 al. Alderney, Channel Is- W. Eagle Clarke 98. 9. 29. 3-6.

lands. (c & p).

2 6. Alderney, Channel Is- G. Barrett-Hamilton 11. 1. 2. 73-74.

lands. {W.Eagle Clarke.) (p).

2 6. Trinity, Jersey, Channel 0. Thomas (p). 8. 9. 2. 10-11,23.

Islands. {R.H. Bunting.)

6. Sark, Channel Islands. Guernsey Society of 10. 2. 27. 1.
{G. F. Derrick.) Natural Science (p).

4 (5, 4 9. Dinan, Brittany. G. Barrett-Hamilton 11. 1. 2. 115-122.

9. Huelgoat, Brittany. 0. Thomas (c & p). 92. 9. 5. 2.

6 (5, 3 9. Solf6rino, Landes. O. Thomas (p). 6. 4. 1. 66-74.
{A. Robert.)

5 6,2 9. Poret deBouconne, Gers, 0. Thomas (p). 6. 4. 1. 59-65.

250 m. {A. Robert.)
2 6,2 9. Cranves - Sales, Haute- A. Robert (c & p). 5. 4. 4. 2-5.

Savoie, 900 m.
3 <^. Cranves - Sales, Haute- 0. Thomas (p). 5.11.18.11-13.

Savoie. {A. Robert.)
8 6,8 9. Barcelonnette, Basses- 0. Thomas (p). 8. 8. 10. 74-89.

Alpes. (C. Mottaz.)
6. Barbizon. G. S. Miller (c). 8. 8. 4. 199.

2 6, 9. Mt. Dore, Puy-de-D6me. G. S. Miller (c). 8. 8. 4. 210-212.

4 (5, 3 9. Biarritz, Basses-Pyr6n6es. J. F. Davison (c & p). 6. 6. 4. 3-9.
3. Nieskv, Silesia, Germany. Lord Lilford (p). 99. 1. 9. 17-18.

{W.Baer.)
6, 3 9. Niesky, Silesia. Dr. E. Hamilton (p). 97. 12. 4. 21-24.

{W. Baer.)
2. Magdeburg, Saxony. Dr. W. Wolterstorff 92. 12. 1. 8-9.

(c & p).
6. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 158.

(Wolterstorff.)
2 9. Marxheim, Bavaria. " Lord Lilford (p). 11. 1. 1. 161-162.

{Wolterstorff.)
6, 9. Sirass. Burgheim, Ba- Lord Lilford (p). 11. 1. 1. 159-160.

varia. [Korbitz.)
2 6, 9. Schwarzburg. Lord Lilford (p). 95. 4. 18. 12-14.

(TF. ScJtlilter.)
6. Kalbe, Germany. Lord Lilford (p). 11. 1. 1. 157.

(Woltersiorff.)
2 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 44-45

4:6,2 9. Strassburg, Alsace. O. Thomas (p). 8. 8. 10. 139-144.

(C. Mottaz.)

2. Csallokoz - Somorja, Budapest Museum (k). 94. 3. 1. 46-47.

Pressburg, Hungary.

APODEMUS

809

6, 2 ?. Hatszeg, Huuyad, Tran- C. G. Danford (c). 3. 2. 2. 25, 30, 32.

sylvania.

1 al. Transylvania. C. G. Danford (p). 80. 10. 28. 1.

4 al. Eszek, Slavonia. Budapest Museum (e). 94.7.23.2-5.

4 J, 4 9. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 17-24.
mania. (W. Dodson.)

4 (5, ?. Wolfhalden, Switzer- Miller Collection. 7. 7. 7. 10-14.
land. {Zollikofer.)

9. Capolago, Ticino. 0. Thomas (c & p). 2. 7. 1. 6.

3 6, 3 9. Locarno, Ticino. 0. Thomas (c & p). 2. 7. 1. 7-12.

6. Faido, Ticino. 0. Thomas (c & p). 2. 7. 1. 19.

3 6. Lucerne, Switzerland. Lord Lilford (p). 11. 1. 1. 154-156.
(Woltersto'rff.)

Apodemus sylvaticus callipides Cabrera.

1907. Micromys sylvaticus callipides Cabrera, Bol. Real Soc. Espafi. Hist.

Nat., Madrid, vii, p. 227, November, 1907 (Villarrutis, La Corufia,

Spain). Type in JMadrid Museum.
1910. Mus sylvaticus callipides Trouessart, Paune I\Iamm. d'Europe, p. 153.

Type locality. — Villarrutis, La CoruSa, Spain.

Geographical distribution. — Mountains of the Pyrenees-
Asturias chain ; southward into Portugal.

Diagnosis. — Size distinctly larger than in Apodemus sylvaticus
sylvoJicus, the hind foot most frequently 23 mm., condylobasal
length of skull usually 23 • 6 mm. or more ; colour dark and rich
as in the typical form.

Measurements. — Type, adult male (from Cabrera) : head and
body, 88-8; tail, 94*5; hind foot, 23. Adult male and female
from Cintra, Portugal : head and body, 100 and 100 ; tail, 107
and 100 ; hind foot, 24*8 and 24 ; ear from meatus, 17 and 17.
Adult male from Estoril, Portugal : head and body, 101 ; tail,
103; hind foot, 23-3; ear from meatus, 16*2 Average and
extremes of ten adults from Pajares, Leon, Spain : head and
body, 100-3 (91-110); tail, 92-3 (80-104); hind foot, 23-0
(22-24) ; ear from meatus, 17 ' 2 (16-18). Average and extremes
of eight adults from the vicinity of Luchon, Haute-Garonne,
France: head and body, 98-7 (98-100); tail, 102-5 (90-115);
hind foot, 22-7 (22-24); ear from meatus, 17-1 (16-18).
Average and extremes of ten adults from Ax-les-Thermes, Ariege,
France: head and body, 98-2 (94-105); tail, 100-8 (90-110);
hind foot, 22-7 (22-23*6). For cranial measurements see Table,
p. 819.

Specimens examined. — One hundred and twelve, from the following
localities : —

France : Port6, Pyrenees-Orientales, 9 ; I'Hospitalet, Ariege, 8 ; Ax-
les-Thermes, Ariege, 31; Luchon, Haute-Garonne, 11; Caterllle, Haute-
Garonne, 5; Bareges, Hautes-Pyr^nuos, 19; Pic-du-Midi,HautGS-Pyr^n6es, 1.
Spain : Pajares, Leon, 10 ; La Coruiia, 2 ; Villalba, Lugo, 5.
Portugal: Cintra, 7 ; Estoril, 2; Oporto,!.
4 S. Port6, Pyr6nees-Oriontales, G. S. Miller (c). 8. 8. 4. 200-203.

Prance.
6,'i. Porte, Pyrenees-Orientales; O. Thomas (p). 8.9.1.57 57.
lG-1700 m. (/I. Hubert.)

810

RODKNTIA

5 6, 9. L'Hospitalet, Ariege. G. S. Miller (c). 8. 8. 4. 204-209.

6, 9. L'Hospitalet, Ariege. O. Thomas (p). 8. 9. 1. 54-55.
{A. Robert.)

4 (J, 4 9. Ax-les-Thermes, Ariege. G. S. Miller (c). 8. 8. 4. 213-220.

5 .5, 3 9. Luchon, Haute-Garonne. 0. Thomas (p). 6. 4. 1. 40-47.

{A. Robert.)

3 6,2 9. Caterille, Haute-Garonne. 0. Thomas (p). 6. 4. 1. 54-58.

(.4. Robert.)

3 6,4 9. Bareges, Hautes-Pyren6es. G. S. Miller (c). 8. 8. 4. 183-189.

c5. Pic-du-Midi, Hautes-Pyr6- 0. Thomas (p). 8. 9. 1. 58.

nees. {A. Robert.)

3 6,5 9. Pajares, Leon, Spain. O. Thomas (p). 8. 2. 9. 110-117.
{N. Gonzalez.)

2 al. Coruna. D. V. L. Seoane 94. 3. 19. 1-2.

(c & p).

5 al. Villalba, Lugo. Dr. V. L. Seoane 94. 1. 1. 10-13.

(c & p). 95, 4. 29. 2.

2 6,2 9. Cintra, Portugal. O.Thomas (c & p). 98! 2. 2. 22-25.

6,9,1. Cintra. {E.Jordan.) Hon. N.C. Roths- 10.9.28.1-3.

child (p).

2 6. Estoril. O.Thomas (c &p). 98. 2. 2. 26-27.

9. Oporto. (J. Searle.) Lord Lilford (p). 11. 1. 1. 164.

Apodemus sylvaticus dichrurus Rafinesque.

1814. Musculus dichrurus Rafinesque, Precis des D6couvertes Somio-

logiques, p. 13 (Sicily).
1844. Mus pecchioli Pecchioli, Atti della quinta Uuioue degli Scienziati

Italiani, Torino, 1843, p. 426 (Tuscany, probably vicinity of

Siena, Italy).
1900. Mus sylvaticus hayi Barrett-Hamilton, Proc. Zool. Soc, London,

p. 410 (]3art).
1910. Mus sylvaticus hayi and M. sylvaticus dichrurus Trouessart, Faune

Mamm. d'Europe, p. 154.

Type localitij. — Sicily.

Geographical distribution. — Mediterranean region from the
Balkan Peninsula to central and southern Spain.

Diagnosis. — Size as in Apodemus sylvaticus callipides, but
colour differing from that of the other continental European
races in a general pallor and dulness resulting from the suppres-
sion of reddish tints and the predominance of yellow and grey.

Colour. — Typical specimens from Sicily have the ground
colour of back a very uniform dvill buffy wood-brown, slightly
darker along median posterior region, and slightly more greyish
anteriorly, the black hair tips, though numerous, nowhere
sufficiently abundant to ]^roduce any evident effect of clouding ;
sides faintly paler and more buffy, the lower portion of cheek
and outer surface of front leg nearly clear buff as in true
sylvaticus ; underparts dull white ; pectoral spot slightly developed
and rarely forming a collar ; tail rather less evidently bicolor
than in the central European form. Grey extreme : ground
colour above a peculiar buffy grey, intermediate between the
ecru-drab and cream-l)uff of Ridgway, but paler than either, the
" lining " produced by the black hairs conspicuous. Yellow

APODEMUS

811

extreme : ground colour a clear buff slightly more yellowish than
that of Ridgway, fading nearly to cream-buff on head and neck,
the black "lining" conspicuous and tending to produce a clouded
effect along middle of back.

All of the Italian skins examined * are essentially like the
typical Sicilian specimens. The same is true of the majority of
those from Spain ; but in this series occur also the extremes of
yellowness and greyness. The skins from Corfu and Cephalonia
are about typical, while those from the mainland of Greece show
a decided tendency toward the yellow extreine.

Measurements. — Adult male and female from Palermo, Sicily :
head and body, 100 and 102 ; tail, 93 and 91 ; hind foot, 23
and 22. Average and extremes of five adults from Sorrento,
Italy : head and body, 100-8 (99-104) ; tail, 94-6 (90-99) ; hind
foot, 21 '9 (21-23). Average and extremes of seven adults from
Genoa, Italy : head and body, 102-8 (9-2-109) ; tail, 87 (80-95) ;
hind foot, 22 (21-23). Adult male and female from Corfu,
Greece: head and body, 98 and 100; tail, 100 and 97; hind
foot, 22 and 21-5 ; ear from meatus, 16-7 and 17. Adult male
and female from Cephalonia, Greece : head and body, 101 and
100; tail, 99 and 98; hind foot, 23 and 21 ; ear from meatus,
17-2 and 16-4. Adult male and female from Tatoi, near Athens,
Greece: head and body, 101 and 100; tail, 106 and 106; hind
foot, 23-4 and 21*8; ear from meatus, 16-2 and 16 3. Three
adult males from Valescure, Var, France: head and body, 104,
105 and 106; tail, 107, 100 and 108; hind foot, 23-4, 22-4
and 23; ear from meatus, 18-2, 17*4 and 17. Average and
extremes of ten adults from the vicinity of Nimes, Gard, France :
head and body, 104-2 (98-110) ; tail, 97-9 (93-105) ; hind foot,
22-9 (21-6-24-3); ear from meatus, 16-3 (15-17-6). Four
adult males from Silos, Burgos, Spain : head and body, 97, 98,
101 and 102 ; tail, 92, 88, 99 and 105 ; hind foot, 22, 23, 23-2
and 23; ear from meatus, 18-6, 17, 17 and 17. Average and
extremes of eight adults from the Province of Granada, Spain
(Guadix and Venta del Baul) : head and body, 105-8 (100-113) ;
tail, 109-5 (100-128); hind foot, 24-1 (21-4-25); ear from
meatus, 17-5 (16-5-19). Average and extremes of ten adults
from the Province of Alicante, Spain (Elche and Alcoy) : head
and body, 104-4 (100-107); tail, 103-2 (92-109); hind foot,
23-2 (21-24); ear from meatus, 18-3 (17-19). For cranial
measurements see Table, p. 820.

Specimens examined. — Two hundred and seventy-three, from the
following localities : —

Spain: Guadix, Granada, 5; Venta del Baul, Granada, 11; Elche,
Alicante, 8 ; Alcoy, Alicante, 3 ; Dehesa de Valencia, 3 ; Barracas, Cas-
tellon, 1 ; Villalba, Madrid, 9 ; B6jar, Salamanca, 10 ; Silos, Burgos, 20 ;

* Except those from the vicinity of Turin, which somewhat approach
true sylvaticus.

812

KODENTIA

Castrillo de la Reiwa, Burgos, 1 ; Panticosa, Huesca, 4 ; Jaca, Huesca, 1 ;
Jerez, Cadiz, 1 ; Inca, Majorca, Balearic Islands, 12 ; San Cristobal,
Minorca, Balearic Islands, 6.

France : Cerbere, Pyr^n^es-Orientales, 1 ; near Nimes, Gard, 21 ;
Valescure, Var, 11 ; Agay, Var, 1.

Italy: Vicinity of Turin, 19 (Turin); Moncalieri, Turin, G (B.INL,
U.S.N.M. and Turin) ; Genoa, 11 (U.S.N. M.) ; Viareggio, 4 ; Sorrento, 12
(B.M. and U.S.N.M.) ; Aspromonte, Lagonegro, 2; Monte Sirino, 1.

Sicily: Acicastello, 1 (U.S.N.M.); Castelbuono,4; Ficuzza,5; Madonna
dell' Alto, 8; Marsala, 4; Palermo, 5 (U.S.N.M.); San Giuglielmo, 4;
Taormina, 4 (U.S.N.M.).

Corsica : Asco, 2 ; La Foce di Vizzavona, 2 ; no exact locality, 3.

Montenegro : No exact locality, 2.

Albania : Northern Albania, 2.

Greece : Corfu, 9 ; Cepbalonia, 13 ; Patras, 1 ; near Athens, 15 ;
Agorianni, 5 (U.S.N.M.).

G. S. Miller (c). 8. 8. 4. 75-79.

G. S. Miller (c). 8. 8. 4. 80-86.

G. S. Miller (c). 8. 8. 4. 87-88.

O. Thomas (p). 8. 2. 9. 127-129.

4c5,l?

4(5,3 9

2<5.

(5,2 9.

Guadix, Granada, Spain.
Venta del Baul, Granada.
Elche, Alicante.
Alcoy, Alicante.

(N. Gonzalez.)
4(5. Villalba, Madrid.

{N. Gonzalez.)
4 (5, 3 9. Bejar, Salamanca.

{N. Gonzalez.)
7 (5, 9. Silos, Burgos.
9. Castrillo, Burgos.

4 6. Panticosa, Huesca.

(N. Gonzalez.)
6. Jaca, Huesca.

(N. Gonzalez.)
9. Jerez, Cadiz.

3 (5, 6 9. Inca, Majorca, Balearic

Islands.

4 (5, 2 9. San Cristobal, Minorca.

(5. juv. Cerbere, Pyrtsn^es Ori-

entales, France.
4 (5, 4 9. Nimes, Gard.

(C. Mottaz.)

4 (5, 3 9. Valescure, Var.

9. Agay, Var.

(5,9. Moncalieri, Turin, Italy.

4 6. Viareggio.
2 (5, 2 9. Sorrento.

2 (5. Aspromonte, Calabria.

[A. Robert.)
i. JMonte Sirino, Calabria.

(.4. Robert.)
3 6, 9. Castelbuono, Sicily.

(A. Robert.)

5 6. Ficuzza, Sicily.

(A. Robert.)

5 c5, 3 9. Madonna dell' Alto,

Petralia, Sicily.

{A. Robert.)
2 (5, 2 9. Marsala, Sicily.

{A. Robert.)

3 (5, 9. San Giuglielmo, Sicily.

(.4. Robert.)

0. Thomas (p).

O. Thomas (p).

G. S. Miller (c),
G. S. Miller (c),
0. Thomas (p).

0. Thomas (p).

8. 2. 9. 122-125.

8. 2. 9. 103-109.

8. 8. 4. 66-73.

8. 8. 4. 74.

8. 2. 9. 118-121.

8. 2. 9. 126.

Abel Chapman (c & p). 8. 3. 26. 5.
0. Thomas and R. 1. 0. 7. 1. 9-17.

PoCOCk (C & P;.

0. Thomas and R. I. 0. 7. 1. 52-57.

Pocock (c & p).
0. Thomas and R. I. 0. 7. 1. 75-76.

Pocock (c & p).
O. Thomas (p). 8. 8. 10. 66-73.

G. S. Miller (c).
G. S. Miller (c).
Dr. E. Festa (p).
0. Thomas (c & p).
R. J. Cuninghame

(c & p).
O. Thomas (p).

O. Thomas (p).

O. Thomas (p).

O. Thomas (p).

0. Thomas (p).

O. Thomas (p).
O. Thomas iv).

8. 8. 4. 190-197.
8. 8. 4. 198.
8. 8. 1. 1-2.

5. 8. 3. 2-5.
98. 5. 2. 12-15.

6. 8. 4. 6-7.
6. 8. 4. 8.

{t:

8. 4. 42-43.

9. 1. 24-25.
(6. 8. 4. 39-41.
18. 9. 1. 22-23.

8. 9. 1. 10-17.

6. 8. 4. 35-38.

[). 1. lS-21.

APODEMUS

813

.5,9. Asco, Corsica. O. Thomas (i>). S. 11. :{0. 7-8.
(Dr. F. Major.)

2. La Pooo di Vizzavona, Col. J. W. Ycubury 'j:3. 'J. 15. 4-5.

Corsica. (r).

3 al. Corsica. Col. J. W. Yerbury 93. 9. 15. 6-8.

2 6. Montenegro. (L. Fiihrcr.) O. Thomas (p). 5. 8. 4. 16-17.

6, 9. North Albania. 0. Thomas (p). 5. 8. 4. 36, 87.
(I/. Fiihrcr.)

6. Potamos, Corfu, Greece. J. I. S. Whitaker (p). 8.10.1.10.

6. Cephalonia. C. Mottaz (c). 8. 11. 3. 11.

8 6,4:9. Cephalonia. (C. Mottaz.) J. I. S. Whitaker (p). 8. 10. 1. 11-22.

6. Itaci, Athens. Hon. N. C. Roths- 8. 10. 2. 26.

(C. Mottaz.) child (p).

4 9. Tatoi, Athens. Hon. N. C. Roths- 8. 10. 2. 33-3G.

(C. Mottaz.) child (p).
9. Kephissia, Athens. Hon. N. C. Roths- 8. 10. 2. .37.

(C. Mottaz.) child (p).

Apodemus sylvaticus creticus Miller.

1910. Aiiodemus sylvaticics creticus Miller, Ann. and Mag. Nat. Hist.,
8th ser., vi, p. 460, November, 1910.

Type locality. — Katharo, Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Size small, essentially as in true A. sylvaficna, but
colour paler and more yellowish.

Colour. — Ground colour of upper parts a clear, light, yellowish
cream-buff without greyish tinge, except occasionally on crown
and neck, scarcely different on sides away from line of
demarcation, where it becomes slightly more yellowish ; back
heavily " lined " with black, this diminishing rather abruptly on
sides. Underparts and feet white, the region from interramia
to base of tail dulled by the blackish-slate undercolour. Tail
sharply bicolor, dark brown above, white below. Ear dark
brown sprinkled with silvery hairs which tend to form a narrow,
faintly defined whitish edging.

Skull and teeth. — The skull and teeth are smaller than in the
other Mediterranean forms, but are not certainly distinguishable
from those of A. sylvaticus sylvaticus,

Measurements. — Type (adult female) : head and body, 88 ;
tail, 88; hind foot, 21; ear from meatus, 17. Average and
extremes of six adults from Katharo and Kania, Crete : head
and body, 84-6 (80-88); tail, 87-8 (86-89); hind foot, 21-5
(21-22); ear from meatus, 16 "5 (16-17). For cranial measure-
ments see Table, p. 823.

Specimens examined. — Six, all from the island of Crete (two from
Kanea, the others from Katharo).

Remarhs.- — The Cretan form of Apjodemus sylvaticus combines
the small size of the typical race with the dull yellowish colour
of A. sylvaticus dichrurus.

2 6,2 9. Katharo, Crete. Miss D. Bate (c). 5. 12. 2. 25-28.

(5. 12. 2. 27. Type of subspecies.)
6, 9. Kanea. Miss D. Bate (c). 5. 12. 2. 29-30.

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APODEMUS

823

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824

APODEMUS HEBRIDENSIS de AViiiton

1895. Mtia lu'bridcnsis de Wintou, The Zoologist, 3rd ser., xix, p. 3G9,
October, 1905. Type in British Museum.

1895. Mus sylvaticHS hebridensis de Wiutou, The Zoologist, 3rd ser., xix,
p. 426, November, 1905.

1900. Mus sylvatlcus hebridensis Barrett-Hamilton, Proc. Zool. Soc.,
London, p. 403.

1910. Mils sylvaticns hebridensis Trouessart, Fauue j\Iamm. d'Europc,
p. 151.

Type Jocaliti/. —Vig, Island of Lewis, Outer Hebrides,
Scotland.

Geographical distribution. — Island of Lewis, and Barra Island,
Hebrides.

Diagnosis. — Size about as in the large southern races of
Apodemus sylvaticus ; head and body, 106 to 112 ; tail vertebra?,
95 to 100 ; hind foot about 25 ; condylobasal length of skull, 24
to 25 mm. ; ear reduced in size, barely or not reaching eye when
laid forward, its height above meatus 15 to 16 mm., its width in
dry specimens noticeably less than in A. sylvaticus; general
colour of upper parts essentially as in ^. sylvaticus sylvaticus, but
underparts so heavily washed with dull buffy brown that line of
demarcation along sides is ill-defined ; pectoral spot present,
usually rather longer than in A. sylvaticus, but not forming a
collar ; skull and teeth as in larger forms of A. sylvaticus.

Meamrements. — Type (adult male) : head and body, 106 ;
tail, 96 ; hind foot, 25 ; ear from meatus, 16. Two adult males
and an adult female from the type locality : head and body, 106,
112 and 108 ; tail, 99, 100 and 95 ; hind foot, 25, 25 and 23-5 ;
ear from meatus, 16, 16 and 15. For cranial measurements see
Table, p. 827.

Specimens examined. — Thirteen, all from the Islands of Lewis and
Barra.

Remarks. — While the colour of this animal is occasionally
duplicated by specimens from Great Britain and the Continent,
the combination of large size, small ear, and dark tints is
sufficient to distinguish the Hebridean Apodemus as a sharply
defined local form.

5. Island of Lewis, W. E. de Winton (c & p). 95. 10. 25. 1.

Hebrides. {Type of species.)

4 (5, 5 9. Island of Lewis. W. E. de Winton (c & p). 8. 10. 11. 13-21.

6. Barra Island. Col. R. W. Finney (c & p). 11. 1. 3. 314.
6 al. Barra Island. J. McRurie (c & p). 11. 1. 3. 415.

APODEMUS 825

APODEMUS HIRTENSIS Banett-Hamilton.

18'J9. Mas liirtcnsis Barrett-Hamiltoii, Proc. Zool. Soc, Loudon, p. 81.
Type in British Museum.

1900. Mus sylvaticiis liirtcnsis Barrett-Hamilton, ]'roc. Zool. Soc, Loudon,
p. 404.

190G. Mas hirtensis Barrett-Hamilton, Ann. Scottish Nat. Hist., Edin-
burgh, XV, p. 3, January, 190G.

1910. Mus sylvaticus hirtensis Trouessart, Faune Mamm. d'Europe, p. 1.52.

Tijpe locality. — Isljind of Saint Kilda, west of Outer Hebrides,
Scotland.

Geographical distribution. — Saint Kilda.

Diagnosis. — Similar to Ajjodenius hehridensis, but skull larger
(condylobasal length, 25 "6 to 26*8) and colour darker.

3Ieasurements. — Type (immature male) : head and body, 81 ;
tail, 85 ; hind foot, 25. Adult male and female from the type
locality: head and body, 107 and 110; tail, 91 and 101 ; hind
foot, 24*6 and 24-6 ; ear from meatus, 17 and 17. For cranial
measurements see Table, p. 827.

Specimens examined. — Fifteen, all from Saint Kilda.

Remarhs. — Although the material representing the two
animals is not entirely satisfactory there seems little reason to
doubt that Apodemus hirtensis is specifically distinct from A.
hehridensis. The skull, as shown by the Table of measurements,
is distinctly larger, while the colour appears to be somewhat
darker than in the Hebrideau form, though of exactly the same
type. Notwithstanding its large size the skull retains all of the
characteristics of that of A. sylvaticus and A. hehridensis, showing
no tendency to assume the angular appearance seen in
A. Jlavicollis.

6. Island of St. Kilda, Outer J. S. Elliott (c & p). 94. 7. 16. 1.
Hebrides. (Type of species.)

6 and St. Kilda. (H. Evans.) G.Barrett-Hamilton 11. 1. 2. 123.
6 skulls. (p).

3(5,3 9. St. Kilda. W. Eagle Clarke 11. 1. 24. 1-^G.

(c & p).

APODEMUS FRIDARIENSIS Kinnear.

1906. Mus sylvaticus fridariensis Kinnear, Ann. Scottish Nat. Hist., xv,
p. 48, April, 1906. Type in Royal Scottish Museum.

1910. Mus sylvaticus fridariensis Trouessart, Faune Mamm. d'Europe,
p. 150.

Type locality. — Fair Isle, Shetland Islands, Scotland.

Geographical distribution. — Fair Isle.

Diagnosis. — Size as in Apodemus hirtensis or somewhat larger ;
skull with rostrum noticeably more slender than in any of the
related forms ; colour above essentially as in A. sylvaticus
sylvaticus, but back and sides less bright and more clouded with

826 ROHENTIA

black, and uuderparts entirely <lull wliitisli, i]\c pectoral spot
absent or barely indicated.

External characters. — In general external characters Apodemus
fridariensis agrees with A. si/lvaticus and A. JlavicoUis, but the
ear is relatively not so large, barely covering eye when laid for-
ward, and the palmar and plantar tubercles are relatively smaller.
The latter peculiarity is especially noticeable in the hind foot,
where the area occupied by the pads as compared with the space
between them is distinctly less than in the related species.

Colour. — ^Upper parts as in the darker less russet specimens
of A2yodemHS sylvaticus sijlvaticus, the black clouding of median
region and " lining " of sides more noticeable than usual in the
smaller animal, the " lining " effect on sides increased by a
certain harsh loose texture of the fur which causes the dark
hair-tips to stand out with unusual distinctness against the light
undercolour. Underparts a peculiar dull bluish white, noticeably
different from the clear white of A. fiavicoUix, and without trace
of the brownish wash so frequently indicated in A. sylvaticus,
though a faint buffy tinge is sometimes present ; pectoral spot
usually absent, bvit occasionally represented by a tuft of
brownish hairs. Tail very sharply bicolor, blackish above,
whitish below. Feet dull white.

Skull and teeth. — In size the skull of Ajwdeyims fridariensis
rather exceeds that of the large Mediterranean forms of
A. sylvaticus, though by no means equalling the maximum of
A. Jiavicollis. In general form it is more slender than in the
related species, a peculiarity especially noticeable in the rostrum.
Brain-case longer and narrower than in A. sylvaticus, its upper
surface never developing lateral ridges, even in extreme old age.
Mandible slender, the coronoid process attenuate, short, noticeably
less developed than in the related species. Teeth as in Apodemus
sylvaticus.

Measurements. — Average and extremes of ten adults : head
and body, 108-5 (103-114); tail, 102-4 (97-5-104); hind foot,
24-26-4 ; ear from meatus, 16-6 (16-18). For cranial measure-
ments see Table opposite.

Specimens examined. — Twenty-three, all from Fair Isle (B.IM., U.S.N.M.
and Edinburgh).

Remarks. — Notwithstanding its large size this species, like
Apodemus hebridensis and A. hirtensis, appears to be an offshoot of
A. sylvaticus rather than of A. jiavicollis. This is indicated
chiefly by the slenderness and lack of angularity of the skull,
the brain-case, even in excessively old individuals, always retain-
ing the smoothness of surface characteristic of the smaller animal.

6. Fair Isle, Shetlands. N. B. Kinnear (c & p). 6. 5. 19. 1.

{Paraiype of species.)
6 St. Fair Isle. N. B. Kinnear (c & p). 6. 11. 18. 1.

2 <5, 1 9. Fair Isle. N. B. Kinnear (c & p). 6. 11. 18. 2-4.

6, 2 9. Fair Isle. Duchess of Bedford 10. 10. 11. 1-3.

(c & p).

APODEMUS

K27

Observations.

o

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Sex.

>

Number.

95. 10. 25. 1*

94. 7. 16. 1*

11. 1. 2. 123

153507

1905. 120. 2. 7t

9t

16t

1905. 120. 4t
1905. 120. If

6.11.18.2
6.11.18.4

196t

289t
153499
153500
153502
153503
103504
103506

Locality.

J '^.

isis.

id of

>>

is.

Islan

isis.

A, hebridei

Scotland : Uig, Islai
>>

A. hirtens

Scotland : St. Kilda

A. fridariei

Scotland : Fair Isle

>>

828

APODEMUS FLAVICOLLIS Melchior.

(Synonymy under subspecies.)

Geoyraphical distribution. — Central Europe from Sweden and
Finland to the Pyrenees, Alps and Greece, and from Great
Britain to Roumania and western Russia ; eastern limits of
range not known.

Diagnosis. — Decidedly larger than Apodenuis sylvaticas syl-
vaticus, the only member of the group together with which it
occurs; head and body about 100 to 115; tail vertebra^ about
105 to 125; hind foot, 23 to 27, most frequently 25; cond}'-
lobasal length of skull, 25 to 28 '8 mm. ; skull in old individuals
becoming decidedly more angular than that of A. sylvaticus ; ear
large, its height above meatus about 17 to 19 mm. ; general
colour of upper parts brighter and more russet than in any of
the European races of A. sylvaticus ; underparts always whitish,
the line of demarcation along sides sharply defined ; chest spot
usually larger than in A. syhaiicus, frequently spreading laterally
to form a complete collar.

External characters. — As in Ajwdemus sylvaticus, except for
such slight peculiarities as result from the larger size.

Colour. — Except for the general tendency toward brighter,
more russet tints on the upper parts, and the constant absence
of all trace of brownish wash below, the colour so exactly
resembles that of Apodemtis sylvaticus sylvaticus as to need no
detailed description.

Skull and teeth. — In individuals old enough to show con-
spicuous wear of the teeth the skull
attains a degree of massiveness
and angularity never seen in
A. sylvaticus. This is especially
noticeable in the lachrymal and
interorbital regions and on the
upper surface of the brain-case. The
angularity of the interorbital rims
in such specimens is continued for-
ward nearly to lachrymal region
and is so strongly developed as to
produce a slight but evident longi-
tudinal furrow along middle of
frontal ; posteriorly the rims are
continued backward as low but
evident ridges extending along sides
of brain-case to outer extremities
of interparietal. In other respects
the skull shows no tangible pecu-
liarities of form as compared with
that of the smaller animal. Large specimens of Ajpodemus flavi-
collis are e([ual in length to skulls of immature A. epimelas; but

Fig. 167.
Apodemits flavicoUis. Nat. size.

APODEMUS gOy

the latter may always be distinguished by their larger, smoothly
rounded brain-case (a character which persists to extreme old
age), relatively deeper rostrum, and larger teeth.

Teeth. — In all respects, except for their slightly larger size,
the teeth resemble those of Apodemus sylraticus.

Bemarls. — The large general size, long hind foot, and massive,
angular skull serve to distinguish adults of this species from
individuals of Apodemns si/lvaticus. Immature specimens are,
on the contrary, difficult to determine. The relatively limited
distribution of the large animal and the fact that it is not yet
known to occur on any of the small islands where A. sylvafims
has been found, are both circumstances which suggest the
probability that Apodemus flaijicoUis has entered western Europe
more recently than the small species. Members of the group
occur in the Himalayas ; but it is not known how much further
east their range extends.

Apodemus flavicollis flavicollis Melchior.

1834. Mus flavicollis ]\Ielchior, Deu Dauske Staats og Norges Pattedyr
p. 99 (Sielland, Denmark). '

1862. ? [^^us sijlvaticus] vrt. major Radde, Eeisen im Siiden von Ost-
Sibu-ien, i, p. 180 (Bureja Mountains). Part, specimens mentioned
from Crimea which are probably this form.

1866. Mus cellarius J. F. Fischer, Zool. Gart., vii, p. 15.3, April, 1866
(near Luga, St. Petersburg, Russia).

1874. Mus sijlvaticus Lilljeborg, Sveriges og Norges Ryggradsdjur, i, p. 263
(description based on the large animal).

1894. Mus flavicollis de Winton, The Zoologist, 3rd ser. xviii p 441
December 1904 (part).

1900. Mus sijlvaticus typicus Barrett-Hamilton, Proc. Zool. Soc, London
p. 404. Not Mus sijlvaticus LinocTus. ' '

1900. Mus sijlvaticus princeps Barrett-Hamilton, Proc. Zool. ,Soc., London
p. 408 (Busteuari, in the Carpathians, N.W. of Bucharest'
Roumania. Altitude 480 m.). Type in British Museum. '

1910. Mus sijlvaticus and M. sijlvaticus princeps Trouessart, Faune Mamm
d'Europe, pp. 149, 158.

Type locality. — Sielland, Denmark.

Geographical distribution. — Continental range of the species.

Diagnosis. — White of underparts usually pure and without
much suffusion of slaty ; chest spot frequently not forming
complete collar.

Measurements. — Four adults from Medelpad, Sweden : hind
foot, 25, 25, 25 and 26. Adult male and female from Hillerod,
Zealand, Denmark: head and body, 104 and 103; tail. 111 and
104; hind foot, 25 and 24; ear from meatus, 17 and 17-5.
Adult male from Nysted, Lolland, Denmark : head and body,
100; tail, 132; hind foot, 27. Average and extremes of five
adults from Caterille, Haute-Garonne, France : head and body.

830 RODENTIA

110 (106-114); tail, 106-2 (99-114); hind foot, 24-8 (24-26);
ear from meatus, 16 '4 (15-18). Adult female from Bareges,
Hautes- Pyrenees, France: head and body. 111 ; tail, 118; hind
foot, 24-6; ear from meatus, 19. Average and extremes of six
adults from Haute-Savoie, France (Lucinges and Cranves-Sales) :
head and body, 98-8 (95-103); tail, 100-3 (95-103); hind foot,
25-5 (25-26); ear from meatus, 17-5 (16-20). Two adult
females from Brunswick, Germany : head and body, 101 and 97 ;
tail, 116 and 102 ; hind foot, 24 and 24. Average and extremes
of five adults from the Harz Mountains, Germany : head and
body, 106-8 (100-110); tail, 110-6 (102-117); hind foot, 25-3
(24-4-26). Adult female from Konigsberg, Germany: head and
body, 105 ; tail, 105 ; hind foot, 26. Adult male from Niesky,
Silesia : head and body, 115 ; tail, 120 ; hind foot, 24 ; ear from
meatus, 17-5. Adult female from Bustenari, Roumania (type
of princeps) : head and body, 108 ; tail, 115 ; hind foot, 25 ; ear
from meatus, 18. Average and extremes of ten adults from
Bustenari, Roumania: head and body, 109-3 (101-119); tail,
108-3 (104-115); hind foot, 24-4 (23-25-5); ear from meatus,
18-4 (17-5-20). Average and extremes of six adults from
St. Cergues, Vaud, Switzerland: head and body, 108-5 (103-
114); tail, 102-3 (98-111); hind foot, 25-9 (25-4-26-4).
Average and extremes of four adults from Les Plans, Vaud,
Switzerland : head and body, 102-7 (95-120) ; tail, 119-7 (11 5-
125); hind foot, 25-7 (25-26). Adult male irom Murgthal,
St. Gallen, Switzerland: head and body, 112; tail, 128; hind
foot, 26-4. Two adult males from Faido, Ticino, Switzerland:
head and body, 98 and — ; tail, 100 and 129; hind foot, 25
and 26. For cranial measurements see Table, p. 832.

Specimens examined. — One hundred and fifty-one, from the following
localites : —

Sweden : Jemtland, 2 (U.S.N.M.) ; Herjedal, 1 (U.S.N.M.) ; Medelpad,
Norrland, 4.

Denmaek: Hillerod, Zealand, 3; Nysted, Lolland, 2 (U.S.N.M.).

France: Ax-les-Thermes, Ariege, 5 (B.M. and U.S.N.M.); Caterille,
Haute-Garonne, 6 ; Bareges, Hautes-Pyren^es, 3 ; St. Paul, Basses- Alpes, 1 ;
Lucinges, Haute-Savoie, 3 ; Cranves-Sales, Haute-Savoie, 4 ; Cenise, Haute-
Savoie, 2 (Mottaz).

Germany : Brunswick, 3 (U.S.N.]\I.) ; Bahrenberg, Harz Mountains, 3
(U.S.N.M.) ; Bodethal, Harz Mountains, 6 (U.S.N.SI.) ; Mauseklippe, Harz
Mountains, 5 (U.S.N.M.) ; Magdeburg, Saxony, 1 ; Tharandt, Saxony, 1 ;
Strass, near Burgheim, Bavaria, 2 ; Niesky, Silesia, 1 ; near Konigs-
berg, 4.

Austria-Hungary: Haida, Arva, Bohemia, 2; Mittelberg, Vorarl-
berg, 3 ; Schwaz, Tirol, 2 ; Hatszeg, Hunyad, 7.

Roumania : Bustenari, Prahova, 15 ; Gageni, Prahova, 1 ; Comana, 1.

Albania : Northern Albania, 2.

Switzerland : Geneva, 1 (U.S.N.M.) ; St. Cergues, Vaud, 7 (U.S.N.M.) ;
Les Plans, Vaud, 3 (U.S.N.M.) ; Chesieres, Vaud, 1 (Mottaz) ; Meiringen,
Bern, 1 (U.S.N.M.); Sitterwald, St. Gallen, 1 (U.S.N.M.); Murgthal,
St. Gallen, 3 (U.S.N.M.); Degorsheim, St. Gallen, 1 (U.S.N.M.); Wild-
kirchli, Appenzell, 1; Albula Pass, Grisons, 4 (U.S.N.M.) ; Julier-Hospiz,
Grisons, 1 (U.S.N.M.) ; Vulpera-Tarasp, Grisons, 4 (Rothschild) ; Vitznau,

APODEMUS 831

3 (B.M. and U.S.N.M.) ; Goschenen, Uri, 1 (Mottaz) ; Faido, Ticino, 4
(U.S.N.M.) ; Lugano, Ticino, 5 (U.S.N.M.) ; Locarno, Ticino, 6.
Greece : Athens, 6 ; Corfu, 3.

6,9. Medelpad, Norrland, Lord Lilford (p). 8.10.19.28-29.

Sweden.
3 6. Hillerod, Zealand, Den- O. Thomas (c & p). 98.6.7.9-10.13.

mark.

2 6, 9. Ax-les-Thermes, Ariege, V. Builles (c & p). 8. 3. 27. 9-11.

France.
6 6. Caterjlle, Haute-Garoune. O. Thomas (p). 6.4.1.48-53.

{A. Robert.)
9. Bareges, Hautes-Pyren^es. G. S. Miller (c). 8. 8. 4. 221.

3 6,9. Granves- Sales, Haute- O. Thomas (p). 5. 11. 18. 14-17.

Savoie. {A. Robert.)
1 al. Tharandt, Saxony. Berlin Museum (e.) 98. 1. 1. 19.

1. Niesky, Silesia. (17. Boer.) Lord Lilford (p). 99.1.9.19.

2 6, 9. Mittelberg, Vorarlberg, 0. Thomas (p). 8. 11. 30. 9-11.

Austria-Hungary.

(Dr. F. Major.)
5 6,2 9. Hatszeg, Hunyad, Tran- C. G. Danford (c). /3. 2. 2. 26-29. 31.

sylvania. (3. 11. 8. 32-33.

7 6,8 9. Bustenari, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 26-40.

mania. {W. Dodson.)

(4. 4. 6. 34. Type of A. princeps Barrett-Hamilton.)
9. Gageni, Prahova. Lord Lilford (p). 4. 4. 6. 25.

[W. Dodson.)
6, 9. North Albania. 0. Thomas (p). 5. 8. 4. 36.

(L. Filhrer.)
4:6,2 9. Locarno, Ticino, Switzer- 0. Thomas (c & p). 2. 7. 1. 13-18.

land.
2 6, 9. Vitznau, Lucerne. 0. Thomas (c & p). 5. 8. 3. 16-18.

3 d, 3 9. Athens. (C. Mottaz.) Hon. N. C. Roths- 8. 10. 2. 27-32.

child (p).
6, 2 9. Corfu. (C. Mottaz.) J. I. S. Whitaker (p). 8. 10. 1. 23-25.

Apodemus flavicollis wintoni Barrett-Hamilton.

1894. Mus flavicollis de Winton, The Zoologist, 3rd ser., xviii, p. 441,

December, 1894 (part).
1900. Mus sylvaticus wintoni Barrett-Hamilton, Proc. Zool. Soc, London,

p. 406 (part). Type in British Museum.
1910. Mtis sylvaticus wintoni Trouessart, Faune Mamm. d'Europe, p. 149.

Ti/pe locality. — Graftonbury, Herefordshire, England.

Geographical (lisfribiiiion. — Great Britain ; not yet recorded
from Scotland.

Diagnosis. — White of underparts usually more tinged with
slaty than in the Continental form ; chest spot usually forming
a complete collar and continuing backward along median line
nearly to belly.

Measurements. — Type (adult male): head and body, 110;
tail, 112; hind foot, 24. Average and extremes of six adults
from the type locality: head and body, 108*6 (102-115); tail,
107-5 (92-115); hind foot, 24-1 (23-25); ear from meatus,
17 "6 (17-1*J). Adult male from Woolpit, Suffolk: head an(l

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836 RODENTIA

body, 112; tail, 120; hind foot, 24. For cranial measurements
see Table, p. 835.

Specimens examined. — Twenty-two, from the following localities in
England : Riding, Mill-on-Tyne, Northumberland, 1 ; Graftonbury, Here-
fordshire, 13 ; Ledbury, Hereford, 1 ; Bishopstone, Hereford, 3 ; Hereford,
no exact locality, 1 ; Swithland, Leicestershire, 1 ; Oundle, Northampton-
shire, 1 ; Woolpit, Suffolk, 1 ; Balcombe, Sussex, 1.

Bemarlis. — The British race of Apodenms flavicoUis, though
not a Avell marked form, seems worthy of recognition on account
of the duller colour of the underparts and the more diiiuse
nature of the pectoral spot.

6. Riding Mill - on - Tyne, Rev. H. H. Slater 11. 1. 3. 413.
Northumberland, (c & p).
England.

6. Graftonbury, Hereford- W. E. de Winton 0. 3. 12. 1.

shire. (c & p). (jP;/pc of subspecies.)

4 <5, 6 9, Graftonbury, Hereford- W. E. de Winton (p). 8. 10. 11. 1-12.
2 juv. shire.

6, 9. Bishopstone, Hereford- H. N. Ridley (p). 85. 12. 29. 1-2.

shire.

6. Bishopstone, Hereford- Rev. S. O. Ridley 81. 1. 1. 4.

shire. (c & p).

6. Ledbury, Herefordshire. Col. J. W. Yerbury 11. 1. 3. 411.

(c & p).

6. Swithland, Leicestershire. F. A. Butler (c & p). 11. 1. 3. 410.

9. Oundle, Northampton- LordLilford (c &p). 11.1.1.165.

shire.

6. Woolpit, Suffolk. W. D. Parker (c & p). 11. 1. 3. 412.

APODEMUS AGRARIUS Pallas.

1778. Mus agrariiis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 95 (Berlin,

Germany).
1801. Ml^iis] agr[ariHs'] albirostris Bechstein, Gemeinn. Naturgesch.

Deutschlands, i, 2d ed., p. 975 (Thiiringen, Germany).
1801. Jlf[M.s] agr[arius'] maculatus Bechstein, Gemeinn. Naturgesch.

Deutschlands, i, 2d ed., p. 975 (Thiiringen, Germany).
1816. Mus rubens Oken, Lehrb. d. Naturgesch., in, pt. ii, p. 898 (Northern

Germany).
1857. Mus agrarius Blasius, Saugethiere Deutschlands, p. 324.
1908. A[j}odemus'] agrarius Thomas, Proc. Zool. Soc. London, 1908, p. 7,

June, 1908.
1910. Mus {Apodnmis) agrarius Trouessart, Faune !Mamm. d'Europe,

p. 155.

Tij2)e locality. — Berlin, Germany.

Geographical distribution. — Eastern and central Europe, west
along the Baltic coast to Denmark.

Diagnosis. — Size and external appearance about as in
Apodenms sylvaticus, but colour of upper parts more reddish, and
back with a narrow, sharply defined, black median stripe ; skull
with interorbital region conspicuously beaded, even in half -grown
individuals ; molars of about the same size as in A. sylraticus,

APODEMUS

837

but vr witli no iniiuito anterior cusp on outer side, and lll^ and
■m.^ without cingulum-like outer ledge.

External characters. — The general external form is less slender
and delicate than in the members of the Apodemus syhciticus
group, the ear is shorter, reaching barely to eye, though not
peculiar in structure, the feet are more robust, and the tail is
relatively shorter and more coai'sely annulated. Palms as in
A. syhaticuH, except that the two intermediate tubercles are
relatively smaller. Feet with metatarsal portion relatively
shorter than in A. sylvatkus, but toes showing no peculiarities.
Plantar tubercles rather small, very sharply defined, the postero-
external minute and sometimes obsolete ; space between tubercles
rugose, though rather less so than in A. sylvatkus. Caudal
aunulations very distinct, about fourteen to the centimeter at
middle ; caudal hairs essentially as in A. sylvatkus but less
abundant, the pencil ill-defined. Mannme : p 2 — 2, i 2 — 2 = 8.

Colour. — Upper parts a reddish brown between the russet
and wood-brown of Ridgway, brighter posteriorly, more didl
anteriorly, fading nearly to ochraceous-buft" on sides and cheeks ;
ears and face between eyes and muzzzle noticeably tinged with
hair-brown ; a clear black median stripe about 3 mm. wide
extends from region just behind eyes to within about 10 mm. of
base of tail ; underparts and inner surface of legs dull white
clouded by the slaty undercolour ; tail not bicolor, the general
efi;ect hair-brown somewhat darker above than below ; feet vary-
ing from pale drab to dull whitish. The
young are essentially like the adults, diflfer-
ing only in a slight general dulness of
colour.

SJiull. — In general appearance the skull
resembles that of Apodemus sylvatkus, but
the brain-case is longer and narrower and
the rostrum is shorter and more noticeably
tapering. Profiles essentially as in the
related species, but occiput more obliquely
truncate, so that a greater portion of the
supraoccipital is visible when skull is viewed
from above. Interparietal with antero-
posterior diameter as in A. sylvatkus, but
with lateral extremities abruptly and
almost squarely truncate at region where
bone has width of about 1 • 5 mm., thus con-
siderably reducing transverse diameter ; j^os-
terior border of interparietal lying along lambdoid suture, the outer
anterior angles of the bone therefore encroaching on parietals.*
Margins of interorbital region conspicuously beaded, the raised

* In Mas nmsculus, which has a similarly ligulate interparietal, the
lambdoid suture passes along the anterior border of the bone, so that the
parietals arc not emarginated (fig. 177, p. 866).

Fig. 168.

Ajiodeniun agrarius.

Nat. .size.

838

edge narrow aiul Hbriipi in adults, lower in innnatuie indi\ iduals,
a faint but evident trace visible in sucklings ; posteriorly the
bead is continued along side of upper surface of brain-case as a
slight ridge to lambdal suture, which it meets about 2 mm.
beyond termination of interparietal. Anteorbital foramen some-
what more widely open than in A. sijlvatlcus. Incisive foramina
noticeably shoi'ter and (relatively) Avider than in the other
European members of the genus, the posterior margin extending
barely or not ta level of alveolus of m\ the anterior margin
separated from alveolus of incisor by a distance fully equal to
combined width of foramina. Auditory bullae more inflated than
in Apodemus syJvaticus, so that hamular is nearly or quite in
contact with inflated portion instead of with a contracted beak-
like process as in the related species. While in general
resembling that of Apodemus sylvaticus the mandible is readily
distinguishable by the narrower less curved angular process and
the higher more curved coronoid, the extremity of wdiich is
distinctly above level of condyle.

Teeth. — The teeth are noticeably smaller than in Apodemus
sylvaticus : alveolar breadth of m^ scarcely more than one-third
greatest width of palate between tooth-rows, the narrowness
chiefly due to the relatively small size of the tubercles of the

outer row. Crown of w} slightly
longer than that of m^ and m-^
together, its area decidedly more
than that of the two other teeth
combined. Crescents less curved
than in A. sylvaticus, and showing
less tendency to become joined by
connecting ridges. First upper molar
with t3 reduced to a mere appendage
to the outer border of f2 ; tl well
developed, distinct, very posterior in
position, this peculiarity together with
the small size of t3 producing great
distortion of the anterior crescent ;
no supplemental loops in angles
between ^3 and t'2 or t2 and ^1.
as in A. sylvaticus except for the
flattening of the second crescent and the more abrupt rounding
of the posterior border resulting from the complete absence of
all trace of t9. Second upper molar with no trace of ^3 and
with re-entrant angle on outer surface of <6 so reduced that the
tubercle is often practically entire. Third upper molar as in
A. sylvaticus but with its elements even more indistinct, and tQ
practically absent, the outer border of tooth consequently entire.
Maxillary teeth essentially as in A. sylvaticus, but paired
tubercles less distinct from each other and outer ledge with its
rudimentary cusps in m^ and j/(.j nearly obsolete.

¥lG. KiO.

Apodcinas agrarius.
X 10.

Clieek-teetli.

Rest of tooth essentially

APODEMUS

839

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840 KODKNTIA

MeasHvcmvuts. — Average and extremes of eight adults t'roin
Berlin, Germany: head and body, 08 "1 (97-112); tail, 78 "5
(73-84); hind foot, 19-2 (18-21); ear from meatus, 11-2 (11-
12). Average and extremes of live adults from the vicinity of
Konigsberg, Germany: head and body, 107*8 (99-115); tail,
76 •4^(71-84); hind foot, 19-1 (18 -6-19 -8). Adult male from
Bustenari, Roumania : head and body, 102 ; tail, 82 ; hind foot,
18 "5. For cranial measurements see Table, p. 839.

Specimens examined. — Sixty, from the followiug localities : —

Germany: Berlin, 9 (B.M. and U.S.N.M.) ; Herrenkrug, Magdeburg,
Saxony, 8 ; Altenburg, 1 ; Zoschau, Saxony, 1 (U.S.N.M.) ; Tausha,
Saxony, 1 (U.S.N.M.) ; Bieberstein, Saxonv, 1 ; Saxony, no exact locality, 2
(U.S.N.M.) ; vicinity of Konigsberg, 32 (UlS.N.M.).

Austria-Hungary : Eszek, Slavouia, 1.

Roumania : Bustenari, 1 ; Gageni, 2.

Bulgaria : Rustschuk, 2 (Andersen).

Memarhs. — This animal is immediately recognizable among
European Murid<e by the sharply defined black dorsal line, and
among the smaller members of the family by the noticeably
beaded margin to the interorbital portion of the skull. A similar
colour pattern occurs in the structurally very different Sicisfa
frizona, and as the ranges of the two animals partly coincide
confusion may easily result. Superficially Sicista irizona is
perhaps most readily distinguished from Apodemus agrarius by
the presence of the large, tufted lobe at base of inner portion
of ear capable of completely closing the meatus.

3 6,0 9. Berlin, Germany. Dr. L. Heck (c & p). 7. 6. 12. 1-8.

4 6. Herrenkrug, Magdeburg, Dr. W. Wolterstorfi 92. 12. 1. 10-17.

4 ? al. Saxony. (c & p).

9 Altenberg, Saxony. E. R. Alston (p). 79. 9. 25. 12.
(Moscliler.)

1 al. Bieberstein, Saxony. Berlin Museum (e). 93. 1. 1. 18.

(Nitsche.)

1 al. Eszek, Slavonia, Hun- Budapest IMuseum 91. 7. 23. 1.

gary. (e).

6 Bustenari, Prahova, Rou- Lord Lilford (p). 1. 4. 6. 50.
mania. {W. Doclson.)

6, ?. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 51-52.
mania. [W. Dodson.)

? Purchased (Ver- 63.11.16.3.

reaux).

GexXUs MICROMYS Dehne.

1841. Micromyi Dehne, Micromys agilis, ein neues Saugthier der Fauna

von Dresden, p. 1.
1857. Mus Blasius, Saugethiere Deutschlands, p. 309 (part).
1905. Micromys Thomas, Ann. and Mag. Nat. Hist., 7th ser., xv, p. 442,

May, 1905 (part).

Tiii)e sjiecics. — Micromys ngilin Dehne = 3Ihs soricinus Her-
mann.

GeoQnii)1i'ical distr'ihat'wn. — Central Europe and Asia from

MICROMYS

S41

Great Britain eastward ; in Europe nurili to .Scotland and
Denmark, south to Italy and southern France.

Ding)iosis. — Skull diflering from that of Apodemus in the
greatly reduced rostrum (distance from gnathion to lower edge
of infraorbital foramen less than depth through lachrymal region)
and in the early fusion of nasal bones along median line ; teeth
essentially as in Apodenms, but ni^ with third outer tubercle
obsolete ; ear with large triangular valve capable of completely
closing the meatus ; tail prehensile, bare above at tip ; two
posterior palmar tubercles united in median line.

Heniarlcs. — This genus is well defined by the peculiarities of
the skull, ear, tail and palm. At present one species only is
known, the widely distributed harvest mouse.

MICROMYS MINUTUS Pallas.*
(Synonymy under subspecies.)

Geoijrapltical Jhtrihidioti. — Same as that of the genus
Micro)iii)i<.

D'kkjhusIs. — General characters as in the genus ; size very
small (head and body, 55 to 75 mm. ; hind foot, 1 3 to 16 mm. ;
condylobasal length of skull, 16 to 18 mm.) ; colour above brown
with a yellowish or russet tinge, below whitish or buffy.

External characters. — General form slender and delicate, but
head rather short and round, owing to the reduced condition of
the rostrum. Ear low and rounded, extending barely half way
to eye when laid forward, its height above crown only about
5 mm. ; lower portion of conch with conspicuous triangular
lobe 2 mm. in height capable of completely closing the meatus ;
surface of ear finely pubescent both inside and out, the meatal
valve with a conspicuous tuft of hairs nearly 5 mm. in length.
Feet relatively broader than those of Apodemus si/lvaticiis, but
proportions of digits not peculiar ; palmar and plantar tubercles
essentially as in Apodenuis si/lvaticiis, but posterior plantar
tubercles relatively larger and more elongate, and posterior
palmar tubercles so enlarged that they are confluent along
median line behind, forming with the thumb a single itubercular
mass opposed to the balls of the fingers. Tail preJiensile, about
as long as head and body, thinly haired below, more thinly
above, the distal fifth or sixth of upper surface bare ; no true
pencil, but an inconspicuous tuft usually projects beyond under
side of naked tip; annulations everywhere more distinct and
regular below than above, this especially true of prehensile ter-
minal area where the rings are completely broken up on dorsal
surface ; at middle there are about sixteen to the centimeter.

Colour. — See descriptions of subspecies.

* 1771. Mus minutus Pallas, Raise clurch verschicdenc Provinzen des
Russischen Reichs, i, p. 454 (Banks of the Volga, Russia).

842 EODENTIA

Skull. — The .skuU ditVers conspicuously tVoiii tliui of ;ill otlior
l*iUi'opeau Murimti in the striking (-ontrast between the short
weak rostrum and the hirge, much elongated brain-ease ; distance
from upper border of infraorbital foramen to tip of nasal less
than one-third that from foramen to posterior border of occiput,
while in other members of the sub-family it is nearly one-half.
Dorsal profile gradually rising and nearly straight from tip of
nasals to interorbital region (sometimes with a faint angle at
middle of nasals), then broadly convex, slightly more gradually
anteriorly than posteriorly, to interparietal, where a slight
break in the profile is produced by the more abrupt convexity
of the overhanging posterior portion of brain-case ; ventral
profile without special features, essentially straight from incisor
to condyle, except for the slight break caused by the auditory
bulla:;. Brain-case oval in outline when viewed from above, a
little flattened posteriorly, the surface very smooth and evenly
inflated ; posterior portion unusually dis-
tinct from main part of brain-case, and
almost marked off by a constriction at
^TN^^^ front of interparietal ; in posterior view

^^__^ the middle portion of brain-case rises

unusually high above interparietal, and
distance from lambdoid suture to upper
margin of foramen magnum is unusually
reduced ; paroccij)ital processes very short,
extending scarcely to level of lower lip
of foramen ; floor of brain-case with no
special features, the median ridge and
Via. 170. lateral furrows well developed, the anterior

Micromys minuUis. •ij_t o ^ • • -i ^ t n ^•

Nat. size. Width 01 basioccipital less than median

length ; auditory bullae rather large, evenly
inflated, without evident beak, the meatus relatively large,
without lip except along anterior border. Interorbital region
rather wide and short, smoothly rounded off at sides though
tending to become slightly angular posteriorly in old indiAdduals.
Zygoma .slender and weak, the arch without special features,
the horizontal plate forming outer wall of anteorbital foramen
rather narrow, producing a very inconspicuous forward projection
on anterior border of zygomatic root when viewed from above, its
own anterior border faintly concave and sloping obliquely forward
from above downward. Nasals not peculiar in general form,
their posterior border together with that of nasal branches of
premaxillaries deeply and irregularly lacinate,* not extending
noticeably behind lachrymal level. Incisive foramina relatively
larger than in Apodemus sylvaticus, their outer border nearly
parallel, the posterior border extending to level of anterior root
of m^, the anterior border separated from incisor by space equal to
greatest width of single foramen. Anterior portion of palate
* Not successfully represented in fig. 170.

3IICR0MYS

SV.

witliuut special peculiarities ; a sliglitly indicated transverse
ridge at level of posterior border of last molar ; niesoptei-ygoid
space beginning about 1 mm. behind this ridge, much narrower
anteriorly than posteriorly, its margins straight to tip of haniulai-,
which is barely or not in contact with auditory bulla. Mandi})le
essentially like that of Apoderuus syhmticus, but with barely
indicated swelling over root of incisor, more concave upper
margin of angular process, and coronoid more abruptly hooked
backward.

Teeth. — General character of teeth much as in Ajjoihnuus
.tylvaticus, but upper incisor perhaps a little more compressed
and root of lower incisor scarcely forming an appreciable
protuberance on outer side of mandible. Relative size of upper
molars more nearly as in Apodcmus sijli-aticus than as in
A. agrarius, but outline of m^ intermediate between that in
these two species. Inner and outer tubercles about equal in size.
First upper molar with anterior crescent
somewhat distorted, t3 well developed
though closely crowded against t'2, tl some-
what posterior in position, the three tuber-
cles less contrasted in size than in the species
of Apodemus ; no intermediate folds in
narrow angles between bases of tubercles,
but a rather well defined ridge from tl to
inner base of to ; /6 very large, filling nearly
tlie entire space occupied by it and /9 in
Anodemus sulvaticus : <9 reduced to a mere ,,. J'' '.' ,

rudiment at posterior base of t% ; postero Cheek-teeth, x lo.
external margin of crown with slight thick-
ening of the enamel as in Apodemus sylvaticus ; tS and tl showing
no special peculiarities. Second upper molar with first crescent
represented by a minute though evident <3 and a small tl ; second
crescent with simple /6 about equal in size to tA ; third crescent
with barely indicated t2 and small tl . Third upper molar much
as in Apjodemus sijlvaticus, its crown showing traces of tubercles
1, 4, 5, 7 and 8. Maxillary molars as in Apodemus syhaticus,
but outer ledge of m^ and m., slightly developed and without
evident cusps.

Remarks. — Two well marked races are recognizable among
the harvest mice of western Europe. True Micromys mimttus is
apparently not found in this region.

KEY TO THE EUROPEAN FORMS OP MICEOMYS MINUTUS.

Head and shoulders reddish brown, essentially

concolor with back (England and western

continental Europe) -"■. mitmtus aoricinua, p. 844.

Head and shoulders greyish, noticeably contrasted

with reddish brown back (Hungary and Rou-

mania) H- »»• iirateasU, p. 846.

8U

MicuuMvs MiNUTUS soKicixus Hermann.

1780. Mits soricimis Hermann in Schrobcr, Siiugcthicre, iv, p. 6G1,

pi. CLXXXiiiB. (Strassbiirg, Germany).
1785. [Mils] triticciis Boddaert, Elcnchus Anim., i, p. Ill (Harapshirc,

England).
1789. Mils minimus White, Nat. Hist, and Antiqu. of Selborne, p. 43 ;

described, pp. 33, 34 and 39 (Selborne, Hampshire, England).
1792. Mus messorius Kerr, Anim. Kingd., ^. 230 (Hampshire, England).
1794. '■'Mus ave?iarius Wolf, Versuche die Feldmiiuse zu vertilgen, p. 16,

315," Hermann, Observ. ZooL, p. 61, 1804 (Locality not men-
tioned).
1804. Mus pendulinus Hermann, Observ. Zool., p. 61 (Strassburg, Germany).
1804. Mus parvul'Us Hermann, Observ. Zool., p. 61 (Strassburg, Germany).
1816. ? Mus arvensis Leach, Syst. Catal. Spec. Indig. ]\lanim. and Birds,

Brit. Mus., p. 7 (Nomen nudum: "Harvest Rat").
1822. Mus campcstris Desmarest, Mammalogie, pt. 2, p. 543 (France).
1832. M[us\ meridionalis Costa, Fauna del Regno di Napoli, p. 13 (Vicinity

of Naples, Italy).

1840. Mus minatus Schinz, Europ. Fauna, i, p. 70.

1841. Micromys agilis Dehne, Micromys agilis, cin neues Siiugthicr der

Fauna von Dresden, p. 1 (Dresden, Germany).

1841. Mus orijzivorus de Selys-Longchamps, Atti della seconda Riunione

degli Scienziati Italiani, Torino, 1840, p. 247 (Rice fields of
Lombardy, Italy).

1842. Mus puinilus F. Cuvier, Hist. Nat. des Mammif., Tabl. gen. et

M6th., p. 4. Described in fasc. xxxii, October, 1821 (Vicinity of

Paris, France).
1857. Mus mvmctus Blasius, Siiugethiore Dcutschlands, p. 326.
1900. Mas minutus agilis Barrett-Hamilton, Ann. and INIag. Nat. Hist.,

7th ser., v, p. 529, June, 1900 (Brunswick, Germany).
1900. Mus minutus campestris Barrett-Hamilton, Ann. and ]\Iag. Nat.

Hist., 7th ser., v, p. 529, June, 1900 (Waremme, Liege, Belgium).
1900. Mus minutus minimus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., v, p. 530, June, 1900 (England).
1910. Mus [Apodemus) minutus agilis, M. {A.) minutus campcstris, M. (A.)

minutus minimus and M. (A.) meridionalis Trouessart, Faune

Mamm. d'Europe, pp. 156, 157.

TijiK' locality. — Strassburg, Germany.

Geographical distrihution. — Central Europe from Great Britain
to eastern Germany, and from Scotland (Aberdeenshire) and
Denmark south to Italy and southern France.

Diagnosis. — Colour of upper parts essentially uniform through-
out, the shoulders, neck and head never conspicuously greyish.

Colour. — AVinter pelage (England and Switzerland) : upper
parts a nearly uniform reddish brown approaching the tawny of
K-idgway but more vivid, especially along median region ; neck
and head sometimes not so bright as back, but never sufficiently
so to produce any marked contrast ; sides of body and outer
surface of legs fading toward ochraceous-bufF ; back very
inconspicuously sprinkled with black-tipped hairs ; ear concolor

MICROMYS 845

with sides ; imderparts and inner surface of legs dull white to
base of hairs, or with a slaty under-tint, the line of demarcation
along sides sharply defined ; feet, a light, indefinite yellowish
brown ; tail obscurely bicolor, the hairs everywhere ochraceous-
buft', the skin light brown below, dark above. Summer pelage
(Germany) : upper parts noticeably darker and duller than in
winter, approaching the russet of Ridgway, but with a tinge of
rufous ; sides dull cinnamon ; underparts didl ochraceous-buff,
the line of demarcation along sides inconspicuous, the median
line, especially of chin and throat, often with traces of white in
the form of ill-defined lines and blotches ; feet and tail as in
winter, but more thinly haired and therefore more obscurely
coloured.

Measurements. — Two adult males from Norfolk, England :
head and body, 64 and 65 ; tail, 61 and 58 ; hind foot, 15 and
14; ear from meatus, 9 and 8. Adult male and female from
Waremme, Belgium : head and body, 68 and 70 ; tail, 60 and
65; hind foot, 14 and 16. Average and extremes of ten adults
from Brunswick, Germany: head and body, 68 "3 (63-76);
tail, 59-7 (51-70); hind foot, 14-9 (14-16). Adult male from
Pavia, Italy; head and body, 63 ; tail, 72 ; hind foot, 15'4; ear
from meatus, 10. For cranial measurements see Table, p. 847.

Specimens examined. — Eighty-seven, from the following localities : —

England : Haddiscoe, Norfolk, 11 ; Swaffham, Norfolk, 2 ; Methwold
Fen, near Brandon, Norfolk, 1; Congham, Norfolk, 1; eastern Norfolk, 3;
Norfolk (no exact locality), 3; Cambridgeshire, 1; Colchester, Essex, 6;
Kingsbury, Middlesex, 1 ; Somersetshire, 1 ; Totton, New Forest, 1 ;
Hampshire, 1 ; no exact locality, 2.

France : Etupes, Doubs, 5 (Mottaz) ; Rugles, near Evreux, Eure, 2
(U.S.N.M.).

Belgium: Waremme, Liege, 3 (U.S.N.M.).

Germany: Holstein, 1 ; Brunswick, 27 (U.S.N. M.) ; near KiJnigsberg, 9
(U.S.N.M.) ; Strassburg, 1.

Switzerland: St. Fiden, St. Gallen, 1 (U.S.N.M.).

Italy : Pavia, 4 (Ghidini).

Remarks. — For the present it seems necessary to regard all
the harvest mice of western Europe, including England and
northern Italy, as belonging to a single race. The differences
which have been observed to exist, notably between British and
German specimens, prove to be merely seasonal.

4:6,9 Haddiscoe, Norfolk, Eng- G. Barrett-Hamilton 11. 1. 2. 124-128.

land. (L.C.Forman.) (p).

1. Congham, Norfolk. OxleyGrabham(c&p). 11. 1. 3. 414.

3 6. Norfolk. G. Barrett-Hamilton 11. 1. 2. 129-131.

{Oxley Grahham.) (p).

1 Cambridgeshire. J. Baker (c & p). 39. 9. 29. 31.

4 c?, 2 9. Colchester, Essex. Dr. Henry Laver 3. 4. 8. 1-6.

(c & p).

1. Kingsbury, Middlesex. E. R. Alston (p). 79. 9. 25. Gl.
{J. L. Harting.)

1 al. Totton, Hampshire. E. B. Wharton (c & p). 83. 4. 30. 1.

846

RODENTIA

1. Somerset. {Hiigel) E. R. Alston (p). 79'. 9. 25. 62.

1. Holstein, Germany. Purchased (Brandt). 47. 4. 5. 2.

1. Strassbm-g, Alsace. C. Mottaz (c). 8. 8. 10. 90.

MiCROMYS MINUTUS PRATENSIS Ockskav.

1831. Mus pratensis Ockskay, Nova Acta Phys.-Med. Acad. Caes. Leop.-
Carol. Nat. Cur., xv, pt. ii, p. 243, pi. lxviii (Western Hungary).

1882. Mus arundinaceus Pet6nyi, Termeszetrajzi Fiizetek, v, p. 142.

1900. Mus 77iinutus pratensis Barrett-Hainilton, Ann. and Mag. Nat. Hist.,
7th ser., v, p. 580, June, 1900.

1910. Mus (Apodemus) minutus pratensis Trouessart, Faune Mamm.
d'Europe, p. 157.

Type locality. — Western Hungary.*

Geograpical distribution. — Hungary and Roumania ; limits of
range not known.

Diagnosis. — Similar to Micromys minutus .soricinus, but head
and anterior half of body decidedly greyish.

Colour. — Posterior half of back together with outer surface
of hind legs as in 31. soricinus. Head and anterior half of body
a buffy grey, darker and more brownish along median dorsal
region, lighter and more nearly smoke-grey on sides of neck and
shoulders, deepening gradually to a buffy ochraceous posteriorly.
Underparts and inner surface of legs sharply defined creamy
white, to which the slaty basal portion of the hairs imparts a
bluish tinge. Feet a dull indefinite buffy brown. Tail rather
sharply bicolor, dark brown above, pale buffy below.

Measurements. — Adult female from Gageni, Roumania : head
and body, 63; tail, 55; hind foot, 14*5; ear from meatus, 9.
Four adults from Csallokoz-Somorja, Pressburg, Hungary : hind
foot, 13-2, 13-6, 14 and 14.

Specimens examined. — Nine, from the following localities : —
AusTRiA-HuNGABY : Csallokoz-Somorja, Pressburg, Hungary, G ; Cepin,
near Eszek, Slavonia, 2 (in spirit).
Roumania : Gageni, Prahova, 1.

Bemarls. — The few specimens examined indicate that this is
a well-defined race.

G. Csallokoz-Somorja, Press- Budapest Museum (e). 94. 8. 1. 53-58.
burg, Hungary.
2al. Cepin, Slavonia. Budapest Museum (e). 94. 7. 23. 11-12.

9 Gageni, Prahova, Ron- Lord Lilford (p). 4. 4. G. 49.

mania. [W. Dodson.)

* "Frequentius occurrit in Comitatus Nittrienses, valle Vagi."

847

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848

Genus EPIMYS Trouessart.

1857. Mus Blasius, Saugethiere Deutschlands, p. 309 (part).

1867. Battus Fitzinger, Sitzungsber. kais. Akad. Wissenscb. Wien, Matb.-

■ Naturwiss. Classe, lvi, p. 63 (not of Donovan, 1827).
1881. Epimys Trouessart, Bull. Soc. d'Etudes Sci. d' Angers, x. p. 117

(sub-genus). Type by subsequent selection Mus rattus Linnseus.
1910. Ejyimys Miller, Proc. Biol. Soc. Wasbington, xxiii, p. 58, April 19,
1910.

Type species. — Mus rattus Linnaeus.

Geographical distribution. — Originally confined to the temperate
and tropical portions of the Old World, but now essentially
cosmopolitan through the artificial dispersal of two species.

Characters. — External form, skull, and teeth with no special
modifications ; molars slightly graduated in size from first to
third, the anterior tooth not tending to assume the main
function of the tooth-row, the posterior tooth not tending to
disappear ; enamel foldings of upper molars directly referable to
a simple 9-cusped pattern and its reductions, the outer margin of
m^ and m^ never with more than three cusps, the inner margin
of the same teeth never with more than two cusps ; m^ with five
roots, its first lamina not distorted by the backward displacement
of tl ; upper incisor compressed, set at such an angle that its
outer side is worn smoothly away by action of lower teeth.

Bemarlcs. — The genus Epimys is the most widely distributed
and abundantly represented group in the sub-family Murinse.
As here defined it contains the great mass of the Asiatic,
Malayan and African species, several hundred in number, to
which the generic name Mus has during recent years gradually
become restricted. Only two of these occur in Europe.

KEY TO THE EUROPEAN FORMS OF EPIMYS.

Greatest breadtb of brain-case across lateral ridges
about equal to length of parietal measured
along ridge; first lamina of vi'- and terminal
heel of vi" with no well developed outer

tubercle ; external form rather heavy ; tail *
decidedly shorter than head and body (Nor-
way rat) E. norvcgicus, p. 858.

Greatest breadth of brain-case across lateral ridges
decidedly more than length of parietal mea-
sured along ridge ; first lamina of m^ with
distinct outer tubercle usually almost as large
as inner tubercle ; terminal heel of m- with
evident though incompletely separated outer
tubercle ; external form slender ; tail about
equal to head and body (usually a little longer) E. rattiis, p. 849.
General colour slaty black, the underparts dark

slaty grey (Distribution general) E. r. voYriw, p. 853.

General colour brown, the underparts buffy or

greyish (Southern) 7-;. r. alexandiniu^, p. 854.

EPIMYS RATTUS Liinutnis.

(Synonymy under subspecies.)

Geoyraphical distrihntlon. — Originally confined to the north
temperate portions of the Old World ; now essentially cosmo-
politan through artificial dispersal.

Didgnosis. — Skull with brain-case broad, its greatest breadth
across lateral ridges decidedly more than length of parietal
measured along ridge ; anterior upper molar without trace of
cingulum at anterior border of crown ; first lamina of m^ with
distinct outer tubercle usually almost as large as inner tubercle ;
terminal heel of m'^ with evident though incompletely separated
outer tubercle ; first and second lower molars usually with
evident small supplemental tubercles at outer extremity of
furrows ; form slender ; tail about as long as head and body
(usually somewhat longer) ; ear when laid forward extending
about to middle of eye ; hind foot of adults usually under
10 mm. in length ; condylobasal length of adult skulls usually
08 to 43 mm.

External cluiradcra. — General form rather slender, iJio tail
usually longer than head and body and never, unless injured,
noticeably shorter. Head slender, tapering anteriorly, the ear
rather large, extending about to middle of eye wlien laid
forward, its substance thin and somewhat translucent, its
general outline broadl}^ oval, the basal portion of anterior border
folded backward (inward) ; posterior border of meatus witli
barely indicated ridge ; surface of ear finely papillose and very
inconspicuously pubescent. Muzzle pad small but well defined,
with deep median groove continuous with cleft in upper lii? ;
nostril crescentic, its inner and lower margins swollen, forming-
an ill-defined, wart-like excrescence below. Feet moderately
large, with no special pecidiarities of form. Front foot with
inner digit reduced to a mere tubercle with rudimentary nail not
extending to its edge ; outer digit slightly shorter than second,
the third and fourth sub-equal, slightly longer than second ; pads
five, large, distinct, that at base of outer toe with slightly
developed supplemental tubercle at outer base. Hind foot with
inner digit extending to middle of first phalanx of second di"it,
outer digit somewhat longer, the second, third and fourth
sub-equal and longest ; pads six, the postero-internal more than
twice as long as broad, widest in front, narrowing posteriorly,
the posteroexternal well developed, fully one-third as large as
that at base of outer toe ; pads at bases of inner and outer toes
with rudimentary supplemental outer tubercles ; surface of sole
naked, smooth except between pads, where it is noticeably
wrinkled. Claws simple, curved, those on hind fec;t largest.
Tail slightly four-sided, uniformly and very thinly haired
throughout, the tip with no distinct pencil ; annulations well

3 1

850

KOPENTIA

(Icfiiicd, rogulai', al)out niuu tt) the ccntiineter at middle ; leufjth
of hairs equal to about one and one-half width of rings. Fur
soft but interspersed with numerous coarse hairs Avhich impart
to it a somewhat harsh quality ; with a lens these coarse hairs
are seen to be very slender bristles, distinctly grooved on the
upper side. Mammiy : p 2 — 2, i 3 — 3 = 10.

Colour. — The two main types of colour are (1) dark slaty,
becoming almost blackish on back, and (2) light brown above,
pale buff or light grey below. All possible intermediate stages
exist connecting the two extremes.

Shill. — In general features the skull shows no marked
peculia-i'ities. Dorsal profile faintly convex throughout, but

FIG. ITi'.
Epiinys raltua. Ivat. size.

flattened over posterior half of nasals, between orbits and on
l^osterior half of brain -case ; occiput s(]uarely and almost perpen-
dicularly truncate ; ventral profile nearly flat. Brain-case squarish
in outline when viewed from above, bvit slightly longer than
wide and somewhat narrowed posteriorly, the sides of its vipper
sui'face marked by conspicuous longitudinal ridges continuous
with the overhanging interorbital beads. The greatest width
of brain-case across these ridges is distinctly greater than length
of parietal measured along ridge. No trace of sagittal crest,
but a low ridge-like lambdal crest present in old individuals and

KPIMYS Sol

a kuife-likc median occipital crest from interparietal to foramen
magnum. Interparietal large, its anterior border nearly straight,
though usually forming a median point, its posterior border
strongly convex. Occiput rather broad and low when viewed
from behind, but central portion of brain-case not rising con-
spicuously above it ; paroccijjital processes slender and rather
long, extending beyond level of lower lip of foramen. Floor of
brain-case with no special features ; width of basioccipital along
anterior border scarcely half median length ; auditory bulla
moderately large, evenly inflated, its greatest diameter about
8 mm., meatus with thickened edges but not tubular. Inter-
orbital region contracted, its least breadth less than that of
rostrum over roots of incisors, its margin overhanging and
conspicuously beaded, the beads passing into lateral ridges of
brain-case behind a slight though evident angle at suture between
frontal and parietal. Zygoma rather slender, not expanded or
noticeably flattened, strongly bent downward, its lowest point
usually opposite m-^ ; anteriorly the arches are not abruptly
spreading, so that greatest zygomatic breadth is at glenoid level ;
plate forming outer border of anteorbital foramen well developed,
its anterior border vertical below, broadly rounded above.
Rostrvim much deeper than broad but without special peculiarities
of form ; nasals truncate or bluntly pointed posteriorly, extend-
ing barely to lachrymal level, somewhat exceeded by nasal
branches of premaxillaries. Incisive foramina parallel sided,
rather short and wide, their posterior border extending to level
of anterior root of wi\ their anterior extremity falling short of
incisors by a distance equal to about one and one-half times the
width of the two foramina together. Palate extending notice-
ably behind level of m'-^. Mesopterygoid space slightly wider
posteriorly than anteriorly, truncate or double rounded in front,
the hamulars slightly turned outward, not in contact with
bulht ; ectopterygoid well developed, tilted noticeably upward
from within outward (skull viewed from below). Mandible
robust, Avith no special peculiarities ; coronoid process directed
strongly backward, slightly curved, its extremity rising notice-
ably above level of condyle.

Teeth. — Teeth moderately large relatively to size of skull ;
upper molars with outer tubercles well developed, nearly as large
as those on inner side, the crescents rather flat but evident
and showing little tendency to distortion. Incisors stronglv
compressed, without special peculiarities, the cutting surface of
upper tooth vertical, flat, its edges entire. First upper mohii-
5-rooted, the anteiior root largest, strongly pr-ojecting forward
beyond base of crown, the median inner and median outer roots
smallest ; crown barely as long as that of m'-^ and i/t-' together,
its area about equal to that of the succeeding teeth ; anterior
border of crown smooth, with no trace of cingulum or minute
supplemental cusps; first lamina moderately and evenlv crescentic,

3 1 2

852 RODENTIA

the outer tubercle nearly as large as Zl, and only a little anterior
to it in position, both small tubercles -well detiued from t2, the
re-entrant angles deep and without supplemental folds ; second
lamina essentially like first, not connected with it by longitudinal
ridges, /6 entire ; third lamina completely distinct from second,
/9 well developed, about as large as /3, <7 absent. Second upper
molar with first lamina represented by large tl ; second and third
laminae essentially as in m^, the third with well developed /9
nearly as large as that of preceding tooth and marked off from
tS by a distinct re-entrant angle. Third upper molar with large,

distinct, terete tl, small ti and tb, the
latter joined by an obsolete tQ to outer
extremity of /8, the only remnant of
the third lamina ; as the crown wears
away the second and third laminae
assume the form of a narrow loop, the
two limbs of which are separated by a
deep re-entrant angle. Anterior lower
molar with crown scarcely as long as
those of the two succeeding teeth, its
area also less ; first lamina narrower
Fig. 173. than second, its posterior border nearly

£piinij!i mttus. Cheek-teeth. straight, its anterior border with deep
re-entrant angle on outer side of
middle ; second and third laminye essentially alike, more strongly
curved posteriorly than anteriorly, the division into two tubercles
ill defined ; terminal heel small, compressed ; a small terete
tubercle usually present at outer edge of space between second and
third lamina, and a similar though smaller tubercle occasionally
present in that between first and second. Second lower molar
essentially like first without anterior lamina, the outer border with
small supplemental tubercle at anterior base of first lamina, and
another usually less well developed in space between laminae.
Third lower molar with two lamina?, the anterior of which shows a
faint indication of division into two tubercles, the posterior similar
to the posterior heel of the other teeth but considerably larger.

Bemarks. — Though now so completely intermingled in many
localities that exact determination of specimens is often im-
possible, there seems no good reason to doubt that two geo-
graphical races of Epimys rattiis have established themselves in
western Europe. True rattus, the blackish, slaty-bellied form,
is a northern animal, normally occurring in a climate similar
to that of central and northern Europe. The buff-bellied
form, Epimi/s rattus alexandrinus, is normally more southern in
its range, finding its optimum conditions, so far as Europe is
concerned, in the Mediterranean region."' Individuals of each

* In the United States it appears never to become naturalized north of
the limits of the Lower Austral life zone, while E. rattus is or has been
locally common in the Transition and Canadian zones.

EPIMYS 853

form have been artificially though unintentionalh^ transplanted
into the area of the other, and to this circumstance, rather than
to individual variation or normal intergradation, should be
attributed the intermediate specimens now so abundantly found.

EplMVS KATTIIS RATTUS Linna'US.

1758. [Mns] rattus Linnteus, Syst. Nat., i, 10th od., p. Gl (Sweden).

1833. Mus tectorum var. fuliginosus Bonaparte, Iconogr. Faun. Ital., i,

fasc. 3, pi. 22, fig. 1, name on plate only (Italy, no exact locality).
1842. Mus subarrulciis Lesson, Nouv. Tabl. du Regne Anim., ^lamm.,

p. 138 (Rochefort, Charente-Inferieure, France).
1857. Mus rattus Blasius, Saugethiere Deutschlands, p. 317.
1867. Rattus domesticus Fitzinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Matb.-Naturwiss. Classe, lvi, pt. i, p. 64 (Substitute for

rattus).
1867. [Rattus domesticus] fuscus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-lSTaturwiss. Classe, lvi, pt. i, p. 64

(Germany).
1867. [Rattus domesticus'] varius Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 64

(Germany).
1867. [Rattus domesticus] fitlvaster Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 64

(Austria and Germany).
1867. [Rattus domesticus] albus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Matb.-Naturwiss. Classe, lvi, pt. i, p. 65

(Austria, Hungary, Germany).
1867. [Rattus domesticus] ater Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, lvi, pt. i, p. 65 (Germany).
1902. Mus alexandrino-rattus Fatio, Revue Suisse de Zoologie, x, p. 402,

December 30, 1902 (Ticino, Switzerland). See Mottaz, Bull. Soc.

Zool. de Geneve, i, p. 163, November 35, 1908.
1905. Mus rattus ater IMillais, Zoologist, 4th ser., ix, p. 205, June, 1905

(London, England). Type in British Museum.
1908. Epimys rattus Satunin, INIitth. Kauk. Mus., Tiflis, iv. Lief. 1-2,

p. 112.
1910. Mus {Epimys) rattus and M. (E.) rattus ater Trouessart, Faune

Mamm. d'Europe, p. 143.

Ti/])e locality. — Upsala, Sweden.

Geographical distribution. — Europe north of the Mediterranean
region ; occurring in isolated colonies and everywhere being
replaced by Epimys norvegicns.

Diagnosis. — General colour slaty, the back darker, frequently
almost black.

Measurements. — Adult male and female from South Ronald-
shay, Orkney Islands: head and body, 189 and 188; tail, 20G
and 205; hind foot, 36-7 and 36; ear, 24-5 and 24. Adult
male from Lundy Island, Devonshire : head and body, 209 ; tail,
233; hind foot, 40; ear, 25-5. Adult male from London,
England (type of ater Millais) : head and body, 235 ; * tail, 235 ; *

* From :\Iillais.

8")4 RODENTIA

hind foot, 36 ; ear, 2;").* Adult female from Hatszeg, Hnn3'ad,
Austria-Hungary : head and body, 190 ; tail, 225 ; hind foot, ."nS.
For cranial measurements see Table, p. 856.

Specimens examined. — Twenty, from the following localities : —
Scotland : South Eonaldshay, Orkney Islands, 5 (Edinburgh).
England : Great Yarmouth, Norfolk, 1 ; Tring, Hertfordshire, 1 ;
London, 6 (including type of ater Millais) ; Lundy Island, Devonshire, 2.
France: Sark, Channel Islands, 1.
Switzerland : No exact locality, 1.
Austria-Hungary: Hatszeg, Hunyad, 2.
Portugal : San Miguel, Azores, 1.

(5 St. Great Yarmouth, Norfolk, W. R. Butterfield 4.1.22.1.

England. (c & p).

9 St. Tring, Hertfordshire. Hon. N. C. Roths- 1. 5. 22. 8.

child (c & p).

2 St. London. A. Stedall (c & p).

<5, lal. London. C. J. Wilson (c & p). 86. 4. 5. 1-2.

St. London. E. Bidwell (c & p). 2. 4. 4. 1.

6. London. J. G. Millais (p). 5. 7. 28. 1.

(Type of afcr Millais.)

2 6. Lundy Island, Devonshire. T. A. Coward (c & p). 7.12.19.1-2.

9. Switzerland. (Nager.) E. R. Alston (p). 79. 9. 25. 40.

6,9. Hatszeg, Hunyad, Tran- C. G. Danford (c). .8.2.2.22-24.
sylvania.

6. San Miguel, Azores. Hon. W. Rothschild 3. G. 5. 15.
(W. B. 0. Grant.) (p).

Epimys rattus alexandrinus Geoffroy.

180-3. Mus alexandrimis Geoffroy, Catal. IMammif. du Mus. Nat. d'Hist.

Nat., Paris, p. 192 (Egypt).
1814. Miiscnliis fritgivorus Rafinesque, Precis des Ducouv. et Travaux

Somiologiques, p. 13 (Sicily).
1825. Miis tectorum Savi, Nuovo Giorn. de' Lettcrati, Pisa, x, p. 74 (Pisa,

Italy).
1827. Myoxus siculn: Lesson, Man. de Mammalogie, p. 274 (Substitute for

Musculus frugivorus Rafinesque).
1841. Miis sylvestris Pictet, ]Mem. Soc. Phys. et d'Hist. Nat., Geneve, ix,

p. 153 (Near Geneva, Switzerland). Alternative for Icncogastcr.
1841. Mus leucogaster Pictet, jNIem. Soc. Phys. et d'Hist. Nat., Geneve, ix,

p. 154 (Near Geneva, Switzerland).
1841. [Mils'] nemoralis de Selys-Longchamps, Atti della secouda Riunione

degli Scienziati Italian!, Torino, 1840, p. 247 (Near Geneva,

Switzerland). Accidental substitute for sylvestris Pictet.
1845. Mus picieti Schinz, Synops. Mamm., ii, p. 142 (Substitute for leuco-
gaster Pictet).
1857. Mus alexandrinus Blasius, Siiugethiere Deutschlands, p. 316.
1882. [Mils 7-atttis] intei'mcditis Ninni, Atti del reale Inst. Veneto, 5th ser.,

VII, p. 574 (Venice, Italy).
1910. Mus [Epimys) rattus intermedius and M. (E.) rattus alexandrinus

Trouessart, Faune IMamm. d'Europe, pp. 143, 144.

Tifpe locality. — Alexandria, Egypt.

(Tt'oi/raphical disfrihution. — Mediterranean region ; eveiy where
aljundant and not tending to be replaced by Epltinix norv^gicus.

* From Millais.

855

D/rtr//(Os/.s.— Upper p;ii'ts light brown ; underparti; strongly
contrasted pale buff or light grey.

Measurements. — Two adult females from near Ninies, Gard,
France: head and body, 180 and 199 ; tail, 222 and 250; liind
foot, 36 • 5 and 38 • 5 ; ear, 24 and 27. Two females from Sorrento,
Italy: head and body, 177 and 178; tail, 243 and 217; hind
foot"^ 37 and 37. Average and extremes of ten adults from
Sorrento, Italy: head and body, 190-7 (181-202); tail, 221
(203-244) ; hind foot, 36 (35-37). Average and extremes of
six adults from Palermo, Sicily: head and body, 202-5 (196-
216); tail, 234 (220-258); hind foot, 35-8 (34-38). Female
from' Silos, Burgos, Spain : head and body, 176 ; tail, 200 ; hind
foot, 36 ; ear, 26. For cranial measurements see Table, p. 8o6.

Specimens examined.— One hundred and twenty-three, from the follow-
ing localities :— -,- -, t i t -ta

England: Sunderland, Durham (on ship), 1; Lundy Island, Devon-
shire, 6. T- . O -D'- O 1

Spain- Pajdres, Leon, 2; Arrechavaleta, Vitoria, 2; Bejar, Sala-
manca 2 ; Silos, Burgos, 4 ; Castrillo de la Reina, Burgos, 1 ; Jerez, 2 ;
Venta del Baul, Granada, 1; Alcoy, Alicante, 1; Barracas, Castellon, 1;
Panticosa, Huesca, 1 ; San Cristobal, Minorca, Balearic Islands, 2.

France: Biarritz, Basses-Pyrenees, 7; near Nimes, Gard, 17; Agay,
Var, 3; Barcelonnette, Basses-Alpes, 1 (U.S.N.M.). •

Switzerland : Lugano, Ticino, 1 (U.S.N.M.) ; Agra, Ticmo, 1 (U.S.N. .M.).

Italy : Giamutri Island, Tuscany, 1 ; Florence, 9 (U.S.N.M.) ; Palermo,
Sicily, 30 (B.M. and U.S.N.M.) ; no exact locality, 1.

Montenegro : No exact locality, 3.

Greece: Corfu, 3; Argostoli, Cephalouia, 14; Kephissia, Athens, 4;
Canea, Crete, 2.

3<J,29.
6,9.

6,9.

2 9.

2 6.

6.
6,9.

9.

6.

9.
6,9.
6,9.

6,29.

9.

1.
6,'69.

Sunderland, Durham (on

board ship), England.
Lundy Island, Devonshire.
Pajares, Leon, Spain.

{N. Gonzalez.)
Arrechavaleta, Vitoria.

{N. Gonzalez.)
Bejar, Salamanca.

(N. Gonrmlez.)
Silos, Burgos.
Castrillo, Burgos.
Jerez, Cadiz.
Alcoy, Alicante.

{N. Gonzalez.)
Barracas, Castellon.

{N. Gonzalez.)
Panticosa, Huesca.

[N. Gonzalez.)
San Cristobal, ]\Iinorca,

Balearic Islands.
Biarritz, Basses-Pyrenees,

France.
Agay, Var.

Giamutri Island, Tuscany.
Italy (Savi).
Palermo, Sicily.

W. B. Tegetmeier

T. A. Coward (c & p).
0. Thomas (p).

O. Thomas (j').

O. Thomas (p).

N. & S. Gonzalez (c).
N. & S. Gonzalez (c).
A. Chapman (p).
O. Thomas (p).

0. Thomas (p).

O. Thomas (i-).

O. Thomas and R. I.

Pocock (c & p).
J. F. Davison (c & p).

G. S. Miller (c).
Genoa Museum (e).
Crisp Collection.
J. I. S. NVhitaker
(c & p).

89. 1. 28. 1.

7. 12. 19. 3-0.

8. 2. 9. 98-99.

8. 2. 9. 94-95.

8. 2. 9. 90-97.

8. 7. 7. 22-23.
8. 7. 7. 24.
8. 3. 26. :3- 4.
8. 2. 9. 101.

8. 2. 9. 102.

8. 2 9. lOf).

0. 7. 1. 50-51.

0. 1. 21. 1-2.

8.8.4.224-220.
7. 2. 28. :3.
84. 6. 3. G.
98. 10. 0. 8 11.

856

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858 RODENTIA

3 6. Montenegro. (L. Filhrer.) O. Thomas (p). 5. 8. 4. 7-9.

5 6,59. Argostoli, Cephalonia, J. I. S. Whitaker (p). 8. 10. 1. 31-37.

Greece. (C. Mottas.) 40-42.

3 (5, 9 Kepbissia, Athens. Hon. N. C. Roths- 8. 10. 2. 40-43.

(C. Moifaz.) child (p).

6, 9. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 4-5.
(C. H. B. Grant.)

EPIMYS NORVEGICUS Erxleben.

1777. Mas norvegicns Erxleben, Syst. Regni Anini., i, p. 381 (Norway).

1778. Mus decumanus Pallas, Nov. Spec. Quadr. Glir. Ord., p. 91 (Europe).

1779. M['H..s] surmolottus Severinus, Tentamen Zool. HuugaricDe, p. 73

(Central Europe).
1800. ilf[Hs] cl[ecmnanus] liybrichis Bechstein, Pennant's Allgom. Ueber-

sicht d. Vierfiiss. Thiere, ii, p. 713, described on p. 497. Walters-

hausen, Germany (melanistic speciinen).
1816. Mus caspius Oken, Lehrb. d. Naturgesch., iii, pt. ii, p. 895.

(Alternative for decumanus.)
1837. Mus liihcrnicus Thompson, Proc. Zool. Soc, London, p. 52 (Rath-

friland, Co. Down, Ireland).
1857. Mus decumanus Blasius, Saugetbiere Deutschlands, p. 313.
1900. Mus norvegicus Rehn, Proc. Biol. Soc, Washington, xiii, p. 1G7,

October 31, 1900.
1908. Einmys norwcgicus Satunin, ]\Iitth. Kauk. Mus., Tiflls, iv. Lief. 1-2,

p. 111.
1910. Mus {Epimys) norvegicus and M. {E.) norvegicus liihcrnicus Trouessart,

Faune Mamm. d'Europe, p. 142.

Type locality. — Norway.

Geographical distribution. — Originally confined to the north
temperate portions of the Old World ; now essentially cosmo-
politan through artificial dispersal.

Diagnosis. — Skull with brain-case rather narrow, its greatest
breadth across lateral ridges about equal to length of parietal
measured along ridge ; anterior upper molar with evident
cingulum at anterior border of crown ; first lamina of m^ with-
out distinct outer tubercle ; terminal heel of m'^ with no outer
tubercle ; first and second lower molars usually without evident
supplemental cusps at outer extremity of furrows ; form robust ;
tail decidedly shorter than head and body ; ear when laid forward
barely or not reaching eye ; hind foot of adult usually over
40 mm. in length, condylobasal length of adult skulls usuall}-
45 to 54 mm.

External characters. — Size larger and form more robust than
in E. rattits, the tail never as long as head and body. Ear short,
extending bai-ely or not to eye when laid forward, its substance
thick and opaque, its surface densely covered with fine short
hairs, its form not peculiar. Feet more robust than in E. rattus,
but proportions of toes and n\nnber and position of tubercles the
same. In size the tul^ercles are, however, relatively smaller than

EPIMYS

859

those of E. rattns, especially on the hind foot. Except for its
greater diameter and relative shortness the tail is essentially
like that of E. raftiiH, though the rings tend to be less well
defined owing to the greater distinctness of the individual scales
of which they are composed. Fur less harsh in quality than
that of E. rattus, owing to the less abundance of grooved bristles
and the excessive slenderness of such of these hairs as occur.

Mammte

i^J-

al-l; / 3-3=: 12.

Colour. — Upper parts a coarse, variable mixture of dull
ochraceous-buft" and slaty grey, the back darker than the sides and
heavily " lined " (though rarely clouded) with black. Occasionally
thei'e is a decided tinge of russet on posterior half of back.
Underparts greyish white, usually with an inconspicuous wash
of cream-buff. Ear dull hair-brown. Feet concolor with under-
parts. Tail obscurely bicolor, the hairs blackish above, whitish
below, the skin everywhere distinctly appearing at surface.

Sl'nll. — Brain-case narrow and almost sub-cyHndrical in aspect,
the lateral ridges evident but not greatly developed ; width of

Via. 174.

Kplmy.'i norveijicun. Nat. size.

hrain-case across ridges about equal to outer length of parietal ;
interparietal with convexity of anterior and posterior borders
sub-equal, seldom if ever showing the conspicuous contrast
normally present in E. raitus ; width of basioccipital along
anterior border decidedlv more than half median length.

860

RODKXTIA

Rostrum deepened anteriorly so that dorsal profile is nearly
horizontal throughout. Upper border of plate forming outer
wall of infraorbital foramen nearly horizontal and very abruptly
rounded off anteriorly, the j^late wider in proportion to its height
than in Ejjiiiii/s raitnx. In other respects the skull closel}-
resembles that of E. rattm apart from its larger general size.
The auditory bullte tend, however, to be less inflated and the
incisive foramina to be slightly shorter and wider, their posterioi-
border usually not extending quite to level of anterior molar
root.

Teeth. — Incisors as in E. rattus. Molars with number of
roots and general size and proportions of crowns as in E. rattus,
but upper teeth showing a greater tendency toward reduction
of the outer cusps. First upper molar with distinct cingulum-

like ridge at anterior base of crown,
this ridge in some specimens showing
a tendency to develop one or more
minute tubercles ; first lamina notice-
ably distorted by the nearly complete
suppression of t3, which is so reduced
as to be practically nothing more than
an outward prolongation of the outer
border of t2, usually without trace
of antero-external re-entrant angle ;
second and third laminae essentiall}'
as in Epimys rattus. Second upper
^^'^' ^'^' molar like that of E. rattus except

Epunysm>rveoicus. Cheek-teeth. ^^^^ ^^ -^ g^.gatly reduced in size and

completely merged in outer base of
/8, the third lamina thus forming a simple biconvex terminal
heel. Third upper molar essentially as in Epimys rattus, tl
equally well developed, but other elements of tooth less distinct.
Lower molars differing from those of Epimys rattus in the less
evident tendency to develop supplemental tubercles at outer
extremities of spaces between the laminse, such rudiments of the
small tubercles as may be present usually appearing as slight
thickenings at outer extremities of the laminae.

Measurements. — Adult male from Cortachy, Forfar, Scotland :
head and body, 226 ; tail, 179 ; hind foot, 415 ; ear, 20. Adult
male from Cambridge, England : head and body, 242 ; tail, 185 ;
hind foot, 42. Adult male from Tiverton, Devonshire,' England :
head and body, 238; tail, 212; hind foot, 42; ear, 21. Adult
male from Roydon, Norfolk, England : head and body, 250 ;
tail, 218 ; hind foot, 45. Adult male from Hampton, near
London, England : head and body, 255 ; tail, 187 ; hind foot, 41 ;
ear, 20 • 5. • Adult male from Kilmanock, Waterford, Ireland :
head and body, 242 ; tail, 209 ; hind foot, 44 ; ear, 20. Adult
male and female from Upsala, Sweden : head and body, 250 and
252 ; tail, 183 and 183 ; hind foot, 41 and 41. Adult male and

861

female from Palermo, .Sicily : head and body, 242 and 222 ; tail,
200 and 186 ; hind foot, 45 and 43. For ci'anial measurements
see Table, p. 862.

Specimens examined. — One hundred and thirty-five, from the following
localities : —

Scotland: Stockhriggs, Lanarkshire, 2; Cortachy, Forfar, '1 (Wilson).

England : Doncastor, Yorkshire, 1 ; East Norfolk Marshes, 2 ; West
Norfolk, 1 ; Roydon, Norfolk, 1 ; Lowestoft, Suffolk, 1 ; Barrow, Suffolk, 1
(U.S.N.M.) ; Swithland, Leicestershire, 1; Cambridge, 2; Epping Forest,
Essex, 1 ; Hampton, Middlesex, 3; Tiverton, Devonshire, 1; Northlew,
Devonshire, 1; Lundy Island, Devonshire, 2; Isle of Man, 1.

Ireland: Castle Hamilton, Cavan, 1; Ballaghmoon, 1; Kilmanock,
Wexford, 8 ; Inch, Wexford, 1 ; Stokestown, Wexford, 1 ; Glare Island, Mayo,
2 ; Inishmore, Arran Islands, 2 ; Co. Wicklow, 1 ; Castle Hamilton, 1.

Norway : Holme, Mandal, 6.

Sweden : Upsala 8 (U.S.N.M.) ; Stockholm, 1 (U.S.N.M.).

France : Trinity, Jersey, 1 ; St. Martins, Guernsey, 1 ; Etupes, Doubs,
1 ; Barcelonnette, Basses-Alpes 1 (U.S.N.M.) ; St. Gilles, Gard. 2 ; Forest
of Bouconne, Gers, 1 ; Montrejeau, Haute-Garonne, 2 (U.S.N.M.) ; Biarritz,
Basses-Pyrenees, 4 ; Cadillac, Gironde, 2 (U.S.N.M.).

Spain: Pajares, Leon, 3; Bejar, Salamanca, 6; Silla, Vallencia, 5;
Alcoy, Alicante, 4; Elche, Alicante, 1.

Germany: Brunswick, 1 (U.S.N.M.); Moritzburg, Saxony, 1
(U.S.N.M.); Uebigau, Saxony, 2 (U.S.N.M.); Jena, Thiiringen, 2;
Nuremberg, Bavaria, 1 (U.S.N.M.) ; near Konigsberg, 2 (U.S.N.M.).

Switzerland : Neuchatel, 1 (U.S.N.M.) ; Roggweil, Thurgau, 4 ;
Bodensce, 1.

Italy: Palermo, Sicily, 26 (U.S.N.M.).

Greece : Corfu, 1 ; Argostoli, Cephalouia, 3.

2 i. Stockbriggs, Lanarkshire,

Scotland.
(5, ?. Clare Island, Mayo, Ireland.

{R. H. Bunting.)
6. 9. Inishmore, Aran Islands.

2 9. East Norfolk, England.
{Dr. P. H. Emerson.)
6. Roydon, Norfolk.

(5. Swithland, Loicestorshirc.

9. Epping Forest, Essex.
6. Tiverton. Devonshire.
6 juv, 9. Lundy Island, Devonshire.
6. Kilmanock, Wexford.

{G. Barrett-Hamilton.)
i. Kilmanock, Wexford.

9. Inch, Wexford.

6. Sulby Glen, Isle of Man.

6. Holme, ]Mandal, 200 ft.

Norway.
9. Trinity, Jersey, Channel

Islands. (R. H. Bunting.
6. St. Martins, Guernsey,

Channel Islands.

{R. H. Bunting.)
9. Etupes, Doubs, Prance.

^ (C. Mottaz.)

E. R. Alston (c .t f). 79. 9. 25. 37. 87

G. Barrett-Hamilton
G. Barrett-Hamilton

iy)-

Dr. E. Hamilton (r).

Rev. T. C. Hose

(c & p).
Hon. F. A. Butler

(c & p).
H. Doubleday (c & p).
T. F. Tracev (c & p).
T. A. Coward (c & p).
W. Eagle Clarke (p).

G. Barrett-PIamiltoii

G. Brooke (c & p).
C.H.B. Grant (c&p).
R. J. Cuninghame

(c & p).
0. Thomas (p).

0. Thomas (p).

0. Thomas (p).

11. 1. 2. 142-

143.
11. 1. 2. 144-

145.
9G. 3. 28. 1-2.

93. 11. 28. 1.

11. 1. 3. 41G.

46. 6. 15. 27.
11. 1. 3. 417.

7. 12. 19. 7-8.
91. 9. 13. 1.

11. 1. 3. 418.

9. 4. 13. 1.
11. 1. 3. 419.

8. 8. 9. 13.

8. 9. 2. 6.
8. 9. 2. 22.

8. 8. 10. 99.

862

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6. St. Gilles, Gard. G. S. Miller (c). 8. S. 4. 22■^.

•?. Bouconne, Gers. O. Thomas (r). (5. 4. 1. W.

{A. Robert.) r 1 oi '^ 1

2 6,2 9. Biarritz, Basses-Pyr6nees, J. F. Davison (c & !•). '^- ,: ,\ °Z^-
100 ft. (G. 6. 4. 1--2.

6. PajAres, Leon, Spain. O. Thomas (]>). 8. 2. 9. 86.

{N. Gonzalez.)
6, 9. Bcjar, Salamanca. O. Thomas (r). 8. 2. 9. 87-88.

{N. Gonzalez.)
2 6, 9. Silla, Valencia. O. Thomas (r). 8. 2. 9. 91-93

(A''. Gonzalez.)
6, 9. Alco}-, Alicante. O. Thomas (p). 8. 2. 9. 89-90.

{N. Gonzalez.)
6, 9. Jena, Thiiringen, Ger- Prof. K. von Bardc- 94. 4. 4. 1-2.
many. leben (p).

2 6,29. Roggwil, Thurgau, Swit- O. Thomas (r). 2. 8. 4. 39-42

zerland.

{E. H. Zollikofer.) "
6. Bodensee, Switzerland. O. Thomas (r.) 2. 8. 4. 38.

{E. H. Zollikofer.)
9. Corfu, Greece. (C. Moitaz.) J. I. S. Whitakcr (r). 8. 10. 1. 30.
2 9. Argostoli, Cephalonia. J. I. S. Whitakcr (r). 8.10.1.38-39.

(C. Mottaz.)

Genus MUS Linn;eus.

1758. .V»i- Linneeus, Syst. Nat., i, 10th cd., p. 59 (musculus, hyta,utonymy).
1814. Musculus Rafinesque, Preces des Decouv. et travaux Somiologiques,

p. 13 (Substitute for Mus).
1837. Leggada Gray, Charlosworth's Mag. Nat. Hist., i, p. 586, November,

1837 (L. boocluga Gray and Mus platytlirix Bennett).
1844. Drymomijs Tschudi, Fauna Peruana, p. 178 (D. parvulus Tschudi ^

Mus musculus Linna3us. See Palmer, Index Gen. Mamm.,

p. 246, 1904).
1857. Mus Blasius, Saugethiere Deutschlands, p. 309 (part).
1876. Nannomys Peters, IMonatsber, k. preuss. Akad. Wisscnsch. Berlin,

p. 480, August, 1876 {N. sctulosus Peters).
1881. Acromys Trouessart, Bull. Soc. d'Etudes Sci. d' Angers, x, p. 133

(Synonym of Drymomys wrongly attributed to Wagner. Sec

Palmer, Index Gen. IMamm., j). 246).
1896. Pseudoconomys Rhoads, Proc. Acad. Nat. Sci., Philadelphia, p. 531,

December 8, 1896. Mus (Pseudoconomys) proconodon Rhoads.
1900. Dryomys Philippi, An. Mus. Nac. de Chile, xiv, p. 20 (modification

of Drynwmys Tschudi).
1910. Mus Miller, Proc. Biol. Soc. Washington, xxiii, p. 59, Aj^ril 19, 1910.

Type species. — Mus musculus Linnaeus (by tautonym}').

Geographical distrihution. — Originally confined to the tropical
and temperate portions of the Old World ; now essentially
cosmopolitan through artificial dispersal of the type species.

Characters. — In general like Epimys, but mechanical scheme
of molars modified by the elongation of crown of anterior tooth
until it forms the main portion of tooth-row ; m^ with three
roots, its crown decidedly longer than those of two succeeding
teeth combined, its first lamina much distorted by the displace-
ment backward of t\ into line with io and /6 ; uv' small and

864 RODENTIA

tending tn disaiDpear, in some species without trace- of first
lamina ; upper incisor compressed, set at such an angle that a

I pper iUL-isois of Epim>ii< (a) ami Mvn (b).
ilufh enlarged.

sub-apical notch is normally cut in its outer side by action f)f
lower tooth.

Beniarls. — The genus Mm as now restricted contains the
Mks viiiscnlm group and the Asiatic and African species usually
referred to Lrggada. Thougli the species differ considerably
among themselves in details of structure the group is very
homogeneous as regards the peculiar specialization of the
mechanics of the molars and the distortioji of the first lamina
of »(^ About twenty -five forms are now known, seven of which
occur in Europe.

KEY TO THE EUROPEAN FORMS OF :\IUS.

HiDcl foot broad and robust, its width at base of
enter toes 4 to 5 mra.
Condylobasal length of skull in adults 21-6 to

22 • 2 mm. ; bind foot 17-4 to 18-8 mm.

(St. Kilda Island, Scotland) M. muralii;, yi 874.

Condylobasal length of skull in adults 23 to

23 "4 mm; hind foot 19 to 20 mm. (Faeroes)... M. fxrocnsis, p. 875.
Hind foot narrow and slender, its width at base of

outer toes about 3 mm.
Tail about as long as head and body, often longer,
rarely much shorter; colour of underparts
usually greyish, rarely contrasted with that
of sides; condylobasal length of skull fre-
quently more than 21 mm. ; hind foot usually

17 to 19 mm. (House mice) M. muscalus, p. 865.

General colour dark and brownish, usually
without evident yellowish tinge, the belly
dusky greyish (central and northern

Europe) jV- '"• musculus, p. 869.

General colour light, usually with a decided
yellowish tinge, the belly bufiy greyish
(Mediterranean region) M. m. azoi-!<:i>.s, p. 871.

MDs 865

Tail noticeably shorter than head and body
(exceptions very rare) ; colour of nnderparts
usually whitish in abrupt contrast with that
of sides ; condylobasal length of skull rarely
attaining 21 mm. ; hind foot usually 15 to

17 mm. (Field mice, rarely eiitering houses)... M. spicilegus, p. 877.
General colour above a clear rather dark greyish
brown with no decided yellowish or russet
tinge (eastern and east-central Europe,

Sweden) M. s. njncUegux, ^. 878.

General colour above not clear greyish brown
unless very pale, the back usually with
decided yellowish or russet tints.
Colour of upper parts ranging from nearly

clear buff to pale huffy grey (Spain) ^f. s. hispaiiicuit, p. 879.

Colour of upper parts brownish grey, usually

with a decided russet tinge (Portugal) ... M. s. lusitanicus, p. 882.

MUS MUSCULUS Linnanis.
(Synonymy under subspecies.)

Geographical distribution. — At present almost cosmopolitan ;
everywhere characteristically associated with human dwellings,
though not infrequently found at a distance from houses.

Diagnosis. — Size medium (head and body about 75 to 100 mm.,
hind foot, 17 to 19*4 mm., condylobasal length of skull, 19-8 to
22 "4: mm.); tail about as long as head and body, often longer,
rarely much shorter ; hind foot narrow and slender, its greatest
width across base of toes about 3 to 3 • 5 mm. ; colour of under-
parts usually greyish buff, rarely forming any decided contrast
with the dusky brown or huffy brown of sides and back.

External characters. — General form rather slender, with no
special peculiarities. Ear moderately large, extending a little
beyond eye when laid forward, its outline broadly oval ; a well
developed but low ridge on inner surface behind meatus ; entire
surface of ear finely pubescent except internally toward base.
Fore foot with thumb reduced to a small tubercle with very
rudimentaiy appressed nail ; fifth finger extending slightly
beyond base of fourth ; second nearly equal to third and fourth
which are sub-equal and longest ; tubercles small, occupying
distinctly less than half the surface of palm ; the three anterior
are nearly circular in outline and about as large as thumb, that
lying at base of fifth finger with small but distinct supplemental
outer tubercle, the two posterior are decidedly larger and some-
what elongated ; surface of palm between tubercles irregularly
tuberculo-reticulate ; scales on under surface of fingers tending
to be divided at middle. Hind foot relatively shorter and more
robust than in Apodemus sylvaticus, its greatest width at base of
toes about 3 to 3 • 5 mm. ; the three median toes sub-equal, the
inner usually not so long as the others ; first digit extending
about to base of second, fifth extending a little beyond base of
fourth ; sole naked except at sides of heel, its surface smooth

3 K

866

RODENTIA

posteriorly, finely tuberculate anteriorly ; plantar tubercles
small, widely spaced, without noticeable contrasts in size or form,
the two anterior slightly the largest, the postero-external slightly
the smallest, their outlines ovate or nearly circular ; a small but
distinct supplemental tubercle at outer margin of tubercles lying
at base of first and fifth toes. Claws simple, curved, those on
hind feet slightly the larger. Tail about as long as head and
body, sometimes a little longer, rarely much shorter unless
injured, its diameter at base about 3 mm., its tip not so
slender as in Apodemus sylvaticus ; annulations well defined but
narrow, about twenty-two to the centimeter at middle, the boun-
daries of the individual scales obscurely marked ; hairs rather
numerous but not concealing annulations, and not forming pencil,
their length equal to about width of two and a half rings. Fur
soft throughout, the grooved hairs so slender as not to suggest
bristles in texture. Mammae : p3 — 3; ^ 2 — 2 = 10.

Colour. — Upper parts a light, dull wood-brown, irregularly
darkened with slaty and blackish along back, faintly more bufFy
on sides, the general effect not far from the broccoli-brown of
Ridgway, but usually with a peculiar dusky cast ; underparts
essentially like sides or a little more yellowish, the line of demar-
cation absent or very obscure ; bases of hairs everywhere slate-
grey, this colour usually appearing at surface on chin and throat,
where it produces a smoke-grey or drab-grey effect ; ear dull
brownish, the outer side of anterior border usually darker ; feet
drab or dusky, not contrasting with back ; tail dull brownish
throughout, occasionally lighter below than above.

Skull. — As compared with that of small European Muridx of
other genera, Micromys excepted, the skull of Mas musculus is
immediately recognizable by its small size. Its general form is
characterized by a peculiar flatness not
found in any of the others. Dorsal
j^rofile slightly convex throughout, the
nasals sloping forward at an angle which
causes an evident break in the general
curve ; occiput low, somewhat obliquely
truncate, so that the entire supraocci-
pital between lambda and foramen
magnum is visible when skull is viewed
from above ; ventral profile faintly con-
cave ; contrast between depth of brain-
case and rostral depth noticeably less
than in Ajjodemus sylvaticus. Brain-case
short-oval in outline when viewed from
above, with a slight though evident ten-
dency toward squaring behind and in
front. Interparietal rather large, its area nearly equal to that
of one parietal, its general form ligulate, the anterior border pro-
jecting forward at middle, the lateral extremities almost squarely

Fig. 177.
Mus )inisctdu)i. Nat. size.

Mus 867

truncate ; lambdoid suture passing along anterior border of inter-
parietal so that the bone lies entirely within occipital ; ■••' sides
of brain-case abruptly rounded oft' or faintly angular, never
beaded. Occiput broad and low when viewed from behind, but
central portion of brain-case not rising conspicuously above it ;
paroccipital process short and inconspicuous, barely extending to
level of lower lip of foramen magnum. Floor of brain-case
with no striking peculiarities ; width of basioccipital along
anterior border contained about 2| times in median length ;
auditory bulla moderately large, less inflated than in Apodemus
sylvaticus but occupying about the same area on base of skull,
its beak rather short and broad, but evident, the meatus with
anterior portion of upper border slightly projecting. Inter-
orbital region distinctly wider than rostrum, slightly convex
laterally, the margins falling away squarely but with no trace
of bead. Zygoma relatively heavy, compressed throughout but
not evidently expanded, moderately bent downward and moder-
ately spreading anteriorly, so that greatest zygomatic breadth is
usually just in front of glenoid level ; plate forming outer wall
of anteorbital foramen well developed, its anterior border vertical,
abruptly rounded above ; a short but very distinct peg-like
process for muscle attachment at antero-inferior angle of plate,
this process nearly as well developed in half-grown young as in
adults. Rostrum slightly deeper than broad, with no special
peculiarities of form ; nasals truncate or bluntly pointed
posteriorly, extending to or slightly beyond lachrymal level, and
usually a little exceeded by nasal branches of pi-emaxillaries.
Incisive foramina very long (nearly equal to diastema), extending
from about 1-5 mm. behind incisors to level of middle of crown
of m^ ; their greatest width is at region of maxillo-premaxillary
suture ; posteriorly they narrow to an almost acute termination,
a peculiarity of form highly characteristic of this group as com-
pared with other European Muridse. Palate extending slightly
behind level of «i^, its length behind incisive foramina less than
width including tooth-rows. Mesopterygoid space moderately
long, wider posteriorly than anteriorly, squarely truncate in
front, the pterygoids and hamulars straight, the latter barely
in contact with bulbw. Mandible rather short and deep as
compared with that of Apodenius sylvaticus, the coronoid process
well developed, strongly curved backward, its tip rising slightly
above level of condyle.

Teeth. — Upper incisors much compressed, the outer and
inner borders nearly parallel through anterior half of their
extent, the transverse diameter contained about 2}, times in
antero-posterior diameter. Enamel unusually thick, scarcely
extending backward on inner side of tooth, but folding notice-

* In Apodenms agrarius, which has a similarly ligulate interparietal,
the suture passes along posterior border so that the small bone lies within
the parietals (fig. 168, p. 837).

3 K 2

868

ably backward on outer side. Main axis of shafts tliverging
posteriorly, the angle at which they are set such that a con-
spicuous sub- apical notch is formed on outer side of cutting edge of
tooth immediately behind margin of enamel. First upper molar
three-rooted, the anterior root largest, projecting conspicuously
forward beyond base of crown, separated from more posterior
roots by an appreciable space ; crown decidedly longer than
that of m^ and m'^ together, its area nearly 1| times as
great ; anterior border of crown smooth or with a slight angle
representing cingulum ; first lamina with crescentic form much
distorted by the displacement backward of tl to a position
essentially in line with t5 and tQ ; tS well developed, nearly
equal to ti or tQ, but not so large as tl, its separation from t2
marked by an evident re-entrant extending almost to base of
crown ; tl separated from i2 by a broad, flattened or slightly
concave area which imparts to the antero-internal border of the

crown a characteristic obliquely-trun-
cate appearance ; second lamina more
regularly crescentic than first, but
with evident traces of the same dis-
tortion ; third lamina represented by
well developed t8 and t9 essentially
similar to t5 and /6, but with no
trace of t7 unless a short enamel
ridge connecting ti and /8 be inter-
preted as representing a trace of this
tubercle. Second upper molar with
first lamina represented by a well
developed, sub-terete tl about as large
as the outer tubercles of m^ ; second
lamina with ti slightly larger than
tQ and connected posteriorly with
/8 as in the preceding tooth ; third lamina about as in m^,
but t9 reduced, decidedly smaller than t6, though marked
off from t8 by an evident re-entrant angle. Third upper
molar relatively smaller than in any of the other European
Miiridse, its area scarcely greater than that of terminal heel of
m'^. In form the crown is sub-circular with a small, often
indistinct tubercular antero-internal supplement representing tl.
Anterior lower molar with crown about as long as those of the
two succeeding teeth together, its area about equal to them ;
first lamina reduced to a single large tubercle forming the entire
anterior border of the tooth and situated slightly internal to the
median line of the crown ; internally it is separated from second
lamina by a deep re-entrant depression, but externally it is
connected with outer tubercle of second lamina by a high, some-
times distinctly tubercular ridge, which occasionally * forms a

Fig. 178.

Mus tnusculus. Clieek-teeth. X 10

(Antero-external re-entrant an^le

of mj usually nuicli less evident.)

* As in specimen represented in fig. 178.

Mus 869

true outer tubercle on jfirst lamina ; second and third lamin;i' and
terminal heel normal and with no trace of supplemental tubercles
on outer side of crown. Second lower molar essentially like
second and third laminae of first, but with tubercles of posterior
lamina distinctly smaller than those of anterior lamina, and
terminal heel greatly reduced. Third lower molar with crown
area equal to about one-third that of second, its anterior lamina
with two tubercles smaller than those of posterior lamina of
second tooth, its terminal heel relatively better developed than
that of m'^.

Bemarks. — Although occasionally straying to considerable
distances from houses, Mus vmscuJus is characteristically an
urban species, and its spread from central Asia to Evu-ope and
thence throughout the world has been chiefly due to human
agency. Two moderately well-marked local races occur on the
continent of Europe, while two more strongly differentiated
forms are confined respectively to St. Kilda Island and the
Faeroes.

Mus MUSCULUS MUSCULUS LinUiVUS.

1758. [Mils'] musculics Lianseus, Syst. Nat., i, 10th ed., p. 62 (Sweden).

1801. lf[?ts] m[usctdus] albus Bechstein, Gemeinu. Naturgesch. Deutscb-
lands, I, 2d ed., p. 955 (Thiiringen, Germany).

1801. M[us'] m[usculiis] flavus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 955 (Thiiringen, Germany).

1801. M[us] 7n[iisciilns] maculatus Bechstein, Gemeinu. Naturgesch.
Deutschlands, i, 2d ed., p. 955 (Thiiringen, Germany).

1801. M[tis\ m[usculus] niger Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 955 (Thiiringen, Germany).

1827. [Mus musculus'] 0 striatus Billberg, Synopsis Faunae Scandinaviaj,

]}. 6 (Siitlne, Sweden).
1827. [Mus musculus'] y albicans Billberg, Synopsis Faunre Scandinaviee,

p. 6 (Sk&ne, Sweden),
1827. [Mus musculus] S nivcus Billberg, Synopsis Faunae Scandinavia,

p. 6 (Molle, Norway).
1837. ?il/«s hrevirostris Waterhouse, Proc. Zool. Soc, London, p. 19 (Mal-

donado, Uruguay).
1844. ^Dryviotnys 2>arvulus Tschudi, Fauna Peruana, p. 178 (Central Peru).
1857. Mus musculus Blasius, Siiugetbiere Deutschlands, p. 320.
1867. [Mas muscuhis] helvolus Fitzinger, Sitzungsber. kais. Akad. Wis-

sensch. Wien, Matb.-Naturwiss. Classe, LVi, pt. i, p. 70

(Hungary).
1867. [Mtis viusculus] varius Fitzinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 70 (Europe).
1807. [Mus musculus] cinerco-maculatus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lAi, pt. i, p. 70

(Europe).
1869. Mus poschiavinus Fatio, Faunc Vert. Suisse, i, p. 207 (Poschiavo,

Grisons, Switzerland).

870 RODENTIA

1872. Mns musculus var. flavescens Fischer, Zool. Garten, xiii, p. 223,

July, 1872 (Berlin, Germany).
1910. Mhs musculus and M. musculus poschiavinus Troiiessart, Fanne

Mamni. d'Europe, p. 145.

Tfipe locality. — Upsala, Sweden.

GeograpJiical distribution. — ^Europe north of the Mediterranean
region.

Diagnosis. — General colour dark and brownish, usually with-
out evident yellowish tinge, the belly dusky greyish.

Measurements. — Adult male and female from Iceland : head
and body, 87 and 89 ; tail, 95 and 92 ; hind foot, 18-6 and 19 ;
ear, 17 and 16. Average and extremes of seven adults from
near London, England: head and body, 78*4 (75-84); tail,
78-6 (76-86); hind foot, 17-3 (16-6-18). Adult male and
female from Richmond, Surrey : head and body, 81 and 80 ; tail,
89 and 77; hind foot, 16-6 and 17; ear, 14-4 and 14. Adult
female from Holme, Mandal, Norway : head and body, 85 ; tail,
89; hind foot,' 17*4; ear, 14. Average and extremes of eight
adults from the Harz Mountains, Germany : head and body, 96
(87-103); tail, 90-2 (84-102) ; hind foot, 18-4 (17'6-19-4).
For cranial measurements see Table, p. 872.

Specimens examined. — One hundred and one, from the following
localities : —

Iceland : Near Eeykjavik, 7 (U.S.N.M.).

Scotland : North Uist, Hebrides, 1 (Edinburgh) ; Dunvegan, Skye, 1 ;
Grantown-on-Spey, Elgin, 2 (Wilson) ; Jardine Hall, Dumfriesshire, 4
(U.S.N.M.).

England : Oundle, Northamptonshire, 3; Broxbourne, Hertfordshire, 1;
Berkhamsted, Hertfordshire, 1 ; Cheltenham, Gloucestershire, 1 (Wilson) ;
London, 3 (B.M. and U.S.N.M.) ; Richmond, Surrey, 5 (U.S.N.M.) ; Wands-
worth Common, Surrey, 4 ; Martock, Somerset, 1 ; Newport, Isle of Wight,
1 ; Ramsay, Isle of Man, 2 ; Ballasalla, Isle of Man, 1 ; St. Ouen's Bay,
Jersey, 3; Alderney, 1.

Ireland: Clare Island, Mayo, 3; Clifden, Galway, 2.

Norway : Graven, Hardanger, 1 (U.S.N.M.) ; Holme, Mandal, 6.

Belgium : Waremme, 2 (U.S.N.M.).

Germany: Harz Mountains, 19 (U.S.N.M.); Dresden, 7 (U.S.N.M.);
Nuremberg, Bavaria, 1 (U.S.N.IM.).

Switzerland : Andermatt, Uri, 1 (U.S.N.M.) ; St. Gothard, Uri, 1
(U.S.N.M.); Untervatz, St. Gallen, 1 ; St. Gallen, 4 ; Poschiavo, Grisons, 4
(U.S.N.M. and Geneva) ; St. Cergues, Vaud, 1.

2 6, 9. Clare Island, Mayo, Ire- G. Barrett-Hamilton 11. 1. 2. 146-148.

land. [E. H. Bunting.) (p).

6, 9. Clifden, Galway. G. Barrett-Hamilton 11. 1. 2. 149-150.

[r. H. Bunting.) (p).

2 6, V. Oundle, Northampton- Lord Lilford (p). 11. 1. 1. 166-168.

shire, England.

S. Wandsworth, Surrey. J. Miller (c & p). 4. 2. 12. 1.

9. Martock, Somerset. C. B. Horsbrugh 11. 1. 3. 420.

(c & p).

S. Newport, Isle of Wight. R. J. Cuninghame 11. 1. 3. 421.

(c & p).

2 i, 9. St. Ouen's Bay, Jersey, 0. Thomas (p). 8. 9. 2. 7-9.

Channel Islands.

(iJ. H. Bunting.)

871

1 al. Alderney, Channel Is- W. Eagle Clarke (p). 98.9.29.7.
lands.
2 6,3 9. Holme, Mandal, Nor- R. J. Cuninghame 8. 8. 9. 14-18.
way. (c & p).

6. Untervatz, Grisons, O. Thomas (p). 4.4.5.45.

Switzerland.

(E. H. Zollikofcr.)
3 6, 9. St. Gallen. 0. Thomas (p). 4. 4. 5. 41-44.

{E. H. Zollikofcr.)

MUS MUSCULUS AZORICUS vScliinz.

1845. M[us] azoricus Schinz, Synops. Mamm., ii, p. 161 (Azores).
1355. Musculus niollissimus Dehne, Allgem. deutsche Naturhist. Zeitung,
Dresden, neue Folge, i, p. 443. (Monte PoUino, Basilicata,

Italy).

Type localittj. — Azore Islands.

Gcograpliical (U>itrihuUon. — Mediterranean region; Azores.

Diagnosis. — General colour less dusky and more yellowish
than in M. musculus 7nusculus, the underparts buffy greyish.

Measurements. — Adult male and female from Elche, Alicante,
Spain : head and body, 84 and 84 ; tail, 92 and 86 ; hind foot,
18 and 18' 4; ear, 14*8 and 14. Two adult females from near
Nimes, Gard, France : head and body, 88 and 91 ; tail, 86 and
88; hind foot, 17 '5 and 17*5; ear, 13*5 and 14*4. Average
and extremes of ten adults from Sorrento, Italy : head and body,
85-5 (78-93); tail, 88-6 (82-94); hind foot, 17-7 (17-18-6).
Average and extreme of ten adults from Palermo, Sicily : head
and body, 85*7 (82-92); tail, 82-2 (78-85); hind foot, 17-4
(17-18" 4). For cranial measurements see Table, p. 872.

Speciyncns examined. — One hundred and ninety, from the following
localities : —

Portugal : San Miguel, Azores, 6 ; Estoril, 1 ; Cintra, 2.

Spain : Pajares, Leon, 2 ; Silos, Burgos, 2 ; Bejar, Salamanca, 3 ;
Dehesa de Valencia, Valencia, 1 ; Elche, Alicante, 2 ; Venta del Baul,
Granada, 5 ; San Cristobal, Minorca, Balearic Islands, 1.

Prance: Biarritz, Basses-Pyr6n6es, 2; Ax-les-Thermes, Ariege, 1;
I'Hospitalet, Ariege, 1 ; Port6, Pyr6n6es-Orientales, 1 ; near Nimes,
Gard, 8 ; St. Gilles, Gard, 1 ; St. Paul, Basses-Alpes, 6 ; La Foce di
Vizzavona, Corsica, 1.

Italy : Acceglio, Cuneo, 1 ; Calappiano, Empoli, 8 ; Florence, 11
(B.M. and U.S.N.M.) ; Rome, 2; Sorrento, 24 (U.S.N.M.) ; Marsala,
Sicily, 1 ; Palermo, Sicily, 59 (B.M. and U.S.N.M.) ; Taormina, Sicily, 10
(U.S.N.M.) ; Lanusei, Sardinia, 1 (U.S.N.M.).

Greece : Corfu, 22 ; Patras, 6 ; Athens, 4 (U.S.N.M.) ; Argostoli,
Cephalonia, 1.

5 6, 9. San Miguel, Azores. Hon. W. Rothschild 3. G. 5. 17-22.
( W. B. Ogilvie-Clranf.) (p).

6. Estoril, Portugal. 0. Thomas (c & p). 98. 2. 2. 3G.

6, 9. Cintra. 0. Thomas (c & p). 98. 2. 2. 28-29.

6, 9. Pajares, Leon. 0. Thomas (p). 8. 2. 9. 133-1-34.
(N. Gonzalez.)

9. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 109.

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RODENTIA

6, 2 9.

Bejar, Salamanca.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 136-138,

6.

Dehesa de Valencia.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 135.

6, 9.

Elche, Alicante.

G. S. Miller (c).

8. 8. 4. 107-108,

6, 2 9.

Venta del Baul, Granada.

G. S. Miller (c).

8. 8. 4. 110-112.

9.

Venta del Baul, Granada.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 130.

6.

San Cristobal, Minorca,

0. Thomas and R. I.

0. 7. 1. 58.

Balearic Islands.

Pocock (c & p).

9.

St. Gilles, Gard, France.

G. S. Miller (c).

8. 8. 4. 222.

4 9.

Ntmes, Gard. (C. Mottaz.

) 0. Thomas (p).

8. 8. 10. 93-96.

6,9.

St. Paul, Basses-Alpes,
1470 m. (C. Mottaz.)

0. Thomas (p).

8. 8. 10. 97-98.

6,9.

Rome. {Coli.)

G. Barrett-Hamilton

11. 1. 2. 38-39.

6.

Marsala, Sicily.

(.4. Robert.)

0. Thomas (p).

6. 8. 4. 44.

3 6, 9.

Palermo.

J. I. S. Whitaker (p).

98. 10. 6. 12-15.

9.

La Foce de Vizzavona,

Col. J. W. Yerbury

93. 9. 15. 9.

Corsica.

(c & p).

2 6,4:9.

Corfu, Greece.

(C. Mottaz.)

0. Thomas (p).

8. 10. 1. 43-48.

3 6,2 9.

Corfu, Greece.

C. Mottaz (c).

8. 11. 3. 12-16.

2 6,9.

Patras, Greece.

Hon. N. C. Roths-

8. 10. 2. 44-46.

(C. Mottaz.)

child (p).

6.

Argostoli, Cephalonia.

C. Mottaz (c).

8. 11. 3. 17.

MUS MURALIS Barrett-Hamilton.

1899. Mus muralis Barrett-Hamilton, Proc. Zool. Soc, London, p. 81.

Type in British Museum.
1910. Mus muscalus muralis Trouessart, Faune Mamm. d'Europe, p. 146.

Type locality. — Island of St. Kilda, Scotland.
Geographical distribution. — Island of St. Kilda.
Diagnosis. — Similar to Mus musculus, but feet and tail less
slender (width of hind foot at base of outer toes about 4 mm.) ;
skull with mesopterygoid fossa coming to a sharp point anteriorly,
the pterygoids nowhere parallel.

External characters. — Except for the less slender
tail and more robust feet the external characters do
not differ appreciably from those of Mus musculus.

Colour. — The colour is essentially as in Mus
musculus, though the underparts appear to be more
frequently a well-defined buffy white.

Skull and teeth. — Apart from the peculiarities of
the palate the skull agrees with that of Mus mus-
culus. Mesopterygoid space conspicuously narx'ower
anteriorly than posteriorly, its lateral borders nowhere
approximately parallel, its width in front only one-
third to one-fourth that between tips of hamulars.
This cuneate form, though less accentuated in immatui'e speci-
mens than in adults, is constantly different from that charac-

Miis muralin
Nat. size.

Mus 8T5

teristic of the large series of European specimens of Mus mnsculus
examined. Teeth as in Mus musculus.

MensHrements. — External measurements of type (adult
female) : head and body, 90; tail, 85; hind foot, 17"4; ear, 14.
Adult male and female from the type locality : head and body,
84 and 89; tail, 89 and 87 ; hind foot, 18-8 and 18; ear, 14
and 14. For cranial measurements see Table, p. 876.

Specimens examined. — Fifteen, all from St. Kilda (B.M. and Edinburgh).

?. St. Kilda, Scotland. G. Barrett-Hamilton (p). 8. 7. IG. 1.
[H. Evans.) {Type of species.)

i, 9 al. St. Kilda. J. E. Halting (p). 94. 7. 16. 2-3.

1 al. St. Kilda. Kelvingrove Museum, 96. 0. 6. 1.

Glasgow (p).
2 (5, 3 9. St. Kilda. W. Eagle Clarke (c & p). 11. 1. 24. 7-11.

MUS FiEROENSIS Clarke.

1904. Mhs musculus fxroensis Clarke, Proc. Roy. Pliys. Soc. Edinburgh,

XVI, pt. II, p. 163. Type in Royal Scottish Museum.
1910. Mus musculus f.rronisis Trouessart, Faune Mamm. d'Europe, p. 146.

Type locality. — Nolsoe, Faeroe Islands.

Geographical distribution. — Faeroe Islands.

Diagnosis. — Decidedly larger than Mus musculus and M.
muralis (hind foot, 19 to 20 mm., condylobasal length of skull,
23 to 23*4 mm.); hind foot very robust, its width at base of
outer toes about 5 mm. ; tail noticeably thickened, its diameter
near base about 4 mm. as opposed to 3*6 mm. in M. muralis and
about 3 mm. in M. musculus.

External characters. — Except as already pointed out the
external characters agree with those of Mus musculus.

Colour. — The few specimens examined indicate that the colour
does not differ essentially from that of Mus musculus.

Shill and teeth. — The skull resembles that of Mus musculus
except for its larger size, relatively more robust rostrum and
apparently more depressed brain-case, though the material is
insufficient to show whether this last character is constant. In
the three skulls examined from Nolsoe the palate is as usual in
Mus musculus, the interpterygoid space broadly rounded or truncate
anteriorly, and the pterygoids, though imperfect, evidently not
far from parallel. The skull from Myggena-s, however, as
pointed out by Dr. Wiuge in the letter published in the original
description of the species, has the pterygoid region exactly as in
Mus muralis. This skull (No. 1229 Copenhagen Museum) is
abnormal, the rostrum having been bent to the left, apparently
by some injury when the animal was young. The teeth are
slightly larger than those of Mus musculus, but I can detect no
peculiarities of form.

Measurements. — Type (adult female) : hind foot (dry), 20.
Adult male from the type locality : head and body, 85 ; tail, 97 ;

876

RODENTIA

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M. muralis.

Scotland : St. Kilda Island

M. faeroensis.

Faeroe Islands : Nolsoe

Myggeuses

Mus 877

hind foot, 20 ; ear, 1 3. Adult female from the type locality :
hind foot (in alcohol), 19 "8. For cranial measurements see
Table opposite.

Specimens examined. — Six, three from Nolsoe (Edinburgh aud Copen-
hagen), and three from Myggenses (Copenhagen).

Bemarks. — The Faeroe house mouse diflers so conspicuously
from all other known members of the group, that however recent
the animal's origin may have been there seems no doubt that
the form is a distinct sjsecies. The enlarged extremities may be,
as suggested by Dr. Winge, the result of changed habits due to
a climbing life among the clitfs of the islands ; but in view of
the fact that a similar tendency is shown by two of the resident
birds of the Faeroes, it seems more probable that the characters
of all these local forms has been determined by some more
general factor of climate or food.

The only species with which Mus faeroensis is likely to be
confused is Mus muralis of St. Kilda. But the size of the Faeroe
mouse is noticeably greater, and the peculiar thickening of the
feet is carried to a more noticeable extreme. The general colour
of the body is darker than in Mus muralis, essentially as in
ordinary specimens of the house mouse, and the underparts and
feet show no tendency to become whitish.

MUS SPICILEGUS Petenyi.
(Synonymy under subspecies.)

GeograpMcal distrihulion. — Eastern and southern Europe,
west to Portugal, Hungary and Sweden.

Diagnosis. — Size less than in Mus itmsculus, the head and
body rarely attaining a length of 90 mm., the hind foot in adults
ranging about from 1.5 '4 to 18 mm., condylobasal length of
skull, 19 to 21*4 mm. ; tail noticeably shorter than head and
body (exceptions very rare) ; colour of underparts usually
whitish in abrupt contrast with that of sides ; upper incisor
with sub-apical notch less developed than in Mus musculus.

Colour. — For detailed descriptions of the colour see accounts
of subspecies. In the yellower races the colour is strikingly
different from that of Mus vmsculus. In the browner forms the
general aspect is much like that of the house mouse, but the
back is a clearer less dusky brown.

Bemarks. — Mus spicilegus is a strictly agrarian animal, found
in fields, scrub, and open, dry forest, and never inhabiting towns.
It is undoubtedly indigenous to the Mediterranean region.
Related forms occur in the corresponding life zone of north
Africa and of Asia Minor and central Asia. Among the
European representatives of the species three races may be
distinguished, one of which ranges somewhat further north than
the others.

878

Mus SPiciLEGUS SPiciLEGUS Petenyi.

1882. Mus spicilegus Pet6nyi, Tcrm6szctrajzi Fiizetek, Budapest, v, p. 114.
1882. Mus accrvator Pet6nyi, Term^szetrajzi Fiizetek, Budapest, v, p. 114

(alternative name).
1882. Mus accrvifex Petenyi, Termeszetrajzi Fiizetek, Budapest, v, p. 114

(alternative name).
1882. M^ls canicularius Petenyi, Termeszetrajzi Fiizetek, Budapest, v,

p. 114 (alternative name).
1882. Mus caniculator Petenyi, Termeszetrajzi Fiizetek, Budapest, v,

p. 114 (alternative name).
1896. Mus musciUus flavcsccns Barrett-Hamilton, Zoologist, 3d. ser., xx,

p. 179, May 1896 (not Mus musculus \&r. flavesccns Fischer, 1872).
1910. Mus spicilcgus Trouessart, Faune Mamm. d'Europe, p. 148.

Type locality. — Hungary.

Geographical distribution. — East central Europe from the
northern portion of the Balkan Peninsula to the shores of the
Baltic ; westward into Sweden.

Diagnosis. — General colour of upper parts a clear greyish
brown without decided yellowish or russet suffusion.

Colour. — Upper parts a light wood-brown, becoming some-
what bufFy on sides, the entire median region from nose to base
of tail rather conspicuously though finely " lined " with black,
this " lining " sometimes producing an evident dark median
dorsal area, and always becoming inconspicuous on sides and
disappearing entirely on narrow area bordering lighter colour
of uuderparts. Basal portion of hairs slate-grey. Underparts
rather sharply defined bufFy white or pale cream-bufF, the slate-
grej bases of the hairs producing a slight greyish cast. Feet
and under surface of tail essentially concolor with belly ; upper
surface of tail brownish without sharp line of demarcation.
Ears sprinkled with brownish hairs externally, with bufFy
internally, neither surface with any decided colour.

Measurements. — Average of seven adults from Upsala,
Sweden: head and body, 86-2 (82-96); tail, 67-8 (60-77);
hind foot, 17 "1 (15 '4-17 -4). Average and extremes of seven
adults from near Konigsberg, Germany: head and body, 81*9
(73-90); tail, 67-8 (60-75); hind foot, 16-8 (16-4-17-4).
Adult female from Hatszeg, Hunyad, Hungary : head and body,
81 ; tail, 62 ; hind foot, 17 ; ear, 12. Adult male from Gageni,
Roumania : head and body, 75 ; tail, 60 ; hind foot, 16 ; ear, 12.
Average and extremes of five adults from Patrily, Roumania :
head and body, 77-5 (72-83) ; tail, 6-21 (59-5-70) ; hind foot,
16-2 (16-16-5); ear, 12 (11-5-135). For cranial measure-
ments see Table, p. 880.

Specimens examined. — Fifty-seven, from the following localities : —
Sweden: Upsala, 10 (U.S.N.M.) ; Stockholm, 2 (U.S.N.M.).
Germany : Baltic coast, west of Konigsberg, 17 (U.N.N.M.).
Austria-Hungary : G&ailokoz-Somorja, Pressburg, 5 ; Hatszeg,
Hunyad, 2 ; Pester Comitat, 2.

Mus 879

RouMANiA : Gageni, Prahova, 1 ; Patrily, 7.
Montenegro : Various localities, 5.
Albania : Various localities, 6.

5. Csallokoz-Somorja, Press- Budapest Museum 94. 3. 1. 48-52.

burg, Hungary. (e).

2 ?. Hatszeg, Hunyad, Trau- C. G. Danford (c). 3. 2. 2. 33-34.

sylvania.
6, 9. Pester Comitat. Budapest Museum 94. 7 26 1-2

i. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 41.

mania. {W. Dochon.)

7 6. Patrily, Roumania. Lord Lilford (p). 4. 4. 6. 42-48
(W. Dodson.)

2 S, 3 9. Montenegro. (L. Filhrer.) O. Thomas (p). 5. 8. 4. 18-22.

3 6, 3 9. North Albania. 0. Thomas (p). 5. 8. 4. 38-43

(L. Filhrer.)

Mus spiciLEGUs iiisPANicus Miller.

1909. Mus spicilegus liispanicus Miller, Ann. and Mag. Nat. Hist., 8th ser

III, p. 421, May, 1909.

1910. Mtis spicilegus hispanicus Trouessart, Fauue Mamm. d'Europe

p. 148.

Type localitij. — Silos, Province of Burgos, Spain.

Geographical distribution. — Central and southern Spain.

Diagnosis. — General colour of upper parts bufty or pale buffy
grey.

Colour. — Back and sides ranging from buff to a pale buffy
grey lighter and less yellow than the cream-buff of Ridgway, the
median dorsal region faintly "lined" with black, the sides
gradually becoming clear buff or buffy grey, this colour con-
tinuing forward over cheeks and above eye to muzzle ; basal
portion of hairs slate-grey. Underparts sharply defined buffy
white, slightly clouded by slate-grey undercolour. Feet and
tail like belly, the tail with a narrow dusky dorsal area extend-
ing to tip. Ear thinly clothed, the colour buffy or greyish in
harmony with surrounding parts.

Measurements. — Type (adult female) : head and body, 79 ;
tail, 50; hind foot, 14-4; ear, 13-6. Average and extremes
of four adults from the type locality : head and body, 76-7
(75-79); tail, 53-7 (50-58); hind foot, 15-4 (14-4-16); ear
from meatus, 12-8 (12-13*6). Average and extremes of ten
adults from Elche, Alicante, Spain: head and body, 81*4 (76-
91); tail, 62-1 (59-72); hind foot, 16-5 (16-17); ear, 13-6
(13-14). Average and extremes of seven adults from San
Cristobal, Minorca, Balearic Islands: head and body, 77 "2
(75-80); tail, 63-2 (59-66); hind foot, 16-1 (15-6-16-8);
ear, 12-5 (12-13). For cranial measurements see Table, p. 880.

Specimens examined. — Eighty-five, from the following localities : —
Spain : Venta del Baul, Granada, 7 ; Elche, Alicante, 24 ; Alcoy,
Alicante, 21 ; Silos, Burgos, 12 ; Castrillo de la Reina, Burgos, 3 ; Pajdres,

880

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M.spicilegus spicilegus.
Sweden: Upsala .

Stockholm
Germany: Konigsberg .

M. spicilegus his-
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Spain : Majorca, Balearic!
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882 RODENTIA

Leon, 1 : San Cristobal, Minorca, Balearic Islands, 7 ; Inca, Majorca,
Balearic Islands, 6.

Feance : Cerbere, Pyr6n6es-Orientales, 1 ; near Nimes, Gard, 2 ;
Valescure, Var, 1. The Gard and Var specimens not typical.

4 6, ?. VentadelBaul, Granada, G. S. Miller (c). 8. 8. 4. 102-106.
Spain.

i 6,2 9. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 89-94.

13 6,89. Alcoy, Alicante. O. Thomas (p). 8. 2. 9. 139-159.
[N. Gonzalez.)

4 6, ?. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 97-101.

(8. 8. 4. 101. Type of subspecies.)

6, 9. Castrillo, Burgos. G. S. Miller (c). " 8. 8. 4. 95-96.

6. Paj4res, Leon. 0. Thomas (p). 8. 2. 9. 132.
(N. Gonzalez.)

A 6,2 9. Inca, INIajorca, Balearic 0. Thomas and R. I. 0. 7. 1. 18-23.

Islands. Pocock (p).

6 6, 9. San Cristobal, Minorca, 0. Thomas and R. I. 0. 7. 1. 59-65.

Balearic Islands. Pocock (p).

6. Cerbere, Pyr^n^es-Orien- O. Thomas and R. I. 0. 7. 1. 77.

tales, France. Pocock (p).

6. St. Genies, Gard. O. Thomas (p). 8. 8. 10. 92.
(C. Mottaz.)

Mus spiciLEGUs LUSiTANicus Miller.

1896. Mus sp. Thomas, The Zoologist, 3d ser., xx, p. 137, April, 1896.

1909. Mus sincilegus lusitanicus ililler, Ann. and Mag. Nat. Hist., 8th ser.,

Ill, p. 422, May, 1909.

1910. Mtis spicilegiis lusitanicus Trouessart, Faune Mamm. d'Europe,

p. 148.

Tyjie Jocalitij. — Cintra, Portugal.

Geographical distribution. — Known only from the vicinity of
the type locality.

Diagnosis. — General colour of upper parts brownish grey
with a decided tinge of russet.

Colour. — Upper parts a light yellowish wood-brown with an
evident russet tinge, the sides lighter and more huffy, the longer
dark hairs producing an evident though not very conspicuous
effect of " lining." A narrow clear buffy area along sides and
accentuating line of demarcation, this continued forward over
cheek but not surrounding eye. Underparts light cream-bufF,
the slate-grey bases of the hairs showing through at surface.
Feet buffy white or pale cream-bufF. Tail obscurely bicolor, buffy
whitish below, dark brown above, the dark area much wider than
in the Spanish form and essentially as in Apodemus sylcatieus.

Measurements. — Type (adult male) : head and body, 77 ; tail,
60; hind foot, 16*2; ear, 13. Average and extremes of six
adults from the vicinity of Cintra: head and body, 75 '5 (74-
77): tail, 61-8(60-65); hind foot, 16-1 (15-16-8); ear, 12-6
(12-13). For cranial measurements see Table, p. 881.

Specimens examined. — Six, all from the neighbourhood of Cintra.

4 <J, 2 9. Cintra, Portugal. 0. Thomas (c & p). 98. 2. 2. 30-35.

(98. 2. 2. 30. Type of subspecies.)

ACOMYS 883

Genus ACOMYS I. Geoffroy.

1838. Acomys I. Geoffroy, Ann. des Sci. Nat., Paris, 2cl ser., x, Zool.,
p. 126, August, 1838.

1841. Acosminthus Gloger, Gemeinn. Hand- u. Hilfsbucb der Naturgesch.,

I, p. 95 (dimidiatus).

1842. Acanthomys Lesson, Nouv. Tab!. Kegne Anim., Mamm., p. 135

{hispidus).

Type species. — Mus cahirinus E. Geoffroy.

Geographical distribution. — Africa and south-western Asia ;
islands of Cyprus and Crete. Not known from any part of the
mainland of Europe.

Characters. — In general essentially like Epimys, but fur
conspicuously spinous, and tail with unusually large scales, its
appearance in most species suggesting that of a lizard ; skull
with anterior half of mesopterygoid fossa closed by plate-like
outgrowth from the palatines, which meet and form a distinct
longitudinal ridge in median line, the open part of fossa thus
reduced to a small triangular space bounded chiefly by the
hamulars ; enamel pattern and relative sizes of molar crowns
about as in Epimys, but m^ three-rooted, its anterior lamina with
inner tubercle somewhat displaced backward, though less so
than in Mus.

Beinarks. — The genus Acomys contains about fifteen species,
one of which occurs within the limits of western Europe as here
defined.

ACOMYS MINOUS Bate.

1906. Acomys dimidiatiis minous Bate, Proc. Zool. Soc, Loudon, 1905, ii,

p. .321, April 5, 1906.
1910. Acomys dimidiatus minous Trouessart, Faune Mamm. d'Europe,

p. 158.

Type locality. — Kanea, Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Like Acomys dimidiatus of Asia Minor, but
smaller in size and darker in colour ; skull noticeably smaller
than in dimidiatus.

External characters. — Tail decidedly longer than head and
body, the scales large and coarse, forming about ten rings to the
centimeter at middle, the boundaries between the rings sharply
defined, but those between the scales in each ring obsolete.
Posterior border of each scale bearing two slender but distinctly
flattened, closely appressed bristles, the length of which equals
the width of about one and a half rings. These bristles are of
a peculiar silvery horn-colour, and are so arranged as to form
continuous longitudinal stria', which, in connection with the
conspicuous rings, impart to the tail a j^eculiar lizard-like appear-
ance. Among the bristles occur numerous longer terete hairs so

3 L 2

884

RODENTIA

slender that they can scarcely be detected without the aid (if a
lens. Hind foot with sole naked to heel, the usual six pads
well developed, the spaces between them occupied by numerous
secondary tubercles, some of which attain nearly the size of the
primary pads. Fore foot with thumb reduced to a mere
tubercle with rudimentary nail. Palm rather broad, with five
main pads and numerous secondary tubercles. Ear rather large,
extending about to middle of eye when laid forward, its surface
finely pubescent. Fur at middle of back consisting entirely of
flattened bristles about 11*5 mm. in length and about 0 • 4 mm.
in breadth at widest region. The few woolly hairs which occur
among the bristles are almost entirely concealed, though a few
extend beyond tips of bristles, especially on rump. On sides,
shoulders, head and underparts the bristles become so slender as
to resemble ordinary coarse hair.

Colour. — Ground colour of sides and upper parts bufl^, clear
and bright along sides, strongly tinged with ecru-drab on head,
neck and shoulders. Spines of back light bluish grey (nearly the
pearl-grey of Ridgway) through basal two-thirds, then bufi" with
the extreme tips horn-colour, the result being a peculiar coarse
grizzle in which the bufi" decidedly predominates. On hind leg
the buff extends down outer side to ankle, but on front leg it
does not reach wrist. Underparts and feet creamy white, the
line of demarcation sharply defined, and extending along sides
of face to muzzle. Ears and tail dull brownish, the tail not so
dark below as above.

Sliidl. — In general appearance the skull resembles a miniature
of that of Epimys rattus, owing to
the approximately similar develop-
ment and form of the ridges in
interorbital region and on sides
of brain-case. The brain-case is,
however, more depressed posteriorly
and relatively wider than in the
members of the black rat group,
the greatest width across lateral
ridges equal to fully 1^^ times
length of parietal along ridge.
Rostrum rather deeper and
narrower than in other European
Murinse. Interparietal very large,
strongly convex posteriorly, straight
or slightly convex anteriorly, its
area about equal to that of parietal.
Incisive foramina extending back
to about middle of first molar,
their length only about 1 mm.
less than that of diastema. Apart from its peculiar backward
extension in median line characteristic of the genus, the bony

Fig. 180.

Acoinyn minous. Nat. size.

885

palate is \"ery .sli<»rt<, its leiitftli from incisive foramen t<j the trans-
rerse ridge homologous with the normal posterior horder scarcely
exceeding breadth at middle of first molar. Hamulars diverging
posteriorly, their tips broadly in contact with well developed
beak of auditory bulla, the greatest distance between them about
equal to length of interpterygoid space. Auditory bullae moder-
ately large but not specially inflated, their form more elongated
transversely to main axis of skull than in members of the other
European genera. Zygomata strong, somewhat expanded
upward anteriorly (conspicuously more than in Epimys rattus or
Mus muscidus). Plate forming outer wall of infraorbital foramen
broad, rather sijuarely rounded oft" above. Infraorbital foramen
rather widely open below owing to the unusually slight pro-
tuberance formed by the root of upper incisor. Mandible with
long, rather narrow and straight angular process, the lower
border of which is folded squarely inward ; coronoid process
reduced to a mere recurved spicule scarcely rising above level of
wide, nearly horizontal area extending from its base to articular
surface. Protuberance over root of lower incisor slight.

Teeth. — Upper incisor strongly compressed, rather more so
than in Mus musculus, the cutting edge normal. Lower incisor
essentially like the upper tooth
in cross section, its anterior and
posterior diameters nearly equal.
Molars suggesting those of 3Iiis
musculus both in relative size and
in arrangement of tubercles, but
the general tendency toward
reduction carried less far, so
that crown area of m^ is about
equal to that of m'^ and nv'
combined. First upper molar
with crown relatively wider
than in 3Ius musculus, but
with tl similarly though less
displaced backward. Second
upper molar with /3 represented
by a rather evident antero-
external shelf, usually bearing
a minute tubercle ; /9 relatively
better developed than in the house mouse. Third upper molar
with crown area equal to fully one-half that of m'^ ; /3 usually
represented by a minute antero-external tubercle ; second and
third lamina " rather distinct, oblique, separated by a narrow-
groove, the extremities of which produce a sHght re-entrant
angle on each margin of crown, the tubercular elements entering
into formation of the laminre obsolete.* Lower molars as in

* In the specimen from which fig. 181 was drawn the crowns of the
upper molars are sufficiently worn to obscure the pattern, particularly of m\

Fig. 181.

Acomys minous. Cheek-teeth,

soinewhat worn. X 10.

886

Observations.

Teeth moderately worn.

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Number.

5.12.2.31
5. 12. 2. 32
5. 12. 2. 33*

H^

O

SPALAX 887

Mus miiscuJus, but anterior portion of ui^ broader and witli
two sub-ecjual tubercles nearly as large as tliose of second lamina,
and »»*.j relatively larger, its crown area slightly more than half
that of m.,.

3lL'aHiirements. — Type (adult female) : head and body, 94 ;
tail, 113 ; hind foot, 18 ; ear, 19 '5. Adult male from the type
locality: head and body, 112; hind foot, 19; ear, 18. For
cranial measurements see Table opposite.

Specimens examined ;- -
Cbete : Canea, 3.

2 S, 9. Caaea, Crete. Miss D. Bate (c). 5. 12. 2. 31--33.

(5. 12. 2. 33. Type of species.)

Family SPALACID^.

1821. Spalacidx Gray, Loudon Med. Repos., xv, p. 303, April 1, 1821.

Geographical distribution. — South-eastern Europe and adjoin-
ing portions of Asia ; south into Egypt.

Characters. — Form mole-like, highly modified for subterranean
life ; eye without external orifice ; external ear reduced to a low
ridge ; tail absent ; claws not specially enlarged ; skull wedge-
shaped, without postorbital processes, nearly half of its upper
surface occupied by a flattened, forward-sloping occipital shield,
the outer margin of which extends to level of outer border of
zygomatic root ; jugal short, extending forward only a little
beyond middle of zygoma and forming no part of anterior portion
of arch ; infraorbital foramen large, simple ; parapterygoid fossa
not closed dorsally, appearing like a large foramen ; cheek-
teeth ^|, rooted, hypsodont, the subterete crowns with one inner
and one or two outer re-entrant folds.

Remarks. — Both externally and in the structure of the skull
the family Spalacid-ee exhibits a higher degree of special adapta-
tion to a peculiar mode of life than any other group of rodents
occurring in Europe. As here understood the family consists of
the single genus Sjxdax.

Genus SPALAX Gueldenstaedt.

1770. Spalax Gueldenstaedt, Nov. Comm. Acad. Sci. Imp. Petrop., xiv,

pt. I, p. 410 (inicrophthalmus).
1783. Myospalax Hermann, Tabula Affinitatum Animalium, p. 83 {lax-

manni Hermann = Spalax microphthalmos Gueldenstaedt).
1799. Talpoides Lac6pede, Tableau des divisions, sous-divisions, ordres et

genres des Mammiferes, p. 10 {tijphlus).
1804. Aspalax Desmarest, Nouv. Diet. d'Hist. Nat., xxiv, Tab. Meth.

Mamm., p. 24 [Mus typihlus Pallas).
1815. A)wtis Rafinesque, Analyse de la Nature, p. 58 (Substitute for

Talpoides Lac6pede).

1840. ? Oiumatostcigus Nordmann in Keysciliiig aiicl Blasius, Wirbelthicre
Europas, pp. vii, 31 (O. I'O'llaHii Nordmann). Nomen nudum.

18'J8. Microspalax Nehring, Sitz.-Ber., Gesellsch. Naturforsch. Freunde,
Berlin, for December 21, 1897, p. 168 (Provisional sub-generic
name for smaller species of Spalax). Not Microspalax Trouessart,
l8S5.

1903. Nannospalax Palmer, Science, N.S., xvii, p. 873, IMay 29, 1903
(Substitute for Microspalax Nebring).

jfj/j^e species.- — Spalax micropJdhahnus Gueldenstaedt.

Geographical distribution. — South-eastern Europe and adjoin-
ing portions of Asia ; south into Egypt. Limits of range
imperfectly known.

Characters. — Essential characters as in the family Spalaeidve.
Fur soft and velvety except for a conspicuous line of stiffened
bristles extending backward along each side of head from outer
extremity of naked muzzle pad. Pattern of enamel folding at
first sigmoid, but later, as the crowns of the teeth wear away,
undergoing considerable changes in form, the re-entrant folds
tending to become isolated as enamel islands (fig. 184, p. 893).

Berimrks. — The members of the genus Spalax are immediately
recognizable among European rodents by their complete adapta-
tion to underground life, the absence of external eyes and tail,
and the presence of the peculiar line of stiffened bristles on each
side of head. About a dozen species have been described ; two of
these inhabit south-eastern Europe west of the Russian boundary,
while a third, of uncertain status, is supposed to occur in Greece.

KEY TO THE EUROPEAN SPECIES OF SPALAX.*

Skull not unusually deep, tbe distance from alveolus
of m^ to sagittal crest about 39 j)er cent of
condylobasal length ; width of auditory bulla
about 7 mm. ; greatest width of infraorbital
foramen slightly more than half its height

(Roumania and Bulgaria) .* S. dolbrogcT, p. 889.

Skull unusually deep, the distance from alveolus of
m^ to sagittal crest about 43 per cent of condy-
lobasal length ; width of auditory bulla about
8 mm. ; greatest width of infraorbital foramen
conspicuously more than half height.
Breadth of rostrum in adult male about 11 mm. ;
alveolar breadth of m^ equal to least palatal

width (Hungary) .S'. liungaricus, p. 894.

Breadth of rostrum in adult male about 12 mm. ;
alveolar breadth of m^ less than least palatal
width (Greece?) S. gr.rcns, p. 895.

* The important work by Dr. L. M6hely on this genus — Species generis
Spalax. A Fold! Kutyak Fajai, Budapest, 1909 — has unfortunately been
received too late to be utilized by the author.- 0. T. (See Appendix,
p. 1000.)

S89

SPALAX DOLBROGEiE Miller.

1901. Spalax huiigarican ]\[atscliie, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 237, November, 1901 (Eoumania).
1903. Spalax dolbrogav Miller, Proc. Biol. Soc, Washington, xvi, p. 161,

November 30, 1903 (Dobrudscha). Type in U.S. National

Museum.
1910. Spalax dolbrogen- Troucssart, Faune ]Mamm. d'Europe, p. 204.

Type locality. — Malcoci, Dobrudscha, Rouuiania.

Oeographical distribution. — Roumania and Bulgaria ; limits of
range not known.

Diagnosis. — Size medium (hind foot about 25 mm., condylo-
basal length of skull about 50 mm.) ; skull moderately deep, the
distance from alveolus of )»•' to sagittal crest about equal to that
from occipital condyle to first molar ; anteorbital foramen large,
much higher than wide ; anterior base of zygoma lightly built ;
auditory bulli¥i rather long and narrow ; breadth of m^ at alveolus
less than width of palate in same region.

External characters. — General form cylindrical, with long
body, short legs and no apparent neck, the broad flattened head
set so closely on shoulders that orifice of ear is almost at anterior
edge of scapula. Head broadly rounded when viewed from
above, wedge-shaped in profile owing to the abrupt reduction in
depth from behind forward, the anterior border of muzzle pad
forming the front edge of wedge, the chin strongly receding ;
external ear absent, the small auditory opening margined by a
low ridge , no trace of external eye ; nostril pad large, lying in
two distinct planes, one on upper surface of muzzle, the other
on abruptly backward-sloping upper lip ; upper portion of pad
broadly triangular in outline, its longest border in front, its
surface smooth except for some indistinct wrinkles crossing it
longitudinally, and a deep groove at each side parallel with
margin ; lower portion similar to upper in general outline, the
simple, round nostrils situated a little below middle ; a narrow
groove curving outward and upward from each nostril to anterior
margin of pad ; lij) thinly haired between pad and mouth. The
shovel-like character of upper surface of head is increased by
the presence of a brush of stiffened hairs directed outward and
upward, and extending from outer angle of muzzle pad about
half-way to ear. Legs short, the feet large and distinctly
fossorial. Fore foot with third digit longest, second and fourth
successively a little shorter, fifth extending to end of first
phalanx of fourth, first very short, not extending to middle of
first phalanx of second ; claws simple, very small, their length
not equal to that of terminal phalanx ; palm naked, the region
at base of phalanges wrinkled but without tubercles, the posterior
portion occupied by a large pad divided into two unequal parts
by a longitudinal fui'row on side next thumb, the anterior
margin raised and thickened, sometimes appearing as a ridge

S90 RODENTIA

crossing palm obliquely outward and backward from base of
thumb. Hind foot longer and more slender ; proportions of
digits as in fore foot, but claws of second and third about three
times as long as the othei*s, which are of the usual small size ;
in old individuals the second and third claws may be worn down
to essentially the same size as the others ; sole naked, finely
wrinkled ; three small, ill-defined tubercles at bases of main
digits, and a fourth at base of hallux ; a faintly indicated fifth on
opposite side of sole from fourth. Tail a mere tubercle barely
1 mm. in height.

Colour. — Back, sides and posterior half of head a light
yellowish brown somewhat paler than the ochraceous-buff" of
Ridgway, the fur everywhere slate-grey beneath surface ; under-
parts and legs slate-grey, the yellowish brown of sides usually
extending as an evident wash across middle of body ; face, cheeks
and region about mouth silvery drab-grey, the two lines of
bristle-like hairs extending back from muzzle whitish in rather
marked contrast.

Skull. — The skull is heavily-built, its surface conspicuously
ridged, its general form strongly cuneate whether viewed from
above or from the side. Dorsal profile slightly convex from
front of nasal to lambdal region, the convexity most marked
between orbits, the nasals and upper surface of brain-case often
flattened or faintly concave ; occiput very gradually sloping at
an angle of about 10°, the length of this sloping portion equal to
distance from lambda to posterior extremity of nasals ; ventral
profile concave immediately behind incisor, then sloping abruptly
downward to front of alveolus, from which point it is essentially
horizontal to back of occipital condyle. Brain case broadly ovate
in outline when viewed from above, the broad occipital shield
causing it to appear almost triangular ; sagittal crest well
developed in adult, extending from lambdoid crest to interorbital
constriction ; occipital shield extending so far forward that the
conspicuous, almost squarely transverse lambdoid crest is con-
tinuous at its outer extremity with anterior border of zygomatic
root, into basal portion of which the antero-external angle of
shield appears to be pressed ; outer border of shield extending as
a thin plate from auditory meatus to base of zygoma, and roofing
over a pit-like area lying between glenoid surface and neck of
bulla ; posterior aspect of brain-case entirely covered by the
pentagonal occipital shield, no portion of brain-case or inter-
orbital region appearing above its upper margin ; lower border
of shield longest, postero-external border shortest, antero-external
border intermediate in length ; foramen magnum squarish in
outline, its height less than half that of shield ; paroccipital
processes distinct but small, not extending to level of lower
border of foramen magnum ; condyles projecting noticeably
behind all other parts of occiput ; base of brain-case with no
specially remarkable features ; width of basioccipital behind

SPALAX 8'Jl

nearly twice length excluding condyles, width along anterior
suture about two-thirds median length ; surface of basioccipital
irregularly depressed at side: auditory bullie rather large, but
displaced upward so that lower border is about on level witli
that of foramen magnum, their outline somewhat flask-shaped,
the neck of the flask projecting horizontally outward, not
abruptly defined from body of bulla ; surface of bulla smoothly

Fig. 182.
Spalax (hilbroyex. Nat. size.

polished. Interorbital region deeply constricted, its upper
surface marked by a narrow median groove extending forward
from sagittal crest. Zygoma slender and weak, not expanded at
middle, nearly horizontal throughout, bowed gradually outward,
so that the greatest zygomatic breadth is at level of anterior
extremity of glenoid surface ; infraorbital foramen large, broadly
crescentic in outline, its greatest width scarcely or not more

892

equal to that of alveolus,

tliau lialf lieight. liostruiu ratlier long, nearly iiacallel-suled
wlieii viewed from above, though shghtly wider somewhat
beyond middle than at base, and tapering rather noticeably in
front, its width everywhere greater than that of iuterorbital
constriction ; nasals wide, abruptly rounded off anteriorly, where
they project so as to conceal incisors, tapering gradually to a
narrow base, wliere they are abruptly truncate at level of
posterior extremity of nasal branches of premaxiilaries, and of
middle of infraorbital foramina ; in lateral view the rostrum is
strongly cuneate, its least depth, immediately behind incisors,
contained about 2| times in that at alveolus of m^ ; incisive
foramina very small, slit-like, at level of anterior border of
zygomatic root, their greatest diameter contained about six times
in diastema (three times in space in front of them, twice in
that behind). Palate very narrow, its least breadth scarcely

its surface marked by a median
ridge and two lateral grooves ; pos-
terior termination of palate simple,,
without pits or changes of level ;
mesopterygoid space parallel-sided,
about twice as long as wide, squarely
truncate anteriorly at level a little
behind that of alveolus of m^ ;
hamulars simple, straight, slightly
diverging, widely separated from
bulla5 ; ectopterj^goid narrow, the
pit deep and widely open at bottom.
Mandible slender, appearing dispro-
portionately small for the very large
incisor which projects conspicuously
behind as well as in front, and to
which the ramus is little more than
a sheath ; coronoid process high,
slightly recurved, its anterior border rising abruptly from level
of front of 7n.2 ; articular process low, its base distorted by the
shaft of incisor ; angular process with rather long, strongly
convex lower border, its free portion short, abruptly curved
outward, not extending behind level of condyle ; between
articular and angular processes and somewhat in front of them
projects outward and upward the encapsuled base of the incisor,
rising in fully adult individuals to a height intermediate between
that of articular and coronoid processes, and forming the most
conspicuous single feature of this part of jaw.

Teeth. — Incisors large and heavy, molars small, the contrast
greater than in any other European rodent. Upper incisor strongly
curved, the exposed portion nearly vertical, the root forming a
slight protuberance at side of palate immediately in front of
first molar ; shaft not compressed, the width of flat anterior face
nearly ecjual to greatest anteroposterior diameter ; posterior

Fig. 183.

S'palax ddlbrogese. Skull from
behind. Nat. size.

SPALAX 8U3

border narrowly rounded ; inner and outer borders about e({ual ;
enamel extending slightly on inner and outer sides of tooth, its
surface finely rugose with minute longitudinal wrinkles, and light
orange in colour. Lower incisor so large that the mandible has
the appearance of a mere sheath to the tooth, its root projecting
between articular and angular processes as a conspicuous sub-
terete capsule, larger and more noticeable than the processes
themselves ; length of capsule increasing with age, scarcely or
not attaining level of condyles in individuals whose molars are
in the early stages of wear, though surpassing this level in

Spalax (hilbrofjex. Cheek- teeth showing enamel pattern at various stages of wear.
Vpper row, maxillary teeth ; lower row, mandibular teeth (semi-diagrammatic). X 5.

older animals ; section of shaft much as in upper tooth, but
antero-posterior diameter relatively greater ; surface of enamel
less evidently rugose, and colour whitish or slightly tinged with
yellow. Molars with high, sub-terete crowns and short roots, of
which there are three in each of the maxillary teeth, the inner
root tending to divide longitudinally into two, and two in each
of the mandibular teeth, each root tending to divide into two ;
size of teeth decreasing from first to third, the crown area of
third when unworn about one-half that of first, when worn
relatively larger; enamel pattern of")/(' and m^ consisting of two
backwardly curving re-entrant folds on outer side and a single

•^94 RODENTIA

foward-curving re-entrant fold on inner side, the anterior outer
fold of m"^ shallower than the others and usually becoming
converted into an " island " at an early stage of wear ; ?«•' with
re-entrant fold on either or both sides ; lower molars each with
a forward-curving more anterior inner fold and a backward-
curving more posterior outer fold, the inner fold in m^ and 5J«2 with
a small supplemental l)asal backward fold which early becomes
isolated as a minute island lying near apex of outer fold ; all
the re-entrant folds in both maxillary and mandibular teeth are
transformed into islands as the wearing away of crown advances.
Measurements. — Type (adult male) : head and body, 230 ;
hind foot, 25 (hind foot with claws, 29). For cranial measure-
ments see Table, p. 896.

Specimens examined. — Nine, from the following localities : —
Bulgaria : Near Rustschuk, 6 skulls (Andersen).
Roumania: Malcoci, Dobrudscha, 3 (B.M. and U.S.N.M.).
S, ?. Malcoci, Dobrudscha, Rou- G. Dalgleish (c). 5. 10. 25. 1-2.
mania.

SPALAX HUNGARICUS Nehrinsf.

1898.

Spalax typhlus hungariais Nehring, Sitz.-Ber. Gesellsch. Natur-

forsch. Freunde, Berlin, for December 21, 1897, p. 173, fig. 3.
Spalax huiigaricus Nehring, Zool. Anzeiger, xxi, p. 479,

September 5, 1898.
1898. ? Spialax monticola Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 6 (Ulog-Obruja, Herzegovina).
Spalax huncjaricus and ? S. monticola Trouessart, Faune Mamm.

d'Europe, pp. 203, 204.

1898.

1910.

Tyye locality. — Hungary ; exact locality not stated.
Geographical distribution. — Hungary ; probably extending
southward into north-western portion of Balkan Peninsula.

Fig. 185.

Anterior base of zygoma in Spalax dolhroijex (a)

and S. hungaricus (,b). Nat. size.

S PAL AX 895

Diagnosis. — Like Spalax dolbrogese, but skull tending to be
somewhat deeper ; anteorbital foramen moderate, nearly as wide
as hiofh ; anterior base of zygoma heavy ; auditory bulke rather
short and wide ; breadth of iii^ at alveolus about equal to width
of palate in same region.

Specimens examined. — Austria-Hungary, Kolozsvar, 2 ; Hungary, no
exact locality, 2 (U.S.N.M.).

2. Kolozsvar, Hungary. 0. Thomas (p). 3. 1. 14. 1-2.

(L. Fiihrcr.)

SPALAX GR^CUS Nehring.

1898. Spalax grivcus Nehring, Zool. Anzeiger, xxi, p. 228, March 21, 1898.

Type in Munich Lluseum.
1910. Spalax gnvcits Trouessart, Faune Mamm. d'Europe, p. 204.

Type localiti/. — Supposed to be the neighbourhood of Athens,
Greece.

Geographical distrihution. — Known only from the two original
specimens and a young male from Stylis too immature to be
positively identified.

Diagnosis. — Similar to Spmlax hungaricus, but skull with
somewhat broader rostrum and palate.

External characters and colour. — As in *S'. hungarictis.

SJcull and teeth. — In all respects the skull and teeth resemble
those of Spalax hungaricus, as opposed to S. dolhrogese (great
depth through frontal region ; form of auditory bullae ; width of
infraorbital foramen), but the rostrum appears to be slightly
broader and the nasals a little more spatulate than in the
northern animal. Teeth exactly as in >S^. hungaricus, except that
molars are relatively somewhat narrower, a peculiarity that may
readily prove to be individual. The enamel pattern of the adult
agrees with that of S. hungaricus in all respects except in the
slightly less development of the longitudinal terminations of the
outer re-entrant folds, a peculiarity probably due to the condition
of wear in which the crowns happen to be.

Measurements. — Hind foot of adult, 27 mm. For cranial
measurements see Table, p. 896.

Specimens examined. — The skeleton and mounted specimen in the
Munich Museum which formed the basis of the species : also an immature
male from Stylis.

Bemarks. — Through the kindness of the authorities of the
Munich Museum I have had the opportunity to compare the
original specimens of Spalax grsecus with S. hungaricus and S.
dolhrogese. Since the species was described the skull has been
removed from the adult individual. This specimen shows that
the teeth, contrary to the supposition of the original describer,
have no characters that can be regarded as specific. If distinct
from S. hungaricus, the larger animal is not likely to be separable
by any other character than its size.

6. Stylis, Phthiotis, Greece. E. MacDonell (c & p). 9. 3. 15, 1.

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Familv SCIURIDiE.

1821. Sciurich'. Gray, London Med. Repos., xv, p. 301, April 1, 1821.

Gcofjrajjliicdl distriJuUion.-, — Eastern and western heinispliorcs
except at tlie extreme north and south ; absent from Madagascai',
New Guinea, AustraHa and the Pacific Islands.

Cltaracters. — Form slender or robust according as the animal's
habits are arboreal or terrestrial ; tail without horny scales ou
lower surface, densely long-haired, usually flattened ; ribs twelve
or thirteen pairs ; skull varying in form but never highly
modified ; postorbital processes present ; infraorbital foramen
small ; jugal long, its anterior extremity wedged between
lachrymal and zygomatic process of maxillary ; palate rathei"
broad ; upper incisor with root in front of anterior cheek-teeth,
lower incisor with root not extending conspicuously into
ascending portion of ramus ; molars rooted, tubercular and
transversely ridged, premolars -| or -}.

RemarJiS. — ^The family Sciuridse, containing the scfuirrols,
spermophiles, marmots and their allies, is one of the most widely
distributed and abundantly represented grou^j of rodents. It
ct)ntains about forty-five living groups, the exact status of many
of which is still in doubt ; three of these occur in Europe.

KEY TO THE EUROPEAN GENERA OP SCIURIDSE.

(For convenience of comparison the cranial and dental cliaractcrs of
Sciuroptcnts are inserted.)

[Orbit large, its vertical diameter equal to more
than one-third basilar length of skull ; posterior
transverse ridge on crown of first and second
upper molar terminating internally in a distinct
cusp isolated by deep groove ; axes of enamel

foldings of lower cheek-teeth not parallel Sciuruptcnta, p. 'Jll.]

Orbit moderate or small, its vertical diameter equal
to less than one-third basilar length of skull;
posterior transverse ridge on crown of first and
second upper molar terminating internally with-
out cusp and groove ; axes of enamel foldings of
lower cheek-teeth parallel.
Incisors compressed, the width of anterior face not

more than half that of lateral face ; small

upper premolar mini;te, its crown simple, with

diameter equal to about one-eighth that of

large premolar ; dorsal profile of skull strongly

depressed posteriorly ; tail nearly as long as

head and body, very bushy (Habits arboreal,

Squirrels) Sciurus, p. 898.

Incisors not comiJi'essed, the width of anterior face

about equal to that of lateral face ; small

upper premolar well developed, its crown

marked by evident ridges and depressions, its

diameter about half that of large premolar;

dorsal profile of skull not very strongly de-
pressed posteriorly (Habits terrestrial).

3 M

yOS RODENTIA

Braiu-case not excessively flatteued, its depth
about twice that of rostrum ; distance from
lachrymal to tip of postorbital process equal
to about one-third condylobasal length of
skull ; size medium or small, form squirrel-
like (Ground squirrels) Citellus, p. 924.

Brain-case much flattened, its depth not con-
spicuously greater than that of rostrum ;
distance from lachrymal to tip of postorbital
process less than one-fourth condylobasal
length of skull ; size large, form badger-
like (Marmots) Marmola, p. 931.

Genus SCIURUS Liuiianis.

1758. Scinrus Linna3us, Syst. Nat. i, 10th ed., p. 63 {vulgaris, by

tautonymy).
1857. Scimus Blasius, Saugethiere Deutschlands, p. 271.
1893. Aphrontis Schulze, Zeitschr. fiir Naturwissensch., Leipzig, lxvi,

p. 165 (vulgaris).

Tjijje f<peeies. — Sciuriis vuhjar/'s Liuiu«us.

Geograpliical (listribiitiou. — Forested regions of the northern
hemisphere, in the Old World from Ireland to Japan and north-
eastern Siberia ; southern limits of distribution at present
uncertain owing to lack of precise definition of the genus.

Characters. — Strictly arboreal Sciuridse, of medium size and
typically sciurine aspect, the tail bushy and conspicuously
flattened, more than half as long as head and body ; skull witla
deep, strongly convex brain-case and weak rostrum ; postorbital
processes slender, directed backward ; incisors strongly com-
pressed ; anterior upper premolar vestigial, usually present as a
practically functioidess terete spike ; crowns of molars low, those
of upper teeth with two moderately developed cross ridges and a
broad, low inner cusp (not forming a (J -shaped pattern with
wear), those of lower teeth distinctly basin-shaped.

Reviarks. — In the present uncertainty regarding the classifica-
tion of the squirrels, it is impossible to frame a satisfactory
diagnosis of the genus Sciurns, or to estimate the number of
forms that should be referred to the group. Only the type
species occurs within the limits of the present work ; it is repre-
sented by thirteen geographical races.

SCIURUS VULGARIS Linmeus.

(Synonymy under subspecies.)

Geograpliical distribntion. — Wooded portions of Europe from
the extreme northern limits of tree growth to the Mediterranean
coast, and from Ireland eastward into Asia.

Diagnosis. — Size medium (head and body ranging in the
diJTerent races from 205 to 280 mm., tail from 160 to 242, hind
foot from 55 to 66, condylobasal length of skull from 45 to 55) ;

SCIURUS 899

tail decidedly more! than half as long as head and body ; ear
conspicuously tufted in winter ; colour reddish, greyish or dark
brown above, whitish (never distinctly yellowish) below ; minute
anterior upper premolar present ; cross-ridges on upper molars
smooth, not tending to become tuberculate ; lower incisors
extremely compressed.

External characters. — Form slender, the head and neck well
differentiated from the trunk, the legs rather long, the tail about
eipial to head and body. Head rounded posteriorly, narrowed in
front, the muzzle deep and not much produced (distance from eye
to nostril about e(iual to depth at region half way between) ; ear
rather large, just covering eye when laid forward, the "width at
middle about three-quarters height from crown ; anterior bordei-
nearly straight below, somewhat convex above, its lower two-
thirds abruptly folded backward, the inner root of this fold lying
nearly over meatus ; tip rather abruptly rounded oft' ; posterior
border convex except for a flattened or slightly convex region
just below tip, its lower border extending forward as an abruptly
angled projection over meatus ; outer surface of ear densely
furred at all seasons, the tip conspicuously tufted in winter
pelage ; muzzle pad hairy above, a narrow bare area extending
between nostrils and across upper lip and sending out a
lateral branch under each nostril. Fore feet long and slender
with long, freely spreading fingers ; thumb reduced to a mere
tubercle with closely appressed nail ; two middle fingers sub-equal
and longest, second and fifth sub-equal and about lialf as long ;
palm with five tubercles, the two posterior largest and best
defined ; claws long, compressed, strongly curved. Hind foot
long, the five toes all well developed and bearing strong claws
shorter and less curved than those of fore foot ; iinier toe
extending about to base of others, outer extending a little beyond
middle of fourth ; fourth toe longest, third and second successively
a little shorter ; sole densely hairy in winter, partly bare in
sunnner ; four small but evident tubercles at bases of toes. Tail
strictly distichous throughout, the hairs at sides and above about
dO to 50 mm. in length. Mammaj, p 1 — 1, a 2 — 2, i 1 — 1 = 8.

Culour. — Though the actual colour shows great difi'erences
according to phase and season the pattern is constant and
characteristic. The head, back, sides and tail are essentially
uniform, the tail usually a little darker than back, the sides of
body and outer surface of limbs somewhat lighter, the middle of
back frequently with more red ; face with an ill-defined pale area
on cheeks and muzzle ; ear-tufts usually concolor with sur-
rounding parts but sometimes darker or more i-ed ; irnier surface
of legs and upper surface of feet not so dark as outer surface of
legs ; tail with a lighter, grizzled or whitish area along median
line below, this sometimes partly or wholly concealed ; white of
underparts extending forward to chin or to back of interramial
space ; posteriorly the white does not reach base of tail, and

3 M 2

900

RODENTIA

laterally it barely extends on inner surface of thigh and forearm ;
line of demarcation well defined. Two colour phases occur, one
characterized by the predominance of red, the other by that of a
dark, sometimes almost blackish brown. The dark phase is often
incorrectly spoken of as melanistic. In a general way the phases
have a geographical significance, as the red is dominant at the
north, the dark brown at the south. The winter pelage is
everywhere lighter, brighter and more greyish than the summer
pelage. In regions where both colour pliases occur four normal
combinations of phase and pelage will be found in any consider-
able series of specimens. Apai't from these combinations and
their intermediate stages individual variation in colour is not
unusually great.

Sktdl. — In general the skull is broad, smooth and rounded,
with large, deep brain-case and small rostrum, the basicranial

Fig. 186.
Sciiirus vulgaris. Nat. size.

axis bent downward so that foramen magnum is entirely below
level of alveolar line. Dorsal profile flat or faintly and irregularly

SCIURUS 901.

convex to region between postorbital processes, then abruptly
and strongly convex over greater portion of brain-case to a slight
but evident concavity immediately in front of lambdoid region ;
ventral profile somewhat parallel to dorsal profile in general
direction, but diverging slightly in its anterior half and con-
verging a little posteriorly ; posterior truncation of brain-case
nearly vertical, slightly convex when viewed from the side, the
occipital condyles not visible from above. Brain-case broadly
ovate in outline when viewed from above, its greatest width
about equal to length, its width posteriorly somewhat greater
than that at postorbital constriction ; surface essentially smooth,
though in old individuals, particularly of the larger races, a low
ridge extends backward from base of postorbital process, across
middle of parietal to lambdoid region, the two ridges together
forming a lyrate figure ; lambdoid crest represented by a mere
angular ridge along course of lambdoid suture ; viewed from the
side the brain-case is conspicuously rounded through anterior
three-fourths, the low occipital region marked off from main
portion by a shallow but evident constriction ; occiput moderately
high, its depth to lower lip of foramen magnum about two-
thirds mastoid breadth, much exceeded by median portion of
l)rain-case, its dorsal and lateral margins forming an evenly
rounded, nearly semicircular arch ; foramen magnum wider than
high. ; occipital with three distinct swellings between foramen
magnum and lambdoid ridge, each about equal to exposed portion
of petrosal ; paroccipital processes small, their extremities hooked
inward and slightly forward, scarcely extending to level of lower
edge of condyle ; basioccipital rather broad and short, its width
in region of suture about equal to median length ; median ridge
slightly developed ; edge of basioccipital in region of suture
slightly raised, but not forming a distinct lateral process ;
auditory bullfe sub-circular in general outline, rather evenly
inflated, but with an evident antero-internal flattening, their
antero-posterior diameter equal to distance from front of bulla
to front of «!•', their lateral diameter to inner margin of meatus
bfirely equal to least width of basioccipital; meatus with a
distinct rim except along postero-upper border. Interorbital
region about as broad as long, its least width nearly the same
as postorbital constriction, its surface nearly flat, though usually
with a low but evident swelling at each side posteriorly, the
bases of postorbital processes about on level with median region ;
notch at anterior base of postorbital process small but well
developed, the orbital border sometimes complete so as to trans-
foi'm the notch into a foramen ; postorbital processes long, slender,
strongly bent downward, the orbit surrounded by bone througli
about three-quarters of its circumference. Zygomata not widely
spreading either in front or behind, the arches essentially parallel
with each other through tlie gi-eater part of their extent ; jugal
projecting posterioi'ly lj(^hind zygomatic process uf squamosal, its

902 RODENTIA

upper horder rising at middle to form a slight but evident angle
marking hinder margin of orbit, its anterior extremity wedged
lietween lachrymal and zygomatic process of maxillary ; anteorbital
foramen small and slit-like, a little in front of level of anterior
pi*emolar, its outer border strongly concave, its lower extremity
marked by a short abruptly projecting process. Rostrum forming
about one-third length of skull, its base rather broad, its anterior
region narrow and compressed, the least depth behind incisors
decidedly greater than width at same region ; nasals rather short,
moderately narrowed posteriorly, their squarely truncate hinder
border lying at level of extreme anterior edge of zygomatic root,
the broadly expanded nasal branches of premaxillaries usually
extending somewhat further back. Palate moderately wide,
slightly concave antero-posteriorly, rather noticeably concave
laterally between tooth-rows ; incisive foramina small, lying
entirely in premaxillaries, their length barely more than one-
third that of diastema and much less than least width of rostrum ,
their anterior extremities very narrow ; posteriorly the palate
extends a little beyond level of last molar, its border truncate
or convex, never with median spine ; a conspicuous foramen or
notch at outer edge of palate behind alveolus of )//■' ; mesop-
terygoid space less than twice as long as wide, its lateral borders
nearly parallel except posteriorly, where the hamulars curve
rather noticeably outward ; ectopteiygoid slightly developed and
inconspicuous. Mandible robust, the ramus deep and noticeably
compressed in front of tooth-row ; coronoid process broad at base,
tapering rapidly to an acute recurved point a little above level
of condyle, its antei'ior border moderately convex ; articular
process with shallow but noticeable concavity on outer surface ;
angular process well defined by the deep curve of its lower
boi'der.

Teeth. — Upper incisor robust, l)ut course of shaft not con-
spicuously marked on side of rostrum, and root producing no
visiljle protuberance ; shaft strongly compressed, its cross section
elliptical in outline, flattened in front, the enamel very thin,
extending backward slightly on both outer and inner side, and
covering less than one-third circumference of tooth, its surface
finely pitted, yellow ; lower incisor with root extending into
ascending portion of ramus and forming a slight protuberance
on outer surface a little above level of crowns of molars, its shaft
much more compressed than that of upper tooth, its outline
ovate, narrowest in front, the enamel confined to narrow anterior
surface. Anterior upper premolar a simple terete spike about
as large as anterior outer root of succeeding tooth, its crown
when unworn with slightly developed anterior tubercle and
postei'ior depression, the area of crown barely more than one-
tenth that of j«?i*. Molariform teeth resembling each other in
general size and form, each with two roots on outer side and a
single largei' I'oot on inner side, the innei- border of crown formed

sciURUs 903

by a single low, elongated tubercle, its height much less than
length of base, the width of base scarcely more than half its
length, the median portion concave transversely, the concavities
of the four teeth forming together a longitudinal furrow, the
outer side, except of m'\ with four low cusps from which trans-
Averse ridges extend inward on or across crown, their general
direction somewhat obliquely forward. The usual scheme of
the outer cusps and their ridges is as follows : first cusp low,
its ridge forming anterior margin of crown and extending to
inner extremity of base of inner tubercle ; second cusp nearly
twice the height of first, its ridge extending to side of base of
inner tubercle ; third cusp like first, its ridge extending only
about half way across crown ; fourth cusp like second, its ridge
extending to slightly behind middle of base of inner tubercle ;
posterior base of inner tubercle continued outward along hinder

Fia. 1S7.
Sciiirtif! vulgaris. Cheek-teeth, x

border of crown as an ill-defined ridge extending nearly to
base of fourth outer cusp. The principal deviations from this
pattern are : in pm*, first outer cusp nearly as large as second,
fourth transverse ridge with an evident tubercle near its inner
extremity ; vv^ all the outer cusps together with their transverse
ridges obsolete except the second, the posterior two-thirds of
crown occupied by a shallow basin-like depression. Lower pre-
molar with three roots, lower molars with four. Crowns squarish
in outline, that of m^ longer than broad, the others slightly
broader than long, that of j'^'h with antero-external border so
rounded off as to give the ci'own a somewhat triangular outline.
Main portion of crown occupied Iw a smooth depression, on both
outer and inner edge of which are thrcio low tubercles, the
anterior and posterior outer ha\ing the broadest and best
defined bases, while the anterior internal is the highest ; on each

904 RODENTIA

side the median tubercle is much smaller than either of the two
1 let ween which it lies.

Bemarls. — The common squirrel in any of its geographical or
seasonal phases is so readily distinguishable among the mammals
of Europe as to require no special comparisons with other
members of the fauna.

KEY TO THE EUROPEAN FORIMS OF SCIURVS VULGAEIS.

White of underparts restricted to median region ;

hind foot about 50 mm. (Southern Spain) S. v. IkMcux, p. 923.

White of underparts not restricted to median region ;
hind foot 55 to 70 mm.
Tail drab, fading in spring and summer to pale
buf^, never strongly reddish or blackish (British

Islands) S. v. levcowvs, p. 907.

Tail varying from red to black, never drab or buff y.
Condylobasal length of skull usually about 49 to
55 mm. ; mandible 34 to 37-6 mm. (Spain).
Tail without white on under surface ; skull

49 to 51 mm. (North-central Spain) ,S'. r. numantius, p. 914.

Tail usually with conspicuous whitish median
area on under side (this sometimes repre-
sented by a few white hairs only).

IMuzzle and cheeks rusty (Central Spain) S. v. -infii.watvs, p. 91G.

Muzzle and cheeks whitish or pale grey

(South-western Spain) S. >\ aegune, p. 917.

Condylobasal length of skull usually about 44 to
49 mm. ; mandible 31 to 34 mm.
Dark brown phase essentially absent (North-
ern).
Winter pelage smoke-grey, the back strongly
tinged with red (Scandinavian Penin-
sula, except extreme north) ,S'. r. mhjuri^, p. 905.

Winter pelage pearl-grey, the back scarcely
or not tinged with red (Extreme north
of Scandinavian Peninsula, east into

Russia) S. ■». vfiriiis, p. 906.

Dark brown phase not infrequent, sometimes
dominant (Central and southern).
Upper parts in dark brown phase much
darker anteriorly than posteriorly

(Greece) .". ,s:. ?•. l!l:eu.-r,^. 913.

Upper parts in dark phase not uoticeablj-
darker anteriorly than posteriorly.
Rostral portion of skull unusually short

and broad (Pyrenees) S. v. alpinn.<>, p. 914.

Rostral portion of skull normal.

Winter pelage of red phase essentially
without suffusion of grey on sides

(Italyj S. V. itaHaia, p. 912.

Winter pelage of red phase strongly
suffused with grey on sides.
Red. in both winter and summer,
clear bright rufous (East-central

Europe) S. v. fum.ater. p 910.

Red in ooth winter and summer,
rufous strongly tinged with chest-
nut (West-central Europe) ,S'. v. nissu.':, p. 909.

sciuRtJs 905

SCIURUS VULGARIS VULCxARIS Linnpfus.

1758. [Sciiinis'] vulgaris Linnaeus, Syst. Nat., i, lOtb ed., p. 63.

1792. Sc[mms'] vulg[aris] rufus Kerr, Anim. Kingd., p. 255 (Tlenaming of
vulgaris).

1827. Sciiirus vulgaris /3. alho notatus Billberg, Synopsis Faunre Scan-
dinavia, p. 2 (Southern Sweden).

1827. Sciurus V2ilgaris y. alhus Billberg, Synopsis Faunas Scandinavia?,
p. 2 (Skiine, Sweden).

1827. Sciurus vulgaris 5. iiiger Billberg, Synopsis Faunas Scandinavia?,
p. 2 (Skune, Sweden).

1857. Scitirus vulgaris Blasius, Saugethiere Deutscblands, p. 272 (part).

1899. Sciurus vulgaris typicus Barrett-Hamilton, Proc. Zool. Soc,
London, p. 6.

1910. Sciurus vulgaris Trouessart, Faune ]\Iamm. d'Europe, p. 117.

Tijijc localiiy. — Upsala, Sweden.

Geograpldciil distrihutum. — Scandinavian peninsula except
extreme 'northern portion. Exact northern limit of range not
known.

Diagnosis. — True dark brown phase absent or extremely rare,
though tail often considerably blackened. Summer pelage :
body dark rufous tinged with chestnut ; tail clear dark rufous.
Winter pelage : body grizzled smoke-grey strongly tinged with
chestnut along middle of back ; tail rufous chestnut.

■ Measurc'iiicnis. — Average and extremes of eight adults from
the type locality: head and body, 209 '7 (203-215) ; tail, 178-7
(172-189); hind foot, 59-1 (58-60). For cranial measurements
see Table, p. 918.

Specimens examined. — Fifty-three, from the following localities : —
Norway: Tolgen, Hedemarken, 1; Foldalen, Hedemarkcn, 1; Gausdal,
Kristiansamt, 1 ; Sundal, Bergenhus, 2 ; Konnerud, Buskerud, 1 ; Spjotsod,
Telemarken, 3 (U.S.N.M.) ; Aas, Akershus, 2; Aker, Akershus, 12 (B.M.
and U.S.N.M.) ; Asker, Cbristiania, 2 (B.M. and U.S.N.M.) ; Maersness,
Christiania, 1; Holmestrand, Jarlesberg, 1; Kristiansand, Maudal, 2;
Frederickshold, Smaalenene, 2 ; Cludbrandsdal, 1 ; no exact locality, 4.
Sweden : Upland, 2, near Upsala, 15 (B.M. and U.S.N.M.).

9. Tolgen, Hedemarken, Christiania Museum (e). 97. 3. 13. 12.

Christiania Museum (e). 97. 3. 13. 13.
Christiania Museum (e). 97. 3. 13. 14.
C. B. Horsburgh (c & p). 99. 9. 5. 1-2.
Christiania Museum (e). 97. 3. 13. 21.
Christiania Museum (e). 97. 3. 13. 17-18.
Christiania Museum (e). 93. 3. 11. 8.
Christiania Museum (e). 97. 3. 13. 1-11.

Eev. H.H. Slater (c&p). 99. 11. 27. 5.
Christiania Museum (e). 97. 3. 13. 22.

Christiania Museum (e). 97. 3. 13. 15-lG.
Frederikshold, Smaale- Christiania Museum (e). 97. 3, 13. 19-20.

nene.
Gudbrandsdal. O. Barrett - Hnmiltou 11.1.2.151.

[J. L. lluuhvtv.) (!■).

Norway.

?.

Foldalen, Hedemarken,

6.

Gausdal, Kristiansamt.

6, 9, 2.

Sundal, Bergenhus.

6.

Konnerud, Buskerud.

2 6.

Aas, Akershus.

6.

Aker, Akershus.

1 6,2 9,

Aker, Akershus.

2 juv.

9.

Christiania Fjord.

6.

Holmestrand, Jarles-

berg.

2.

Kristiansand, Maudal.

906 RODENTIA

2 6,2 9. Near Christiania,. W. E. de Winton (c & p). 97. 4. 11. 1-4.

6 juv., 9. Upland, Sweden. Lord Lilford (p). 11. 1. 1. 48-49.
(G. Kolthoff.)

2 S. Upsala. (G. KoUliujJ.) Lord Lilford (i'). 11. 1. 1. 50-51.

SciURUS VULGARIS VARIIIS Giiielin.

1789. [Sciurus vulgaris] fi varius Gmelin, Syst. Nat., i, 13tli ed., p. 146.

1792. Sc[i2i7-us'] vulglaris'] varius Kerr, Anim. Kingd., p. 250.

1829. [Sciur^is vulgaris] /3 cincreiis Fischer, Synopsis Mammalium, p. 353.

Not Sciurus cinereus Linnseus.
1857. Sciurus vtilgaris Blasius, Saugethiere Deutscblands, p. 272 (part).
1899. Sciurus vulgaris varius Barrett-Hamilton, Proc. Zool. Soc,

London, p. 6.
1906. Sciurus vulgaris varius Trouessart, Bull. Mus. d'Hist. Nat. Paris,

XII, p. .300.
1910. Sciurus vulgaris variiis Trouessart, Faune Mamm. d'Europe, p. 117.

Type locality. — Northern Europe.

Geor/rajjhiral distribution. — From the extreme north of Scan-
dinavia eastward across Finland and into Rnssia.

DiaijnosiH. — Similar to Sciiirus vulgaris vulgaris but with grey
of winter pelage much lighter, closely approaching the j^earl-
grey of Ridgway, and median dorsal region scarcely if at all
washed with chestnut.

Colour. — Winter pelage : hairs of upper parts slate grey at
base {10 mm.), the underfur tipped (4 mm.) with a pale grey
most nearly approaching the grey No. 10 of Eidgway, but less
blue and with a slight smoky cast, the longer hairs (30 mm.)
greyish to level of tips of underfur, then light hair-brown with
from one to three whitish annulations. The general effect is a
delicately grizzled, light bluish grey, approaching the pearl-grey
of Ridgway, the sides lighter and clearer ; the back with a
slight drab cast. In some specimens the drab becomes faintly
reddish. Grown and face not so pale as back, occasionally with
a distinct brownish or reddish wash. Cheeks like sides of body ;
muzzle buffy whitish. Ears grizzled grey like sides of body
internally, the tufts varying from orange-rufous to blackish.
Outer surface of legs like sides of body, usually tinged to a vary-
ing degree with dull orange-rufous or light russet, this generally
the dominant colour on upper side of feet ; soles light buffy gre}'.
Tail greyish or light reddish russet above, clouded by the blackish
sub-terminal bands (20 mm.) of hairs ; tips of hairs incon-
spicuously light russet. Underside of tail with median area
(25 mm.) like sides of body but more coarsely grizzled, this
edged with yellowish russet (10 mm.) and blackish. Underparts
and inner surface of legs creamy white to base of hairs ; chin
sometimes a little grizzled with grey.

Measurements. — Adult male from Kirkkonunnni, Finland :
head and body, 205; tail, 160; hind foot, 55; ear, 30. For
cranial measurements see Table, p. 918.

sciuRus 907

Specimena pxamined. — Eight, from the following localities : —
Norway: Hammcrfest, Fiiimarken, 2; Alteu, Finmarkeu, 1.
Sweden : Enontekis, Norrbotten, 1.
Finland: Muonionniska, 3 ; Kirkkonummi, 1.

2. Hammerfest, Finniarkcn, G. Barrett-Hamilton 11. 1. 2. 152-153.

Norway. (J. L. Bonhotf.) (p).
9. Alten, Finmarkcn. Christiania Musemn 93. 3. 1. 9.

6. Enontekis, Norrljotten, D. IMeinertzhagcn 97. 10. 21. 5.

Sweden. (c & p).

4. Muonionniska, Finland. D. Meinertzhagen 97. 10. 21. 1-4.

(C (fe p).
6. Kirkkonummi, Finland. Lord Lilford (i'). 98. 11. 4. 2.

(P. Mcrilaincn.)

SCIURUS VULGARIS LEUCOURUS KciT.

1792. Sc\i7irufi] vnlg[ciris] lencourus Kerr, Anim. Kingd., p. 25G.

1857. Sciurus vulgaris Blasius, Silugethiere Deutschlands, p. 272 (part).

189G. Scmrua vulgaris leucurus Thomas, The Zoologist, 3d ser., xx,

p. 402, November, 1896.
1899. Sciuvjcs leucurus Barrett-Hamilton, Proc. Zool. Soc, London, p. 3.
190G. Sciurus vulgaris leucurus Trouessart, Bull. IMus. d'Hist. Nat., Paris,

XII, p. 3G'2.
1910. Sciurus vulgaris leucurus Trouessart, Faune ]\Iamm. d'Europe,

p. 118.

Type locaUti/. — England.

Geo(jrnpliical distribution. — Great Britain and Ireland.

Diagnosis. — Dark phase absent : tail drab, fading in summer
to cream-buff, never tinged with red ; summer pelage : general
colour above light hazel or dull rufous, clearest and brightest on
sides and legs ; winter pelage : general colour above a dull
hroccoli-bi'own. Size small, the condylobasal length of skull
I'arely if ever exceeding 48 mm.

dolour. — Summer pelage : upper parts a reddish brown ^ iirN'ing
from a light hazel to a dull rufous, darker and less reddish along
median Jirea from face to rump, brighter on sides, on outer
surface of limbs, and on feet ; underparts dull white, usuall\'
marked off from brown of sides by a narrow line of tawny-
ochraceous, this spreading to suffuse inner svirface of limbs ; tail
a rather dark drab bleaching irregularly to cream-buff; a similar
bleaching takes place in the remnants of the ear-tufts which
occasionally persist. Winter pelage : upperparts an incon-
sjiicuously grizzled greyish brown approaching the broccoli-
bi'own of Ridgway ; the middle of back faintly darker, tlu^
sliouklcrs and sides of neck usuallv tinged with russet ; outer
surface of limbs and narrow line along sides russet or hazel,
rarely with the; decided I'eddish tinge seen in summer ; under-
parts dvdl white ; tail and ear-tufts dral), occasionally with a
dusky cast.

Measure )ne)itx. — Adult male and female from Dunphail, Elgin,
Scotland: head and body, 218 and ^l\'.^ ; tail, 179 aiul 165;

908

RODENTIA

hind foot, 58 and 50. Average and extremes of ten adults from
Saffron Walden, Essex: head and body, 227-5 (220-238); tail,
171-9 (159-178); hind foot, 56-0 (55-58). Average and
extremes of ten adults from Blandford, Dorset : head and body,
216-4(211-224); tail, 172-1 (162-190); hind foot, 57-0 (55-
59); ear, 27*7 (27-29). For cranial measurements see Table,
p. 919.

Siiccimens examined. — One hundred and seventy-four, from the follow-
ing localities : —

Ireland : Craigie, Kilmanock, 1 ; Ahascragh, Galway, 1 ; Woodpark,
Clare, 1 ; Graigue, Kilkenny, 1.

Scotland: Black Isle, Cromarty, 1 ; Dunphail, Elgin, 3; Ballends-
loch, Banffshire, 2 ; Loch Earn Head, Perthshire, 3 ; Pitlochry, Perth-
shire, 2 : Stockbriggs, Lanarkshire, 3 ; Tillicoultry, Clackmannanshire, 3.

England : Guishorough, Yorkshire, 2 ; Leeds, Yorkshire, 2 (U.S.N.M.) ;
Oswaldskirk, Yorkshire, 1 ; Greta Bridge, Yorkshire, 1 ; Cowfold, Horsham,
Norfolk, 1 ; Bury St. Edmunds, Suffolk, 3 ; Barrow, Suffolk, 3 (U.S.N.M.) ;
Cambridge, 1 ; Graftonbury, Herefordshire, 4 ; Dudley, Worcestershire, 1 ;
Middlehill, Broadway, Worcestershire, 4 ; Chipping, Sodbury, Gloucester-
shire, 1 ; Newland, Coleford, Gloucestershire, 1 ; Clarberton Road, Pem-
brokeshire, 1 ; Glamorganshire, 1 ; ]Monmouthshire, 1 ; Clevedon,
Somersetshire, 1 ; W^ormsley, Oxfordshire, 5 ; Thornhaugh, Northampton-
shire, 4 ; Luton, Bedfordshire, 6 ; Felden, Hertfordshire, 3 ; Tring, Hert-
fordshire, 1 ; Saffron Walden, Essex, 40 ; Whitley, Surrey, 1 (U.S.N.M.) ;
Milford, Surrey, 2 (U.S.N.M.) ; Horsham, Surrey, 1 ; Knockholt, Kent, 5 ;
Upton, near Andover, Hampshire, 1 ; Blandford, Dorsetshire, 55.

Memarl^s. — By its small size and the peculiar colovir of its
tail the British squirrel is one of the most sharply differentiated
races of Sciurus vulgaris. Nothing like the peculiar summer
whitening of the tail occurs in the other forms ; and this
character is the more remarkable for its occurrence in an animal
subjected to climatic conditions the reverse of those which would
be expected t(5 produce bleaching. Even when not bleached the
tail is of a drab hue unlike that in any of the Continental forms.
It would probably be more logical to regard the animal as a
distinct species, but for conxenience it is here treated as a race.

9. Ahascragh, Galway, Ire- G. Barrett-Hamilton 11. 1. 2. 154.

land. (W.S.Witherby.) (p).

6. Woodpark, Clare. G. Barrett-Hamilton 11. 1. 2. 155.

{R. F. Hibberf.) (p).

9. Graigne, Kilkenny. G. IBarrett-Hamilton 11. 1. 2. 15G.

(I/. I. Finn.) (p).

?. Black Isle, Cromarty, W! R. Ogilvie-Grant 11. 1. 3. 422.

Scotland. (c & p).

9. Dunphail, Elgin. W\ R. Ogilvie-Grant 11. 1. 3. 423.

(c & p).

3 o'. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 2G-28.

c^, 2 9. Tillicoultry, Clackman- R. G. Wardlaw Ram- 11. 1. 3. 424-42G.
nan shire. say (c & p).

1. Glamorganshire, Wales. Rev. A. Morgan (c & p). 71.7.9.1.

1. Monmouthshire. Rev. A. Morgan (c & p). 73.7.12.1.

6. Oswaldskirk, Yorkshire, A. Houston Boswoll 11. 1. 3. 427.

England. (c & p).

9. Cruisborough, Yorkshire. A. E. Pease (c & p). 11. 1. 3. 428.

sciuRus 909

J, 29. Thornhaugh, Northanap- Rev. H. IT. Slater 11.1.3.429-431.

tonshire. (c & p).

2 6. Bury St. Edmunds, Suf- Dr.A.Gimthcr (c&r). 83.2.3.1-2.
folk.

6. Horsham, Surrey. E. R. Alston (i). 7'J. 9. 25. 29.

{J. E. Hurting.)

55. Blandford, Dorset. J. C. Manscl I'lcydcU 97. 1. G. 1-55.

(c & i>).

SCIUKUS VULCiAKIS liUSSUS Millor.

1857. Sciurus vulgaris Blasius, Siiugethiere Deutschlauds, p. 272 (part).
1899. Sciartis vulgaris rufns Barrett-Hamilton, Proc. Zool. Soc, London,
p. 5 (part). Not of Kerr, 1792.

1906. ISciiirus vulgaris rufus Trouessart, Bull. Mus. d'Hist. Nat. Paris,

XII, p. 360 (part). Not of Kerr, 1792.

1907. Sciurus vulgaris russus IMiller, Ann. and Mag. Nat. Hist., 7tli ser.,

XX, p. 427, November, 1907. Type in British Museum.
1910. Sciurus vulgaris russus Trouessart, Faune Mamm. d'Europe, p. 119.

Ti/jtc lucalitij. — Dinaii, Cutes-du-Nord, France.

GeogrcqjMcal distribution. — West-central continental Europe
from Brittany to Holland, east to central France and perhaps
further ; details of distribution not known.

Diagnosis. — Dark phase present but less common than red
phase. Summer pelage : red phase : essentially as in *S'. viihjaris
vuhjaris, but red of back more strongly tinged with chestnut •
dark phase : body very dark, almost blackish mummy-brown,
the sides and shoulders washed with hazel ; tail blackisli.
AVinter pelage : light phase : body hazel, the sides a little
suffused with light smoke-grey, tail rufous, much darker than
that of Ridgway ; dark phase not seen.

Mcasuroaenta. — Type (adult male) : head and body, 202 ;
tail, 166; hind foot, 54; ear, 30. A second adult male from
the type locality : head and body, 208 ; tail, 173 ; hind foot,
54 ; ear, 28. Adult male and female from Oosterbeek, Guelder-
land, Holland: head and body, 217 and 227; tail, 175 and
172; hind foot, 57*5 and 57 "5; ear, 31 and 32. For cranial
measurements see Table, p. 919.

Specimens examined. — Twenty-six, from the following localities : —
Holland : s'Graveland, 8 ; Oosterbeek, Guelderland, 2.
France : Seine-Inferieure, 5 ; Dinan, C6tes-du-Nord, 4 ; Meurthe-et-
Moselle, 7 (perhaps referable to S. v. infuscatiis).

2 tJ, 4 ?. s'Graveland, Hilversen, F. E. Blaauw (c & p). 98. 5. 17. 1-2.

Holland. 98. G. 11. 1-2.

98. 11. 13. 1-2.

6, 9. Oosterbeek, Guelder- 0. Thomas (c & p). 98. 2. 1. 13-14.
land.

6. Duclair, Seine - Inferi- G. Barrett-Hamilton 11.1.2.157.
eure, France. (p).

4 6. Dinan, Brittany. W. J. Bramley (c & p). 97. 11. G. 1-4.

(97. 11. 6. 2. Tijpe of subspecies.)

5 9. Manonville, Meurthe- Lord Lilford (r). 11. 1. 1. 97-101.

et-Moselle. (Lomont.)

910 KODENTIA

SCIURUS VULGARIS FUSCOATKK Altum.

1804. Sciurus vulgaris var. cinerea Hermann, Obscrv. Zool., p. 65

(Germany?) Not Sciurus cinereus Linnaeus 1766.
1857. Sciurus vulgaris Blasius Siiugethiere Deutschlands, p. 272 (part).
1876. [Scittrtis vulgaris] var. fuscoatra Altum, Forstzoologie, 2d cd., i,

p. 75 (Harz Mts. and Silesia).
1876. [Sciurus vulgaris] var. nigrescens Altum, Forstzoologie, 2d cd., i,

p. 75 (Silesia).
1876. [Sciurus vulgaris] var. brunnca Altum, Forstzoologie, 2d ed., i,

p. 75 (Alsace-Lorraine).
1876. [Sciurus vulgaris] gnuca Altum, Forstzoologie, 2d ed., i, p. 75

(synonym of brunnca).
1876. [Sciurus vulgaris] al/nna Altum, Forstzoologie, 2d ed., i, p. 75

(synonym of brunnca). Not Sciurus alpinus F. Cuvier, 1842.
1809. Sciurus vulgaris rufus Barrett-Hamilton, Proc. Zool. Soc. London,

p. 5. Not of Kerr, 1792.
1905. Sciurus vulgaris var. gottharcli Fatio, Arch. Sci. Phys. et Nat.,

Geneve, 4.th ser., xix, p. 512 (South slope of Mt. St. Gothard,

Switzerland).

1907. Sciurus vulgaris rutilans ]\Iiller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 426, November, 1907 (Rudolstadt, Thiiringen, Germany).
Type in British Museum.

1908. Sciurus vulgaris fuscuatcr Miller, Ann. and ]Mag. Nat. Hist., 8th ser.,

I, p. 128, January, 1908.
1910. Sciurus vulgaris fuscoatcr Trouessart, Faune Mamm. d'Europe,
p. 118.

Type locality. — Harz Mountains, Geiinany.

G coijraphical (1 istrlhution. — East-central Europe from Germany
through Austria-Hungary to Roumania; south through the Alps.

l)ia<inoxif<. — Similar to *S'. cuhjaris runtmt^, but red phase in
])otli winter and summer a clear bright rufous, tinged with grey
on sides and sometimes on back. Dark phase present but less
common than red phase, the relative abundance of the two
subject to much local variation. Sunnner pelage : light phase :
body clear rufous, closely agreeing with that of Ridgway, and
without chestnut suffusion, tail the same clear rufous but more
intense ; dark phase as in winter but general effect more sooty.
Winter pelage : light phase : body rufous tinged with light
smoke- grey along sides, tail clear rufous; dark phase: body a
grizzled hair-brown suffused with mummy-brown over back ; tail
slaty black.

Measurements. — Average and extremes of five adults from
Ingelheim, Rheinhessen, Germany: head and body, 229 '4 (220-
236); tail, 192 (189-198); hind foot, 60-4 (59-61); ear, 32-4
(31-34). Young adult male from Rudolstadt, Thiiringen,
Germany (type of ri(tilans Miller) : head and body, 223 ; tail,
175 ; hind foot, 57 ; ear, 27. Adult female from Niesky, Silesia,
Germany: head and body, 228; tail, 195; hind foot, 62; ear,
32. Adult male and female from St. Gallen, Switzerland : head
and body, 230 and 229; tail, 184 and 190; hind foot, 61 and

SCIURUS 911

59. Adult male from Hatszeg, Hunyad, Hungary : head and
body, 220 ; tail, 190 ; liind foot, 59. Average and extremes of
live adults from Bustenari, Roumania : head and body, 220 "8
(213-225); tail, 181-2 (170-185); hind foot, 58-6 (57-60);
ear, 30 • 9 (29-32 • 5). For cranial measurements see Table, p. 920.

Sj'cciinens examined. — One hundred and seventy, from the following
localities : —

France: Etupes, Doubs, 1 (Mottaz) ; Montauban, Haute-Savoie, 2;
Les Pitous, Haute-Savoie, 11 (Mottaz) ; Scientriers, Haute-Savoic, 2
(Mottaz) ; Barcelonnette, Basses-Alpos, 5 (B.M. and Mottaz) ; Dignc,
Basses- Alpes, 3 (Mottaz).

Germany : Marburg, Hcssen-Nassau, 4 ; Blumenthal, Hanover, 2 ;
Buchholz, Saxony, 1 (U.S.N.M.) ; Moritzburg, Saxony, 2 (U.S.N.M.) ;
Wernigerode, Saxony, 1 ; Neustadt, Saxony, 2 ; Ilscnburg, U ; jMagdeburg,
1 ; Ammenslablen, 1 ; Rudolstadt, Thiiringcn, 5 ; Ummerstadt, Thiiringen,
12 ; Ingelheim, 10; Hartwald, Baden, 1 (U.S.N.M.) ; Nuremberg, Bavaria,

1 (U.S.N.M.) ; Strass, near Burgheim, Bavaria, 9 ; ]\Iarxheim, near
Monheim, Bavaria, 6 ; Niesky, Silesia, 1.

Austria-Hungary: Csallokoz-Somorja, Pressburg, 2; Trcfail, Steior-
mark, 1 ; Haida, Arva, Bohemia, G ; Karlsbad, Bohemia, 1 ; Hatszeg,
Hunyad, Bohemia, 1 ; Hungary, no exact locality, 1 (U.S.N.M.) ;
Zuberec Hungary, 2.

Roumania : Bustenari, Prahova, 5.

Switzerland : Geneva, 2 (Mottaz) ; St. Cergues, Vaud, 1 (U.S.N.M.)
Morat, Fribourg, 1 (Mottaz) ; St. Gallon, 5 (B.M. and U.S.N.M.) ; Gais
Appenzell, 1 (U.S.N.M.) ; Hcrisau, Apponzell, 1 (U.S.N.M.) ; Untervatz
Grisons, 3 (B.M. and U.S.N.M.) ; Rabins, Grisous, 1 (U.S.N.M.) ; Albogasio
Ticino, 1 (U.S.N.M.); Curregia, Ticino, 1 (U.S.N.M.); Davesco, near
Lugano, Ticino, 2 ; Lugano, Ticino, G (Mottaz) ; Lumino, Ticino, 1
(U.S.N.M.) ; no exact locality, 1.

Italy: Padola, Cadore, 4 (Turin); Crevacuoro, Novara, 3 (Turin);
Coggiola, Novara, 7 (Turin) ; Val d'Aosta, 3 (Turin) ; near Turin, 2 (Turin) ;
Sommariva del Bosco, Cuneo, 2 (Turin) ; Dronero, Cuneo, 1.

1. Montauban, Haute- A. Robert (c it r). 97.1.9.4.

Savoie, France.
6. Montauban, Haute- O. Thomas (i'). G. 4. 2. 9.

Savoie. (.1. Robert.)
6, 2 9. Marburg, Hesseu-Nassau, Lord Lilford (r). 11. 1. 1. 76-78.

Germany. (Sclmeider.)

5. Blumenthal, Hanover. Lord Lilford (p). 11. 1. 1. 87.

( Woltcrsturff.)
9. Wernigerode, Saxony. Lord Lilford (p). 11. 1. 1. 88

(Woltcrstorff.)
6, V. Noustadt, Saxony. " Lord Lilford (p). 11. 1. 1. 83-84.

(Sclmeider.)
6, 9 9. Ilsenburg, Saxony. Lord Lilford (p). 11. 1. 1. 52-60

(Bartells.)
1. Gross Ammenslablen, Lord Lilford (p). 11. 1. 1. 91.

Saxony. (Schucltardt.)

6. Magdeburg, Saxony. Lord Lilford (p). 7. 9. 7. 1.

5 6, 9. Ummerstadt, Thiiringen. Lord Lilford (p). 11. 1. 1. 70-75.

(Schucltardt.)

2 6,3 9. Rudolstadt, Schwartz- Lord Lilford (p). 95. 4. 18. 7-11.

burg, Thiiringen. (95. 4. 18. 7. Type of S. v. rutilans Miller.)

6, 5 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 32-37.

3 6, 9. Ingelheim, Rheinhessen. Lord Lilford (p). 11. 1. 1. 79-82

(Hilrjcrt.)

9, 1. Burgheim, Bavaria. Lord Lilford (p). 11. 1. 1. 85-86.

{Kurbitz.)

912

RODENTIA

9. Strass, Bavaria. Lord Lilford (r). 11. 1. 1. 89.

6, 2 ?. Marxheim, Bavaria. Lord Lilford (p). 11. 1. 1. 67-69.

{Wolterstorg)

?. Niesky, Silesia. ' Dr. E. Hamilton (r). 97. 12. 4. 20.

2. Csallokoz-Somor j a, Budapest Museum (e). 94.8.1.29.

Pressburg, Hungary.
6. Tr6fail,Steiermark,Hun- Lord Lilford (p). 11. 1. 1. 90.

gary.
6 i. Haida, Bohemia. Lord Lilford (p). 11. 1. 1. 61-66.

{Woltcrstorff.)

1. Karlsbad, Bohemia. Lord Odo llusscll 44. 9. 7. 3.

(c & p).
i. Hatszeg, Hunyad, Tran- C. G. Dauford (c). 3. 11. 8. 20.

sylvania.

2. Zuberec. BudapestMuscum(K). 94. 3. 1. 31-32.
3 J, 2 9. Bustenari, Prahova. Rou- Lord Lilford (p). 4. 4. 6. 67-71.

mania. {W. Dodson.)
2 (5, 2 9. St. Galleu, Switzerland. 0. Thomas (p). 4. 4. 5. 35-38.

(Zollikofer.)
2 9. Uutervatz, Grisons. O. Thomas (p). 4. 4. 5. 39-40.

{Zollikofer.)
6, 6 juv. Davcsco, Ticino. O. Thomas (p). 2. 8. 4. 28-29.

{Zollikofer.)
6. Switzerland. E. R. Alston (p). 79. 9. 25. 30.

SciUEUS vuLUARis iTALicus Bonaparte,

1838. Sciurus italicns Bonaparte, Iconogr. Faun. Ital., i, fasc. 23 (Italy).
1857. Sciurus vulgaris Blasius, Saugethiere Deutschlands, p. 272 (part).

1906. Sciurus vulgaris italicns Trouessart, Bull. Mus. d'Hist. Nat. Paris,

XII, p. 364 (part).

1907. Sciiirus mcridionalis Lucifcro, Revista Ital. di Sci. Nat., Siena,

XXVI, p. 45, June, 1907 (Sila, Calabria, Italy). Renaming of
S. alpinus Costa (not of Desmarest).
1910. Sciurus vulgaris italicns Trouessart, Faune Mamm, d'Europc,
p. 122.

Tijjpe Ivcalitj/. — Italy.

Geographical distrihution. — Italy except the region north of
the Apennines.

Diagnosis. — Red phase lighter and more ochraceous than in
Sciurus vulgaris fuscoater ; brown phase less dark than in
fuscoater, the tail with broad grizzled median area ; back rarely
if ever showing any evident trace of n darker median line, and
sides rarely with any noticeable greyisli suffusion even in winter.

Colour. — Red phase, winter : upper parts clear ochraceous-
rufous, the crown, ears and median dorsal region brighter and
more red, the sides paler and duller but with no distinct inter-
mixture of grizzled grey ; tail dark orange-rufous, the median
region below tinged with light clay-colour or occasionally some-
what grizzled ; feet like median region of back, the soles drab ;
face and muzzle, especially the region surrounding base of
whiskers, fading to ochraceous-buff; underparts from throat to
base of tail buffy white ; interramia tinged vrith. grey ; inner
side of legs paler and less red than outer side, closely resembling

sciURUs 913

median region of tail. Brown phase, winter : back and sides
a uniform, rather fine grizzle of wood-brown, cream-butt' and
black, in which the brown predominates, and in which there is
f retpiently a decided tinge of drab ; ear tufts dark brown,
approaching black, some of the hairs usually with indistinct
light annulations ; tail blackish brown above, the underside with
a broad median band essentially like back except that it is much
more coarsely grizzled ; feet varying from dull russet to blackish,
the soles usually hair-brown or drab ; muzzle dull ochraceous-
bufF ; underparts as in red phase except that interramial region
is darker and often with a slaty tinge. Summer pelage not seen.
Measurements. — Average and extremes of five adults from
western Liguria, Italy: head and body, 230 (218-242); tail.
188-2 (176-197); hind foot, 58 (.57-60); ear, 29-4 (27-31).
Average and extremes of four adults from the province of Rome,
Italy : head and body, 237-7 (220-250) ; tail, 196-2 (192-200) ;
hind foot, 58-5 (57-60). For cranial measurements see Table,
p. 921.

Specimens examined. — Thirty-eight, from the following localities in
Italy : neighbourhood of Genoa, 28 (Genoa) ; Spezia, 1 (Turin) ; Quiesa,
Lucca, 1 ; San Casciano, Florence, 2 ; Siena, 2 ; Arsoli, Rome, 2 ; Viterbo,
Rome, 2.

6. Quiesa, Lucca, Italy. Dr. E. Hamilton (p). 98. 10. 2. 9.

6. San Casciano, Florence. Dr. E. Hamilton (p). 98. 10. 2. 10.

2. Siena. (Broqi.) Paris Museum (e). 6. 7. 15. 1-2.

?. Arsoli, Rome. {Coli.) G. Barrett-Hamilton 11. 1. 2. 36.

(P)-
6. Viterbo, Rome. iColi.) G. Barrett-Hamilton 11. 1. 2. 35.

SciURUS VULGARIS LiL^us Miller.

1906. Sciurtis vidgaris italicus Trouessart, Bull. ]\Ius. d'Hist. Nat. Paris,

xii, 13. 364 (part).

1907. Sciurus vulgaris lihnis Miller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 429, November, 1907. Type in British Museum.
1910. Sciurus vidgaris lilmus Ttouessart, Faune Mamm. d'Europe, p. 123.

Type locality. — Agoriani, north side of Lyakura (Parnas.sus)
^Mountains, Greece.

Geograjjldcal distrihutton. — Greece ; limits of range not known.

Diagnosis. — Size as in S. vulgaris italicus ; colour in brown
phase peculiar in the noticeable contrast of the very dark,
almost blackish, posterior half of l)ack with the hair-brown
shoulders and neck ; red phase not known and probably rai-e oi-
absent.

Colour. — General colour above a grizzled hair-brown, paler
and more grey on cheeks and across muzzle, much darkened with
blackish on posterior half of back and on outer surface of hind
legs. Inner surface of hind legs and line along sides of body
bordering white of underparts tawny-ochraceous, becoming

3 X

914 RODEXTIA

lighter and duller anteriorly and continuing over outer surface
of fore leg and along side of neck. Feet dull tawny-ochraceous
clouded with blackish. Tail blackish suffused with tawny-
ochraceous beneath surface. Underparts creamy white ; chin
and interramial region light drabby grey.

Measuremenis. — Type (young adult female) : hind foot, GO ;
ear from meastus, 29. For cranial measurements see Table,
p. 921.

Specimens examined. — Three, all from the type localitj".

1. Agoriani, Lyakura Mts., Greece. Lord Lilford (p). 7. 9. 8. 1.

{Schliiter.) {Tijpe of subspecies.)

1. Parnassus. Lord Liiford (p). 8. 10. 17. 1.

SciURUS VULGARIS ALPiNus Desmarest.

1822. Scmrus alpinus Desmarest, Mammalogie, ii, p. 543 (Pyrenees).
1842. Sciurus alpinus P. Cuvier, Hist. Nat. des Mamm., iv, tabl. gen., p. 4

(described and figured in fasc. 24, pi. 237, 1821).
1857. Sciums vulgaris Blasius, Siiugethiere Deutschlands, p. 272 (part).
1905. Sciurus alpinus Cabrera, Bol. Real Soc. Espafi. Hist. Nat., v, p. 230,

April, 1905.
190G. Sciurus vulgaris alpinus Trouessart, Bull. ilus. d'Hist. Nat., Paris,

XII, p. 363.
1910. Sciurus vulgaris alpinus Trouessart. Faune Mamm. d'Europe, p. 119.

jT/ype localitj/. — Pyrenees.

Geographical distribution. — Pyrenees ; limits of range not
known.

Diagnosis. — Size essentially as in Sciurus vulgaris italicus ;
colour in dark phase similar to that of S. v. riissns and S. v.
fascoater ; skull with rostral portion noticeably broadened and
shortened.

Measurements. — -Adult male from Espot, Lerida, Spain : head
and body, 240 ; tail, 205 ; hind foot, 58 ; ear, 22. For cranial
measurements, see Table, p. 921.

Specimens examined. — Two from Espot, Lerida, Spain (U.S.N.M. and
Cabrera).

MemarTis. — The status of this race is not clearly understood.

Sciurus vulgaris numaxtius Miller.

1905. S^ciurusl sp. Cabrera, Bol. Real Soc. Espafi. Hist. Nat., Madrid, iv,

pp. 224, 231, April, 1905.
1905. S\ciurxis'] rufus Cabrera, Bol. Real Soc. Espau. Hist. Nat., ]\Iadrid,

IV, p. 225, April, 1905 (not of Kerr, 1792).
1907. Sciurus vulgaris nurnantius ]\Iiller, Ann. and l\Iag. Nat. Hist., 7th

ser., XX, p. 428, November, 1907. Type in British ]\Iuseum.
1910. Sciurus vulgaris nurnantius Trouessart, Faune Mamm. d'Europe,

p. 120.

Type locality. — Pinaies de Quintanar de la Sierra, Burgos,
Spain.

SCIURUS

915

Geogrfqjhical dislrlhution. — Northern and north-central Spain
(provinces of Leon, Vitoria, Huesca and Burgos) ; limits of range
not known.

Diagnosis. — Size greater than in the central European forms,
nearly equal to that of S. v. infuscatus ; colour in light phase not
so dark as in S. v. infuscatus, and tail never with whitish median
area on lower surface.

Colour. — Light phase (type) : head, back, sides, and outer
surface of legs a uniform indistinctly grizzled bi-own, intermediate
between the broccoli-brown and wood-brown of Ridgway ; a faint
russet tinge along middle of back. Muzzle and fore part of face
between ochraceous-buffand clay-colour. Cheeks drab. Ear-tufts
l)lackish brown, sides of neck pale dull wood-brown. Inner
surface of legs and ill-defined stripe along sides of belly dull light
hazel. Feet like inner side of legs, but paler. Tail a very dark
rufous, approaching the chestnut of Ridgway, especially near
base, but rather more red ; median portion of tail below lighter,
the hairs bufFy grey through basal half, each with two drab
anuulations. Underparts buffy white, the chin and interramial
region light ecru-drab. Dark phase : tail clear bluish black, very
faintly grizzled along median region below ; back much darkened
by a blackish suffusion. Colour variation shows itself chiefly in a
greater or less tendency to assume the dark phase. Occasionally
the hazel of sides brightens almost to a dull rufous and spreads
to lateral portion of dorsal area, the region immediately bordering
white of ventral surface becoming nearly buff. Ear-tufts either
blackish or reddish.

Skull and teeth. — The skull and teeth are larger than those of
the central European races, but in form they show no tangible
peculiarities.

Measurements. — Type (adult female) : head and body, 2.37 ;
tail, 200 ; * hind foot, 66 ; ear, 34. Two other adult females
from the type locality: head and body, 235 and 250; tail, 190
and 210; hind foot, 63 and 64; ear, 34 and 34. Adult male
from Panticosa, Huesca : head and body, 262 ; tail, 242 ; hind
foot, 64 ; ear, 36. For cranial measurements see Table, p. 922.

Specimens examined. — Twenty-two, from the following localities : —
France : Solf^rino, Landes, 1 ; St. Jean de Luz, Basses-Pyrenees, 1.

(These specimens intermediate between numantius and russus.)

Spain : Sierra de Dubros, Asturias, 2 ; Pajares, Leon, 1 ; Arrechavaleta,

Vitoria, 2 ; Panticosa, Huesca, 6 ; Palacios de la Sierra, Burgos, 2 (U.S.N. M.) ;

Pinares de Quintanar, Burgos, 7.

9. Solf(irino, Landes, 0. Thomas (p). G. 4. 1. .^

France. {A. Robert.)

6. St. Jean de Luz, Basses- W. Eagle Clarke (p). 92. 10. 30. 1.

Pyr6n6es.

6, 9. Sierra de Dubros, Astu- J. W. Richmond Lee 97. 1. 7. 1 2.

rias, Spain. (c & p).

* Misprinted 230 in original description.

3 N 2

916 RODENTIA

?. Pajares, Leon. 0. Thomas (p). 8. 2. 9. 61.

{N. Gonzalez.)

2 6. Arrechavaleta, Vitoria. O. Thomas (p). 8. 2. 9. 65-66.

{N. Gonzalez.)

2 6, 9. Panticosa, Huesca. O. Thomas (p). 8. 2. 9. 62-64.

(N. Gonzalez.)

5 ?. Pinares de Quintanar, G. S. Miller (c). 8. 8. 4. 52-56.

Burgos. (8. 8. 4. 52. Type of suhspecies,)

SciURUS VULGARIS INFUSCATUS Cabrera,

1905. Sciurus infuscatus Cabrera, Bol. Real Soc. Espail. Hist. Nat., v,
p. 227, April, 1907. Type in U.S. National Museum.

1909 Sciurus vulgaris infuscatus Miller, Ann. and ]\Iag. Nat. Hist., 8th
ser., Ill, p. 418, May, 1908.

1910. Sciurus vulgaris infuscatus Trouessart, Faune Mamm. d'Europe,
p. 121.

Ti/pe locality. — -Las Navas, Avila, Spain.

Geographical distrihniion. — Central Sjaain.

Diarjnosis. — Largest of the European squirrels (condylobasal
length of skull, 52 • 4 to 55 mm.) ; colour very dark, the two
phases if present not well defined ; tail rusty red, its under
surface conspicuously sprinkled with pure white hairs ; cheeks
and throat strongly contrasted in colour.

Colour. — Upper parts a rich, faintly grizzled dark brown, the
general efiect nearly the mummy-brown of Ridgway, the posterior
half of back darker and less grizzled than shoulders and neck ;
underfur slate-grey, the tips of the hairs russet ; face rather
abruptly russet, the muzzle tinged with bufi"; cheeks below and
behind eyes an indefinite drab brown, noticeably contrasted with
svu-rounding parts, especially with white of throat ; ears russet,
the longer hairs dusky ; feet rufous, somewhat darker than that
of Ridgway, this colour sufi'using outer surface of fore leg and
extending up inner surface of hind leg to thigh, though in both
instances duller and more brownish on leg than on foot ; under-
parts buffy white, the extreme bases of hairs inconspicuously
gi-eyish, the white covering entire inner side of fore leg to wrist,
and extending as a narrow line down inner side of hind leg to
lieel and sole ; tail a rich dark red, between the rufous and
ferruginous of Ridgway, the distal half usually with a slight
dusky cast, the entire tail sprinkled with pure white hairs, this
inconspicuous above but tending to form a whitish, often very
noticeable median area below.

,S7i«// and teeth. — Except for their lai'ger size the skull and
teeth do not difter appreciably from those of the central European
forms.

Measurements. — Type (adult male) : head and bod}-, 280 ;
tail, 230 ; hind foot, 65 ; ear, 33. Adult female from La Granja,
Segovia : head and body, 256 ; tail, 220 ; hind foot, 66 ; ear, 33.
For cranial measurements see Table, p. 922.

SCIURUS 917

Specimens examined. — Five, from the following localities iii coutral
Spain : Las Navas, Avila, 1 (U.S.N.M.) ; La Granja, Segovia, 3 (B.M. and
Cabrera) ; no exact locality, 1.

6.

Old Castile, Spain.

Lord Lilford (p).

Ui. 6. 11. G.

6.

La Granja, Segovia.

(Escalera.)

A. Cabrera (p).

7. 10. 17. 1.

Sciuiius VULGARIS SKGUK.E Miller.

I'JO'J. Sciiu'us vulgaris segur;>'. ]\Iiller, Ann. and IMag. Nat. Hist., 8th scr.,

Ill, p. 418, May, 1909. Type in British Sluseum.
1910. Sciurus vulgaris scguriv Trouessart, Faune ]\Iamm. d'Europc, p. 121.

Tifpe locaJit)/. — Moliniccs, Sierra de 8egura, Albacete, Spain.

Geotjraphical rlistrilmtion. — Known only from tlie Sierra de
Segura, provinces of Albacete and Jaen, south-eastern Spain.

Diagnosis. — Related to Sciurus vulgaris infuscatus, but back
less blackish, its underfur light grey, tail less red and with white
area on under surface less well developed, and cheeks light grey,
forming no decided contrast with white of throat.

Culour. — Summer pelage : upper parts a fine, inconspicuous
grizzle of wood-brown and blackish, the general effect resembling
the mars-brown of Ridgway, blackening on flanks, across posterior
half of back, and on postero-outer side of thighs ; in some
specimens the light element is more nearly russet and the black
is essentially absent ; underfur pale ecru-drab, the hairs with
faintly brownish tips ; ears and crown like back, but face with a
rusty wash, and muzzle and cheeks to behind base of ear light
clear ecru-drab, so pale as to form no marked contrast with white
of throat ; underfur of head, back, sides, and limbs pale ecru-
drab like that of cheeks, appearing conspicuously at surface
in specimens with abraded pelage ; feet a dull ferruginous, this
colour extending up outer side of thigh and over entire fore leg,
in both regions diluted by the ecru-drab of underfur ; entire
underparts and inner surface of legs buffy white to base of hairs ;
tail blackish, slightly tinged with dull red, the hairs becoming
ecru-drab at base ; whole tail sprinkled with pure white hairs
more numerous along median line below than elsewhere, and
usually (in eight of the eleven skins examined) forming a distinct
white median area as in S. v. infuscatus.

SliiU. and teeth. — The skull and teeth do notdifier apprecial)ly
from those of Sciurus vulgaris infuscatus.

Measureincnts. — Type (adult female) : head and body, 245 ;
tail (probaVjly injured in life), 195 ; hind foot, 61. Avei-age and
extremes of seven specimens from the Sierra de Segura : head and
body, 235-7 (230-245); tail, 210 (200-220); hind foot, 61-8
(60-64-4); ear, 25-8 (25-30). For cranial measurements see
Table, p. 922.

Specimens examined. — Eleven, all from the Sierra dc Segura.

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sciuRus 923

Rcmarlcti. — While this animal is readily distinguishable from
the squirrels of central and northern Spain its distinctness from
the imperfectly known Sciurus vulgaris bseticus is perhaps open to
doubt. In an obviously immature individual, however, the hind
foot is 60 mm. in length as opposed to 45 mm. in the type of
huiticus.

2 6,4:9 Sierra do Segura, Albacote, IM. de la Escalcra (c) 8. 9. 24. 1-6.
Spain. (8. 9. 23. 3 Tjjpc of subspecies.)

Sciurus vulgaris b.^^ticus Cabrera.

1905. Sciurus bn'ticus Cabrera, Bol. Real Soc. Esx^aii. Hist. Nat., v, p. 228,
April, 1905. Type in museum of University of Seville.

1910. Sciitriis vulgaris bivticus Cabrera, Asoc. Espau. Progr. Cien., Congr.
Zaragosa, 1908, p. 11, June, 1910.

1910. Sciurus vulgaris hivticus Trouessart, Faune Mamm. d'Europc, p. 122.

Tjipe locality. — Alanis, Province of Seville, Spain.

Geographical distribution. — At present known from the type
locality only.

Diagnosis. — In general much like Sciurus vulgaris segurse, but
size considerably less (hind foot only about 50 mm.) and white
area on underparts unusually narrow.

Bonarls. — This squirrel, of which I have seen no specimens,
appears to be similar *to Sciurus vulgaris segurm except for its
small size, the absence of white in the tail, and the narrowness
of the white ventral area. The type is an immature mounted
specimen in summer coat ; and its skull has not been examined.
The more important parts of the original description are
essentially as follows : Probably the smallest of the European
forms of the genus Sciurus ; further distinguished * by having the
tail unicolor and similar to the body. Pelage a bright reddish
chestnut above, white below. The white is not so extensive as
in S. vulgaris iufuscatus, being limited to the median portion of
the ventral surface. Sides of the face in region of eyes much
paler than the rest of the head, approaching a dirty yellowish.
Tail of the same colour as the body, uniform, without white
hairs on its lower sui'face. Dimensions (approximate) : head and
body, 200; tail, 160; hind foot without claws, 50. In a later
paper Mr. Cabrera corrects the measurements of tail and hind
foot to 135 and 45 mm. respectively, adding that, from informa-
tion which he has received he believes the size of the adult
animal to be about as in numantius. If this is true it seems not
improbable that segurse will prove to be the same as hn'ticus.

* (from infuscatus).

924

Genus CITELLUS ()k(ui

181G. Citellus Okcn, Lcbrb. dcr Ncaturgcsch., Th. in, Abth., ii, p. 824

(citelltos).
1825. Spermopliil'iis F. Cuvier, Dents dcs Mammiferes, p. IGO {ci(cllus).
1827-34. Citillus Lichtenstein, Darstell. ncuer oder wcnig bekanntcr

Siiugetb., pi. XXXI (citellus).
1857. Spcrmoj)hibis Blasius, SJiugetbiere Deutscblands, p. 275.
rj02. Citellus Allcu, Bull. Amer. Mus. Nat. Hist., xvi, p. 375, October 11,

1902.

Ti/pe i<pec}es. — Mus citellus Linn;eus.

Geographical distribution. — Northern Hemisphere from
Hungary eastward through Asia and North America to the
central United States.

Characters. — Strictly terrestrial Sciuridse of medium size and
slightly modified Sciurine aspect, the tail flattened but relatively
shorter and less bushy than in Sciurus ; cheek pouches present ;
skull more massive than in Sciurus, the brain-case much less
convex above, its depth rather more than twice that of rostrum ;
incisors sub-terete ; molars with cross-ridges and inner tubercle
relatively high, obscuring the basin-shaped form of crown, and
forming a conspicuous [J -shaped pattern in moderately worn
teeth ; anterior upper premolar well developed.

Beuiarls. — Though the limits of the genus Citellus are not at
present well understood, owing chiefly to the difliculty of com-
paring the Old World and American forms, the group may be
regarded as containing about eight Pahearctic species, two of
which occur in central Europe. Remains of extinct membei-s of
the genus are found as far west as England.

KEY TO THE EUROPEAN SPECIES OP CITELLUS.

Back without distinct spots ; skull with incisive fora-
mina normal, their length greater than diameter of
alveolus of upper incisor (Hungary in general, and
eastward) C. citellus,]}. 924.

Back with sharply defined whitish spots about 4 mm. in
diameter ; skull with incisive foramina very small,
their length about equal to diameter of alveolus of
upper incisor (Hungary cast of the Carpathians, and
eastward) C. suslica, p. 929.

CITELLUS CITELLUS Linnfeus.

1766. [Mus] citellus Linnseus, Syst. Nat. i, 12th ed., p. 80 (Austria).
1778. Mus citillus Pallas, Nov. Sp. Quadr. Glir. Ord., p. 19.
1857. S'pcrmophilus citellus Blasius, Saugethiere Deutscblands, p. 275.
1904. [Citellus'] citellus Trouessart, Catal. IMam. Viv. Foss., Suppl., p. 339.
1910. Citellus [Citellus) citellus Trouessart, Fauue Mamm. d'Europo,
p. 127.

Type locality. — Austria.

Geographical distribution. — From Silesia and Bohemia, east-

CITELLUS 925

wai'd through llussia into Asia ; eastern limits of range not
known.

Diagnosis. — Size medium (head and body, 195 to 220 mm. ;
tail, 60 to 75; hind foot, 35*4 to 38 "4; condylobasal length of
skull, 41 • 4 to 45) ; tail short, about twice as long as hind foot ;
colour of upper parts mingled bull' and dark brown so arranged
as to produce an evident effect of light mottling on a dark
ground ; skull with rather low, broad brain-case ; incisive
foramina not unusually reduced in size ; incisor teeth both above
and below exhibiting the minimum degree of compression.

External characters. — General form less slender than in
Scim-us vulgaris, the short, narrow tail extending little beyond
outstretched hind feet, the ear short and withovit tuft, the fur
thin, coarse and lying close to body. Ear low, obscurely jDointed
above, extending much less than half-way to eye when laid
forward, densely clothed with fine short hairs on both surfaces ;
muzzle pad naked, the bare area extending downward as a
narrow line across middle of upper lip. Feet shorter and more
robust than in Sciurus vulgaris, the digits less elongated, the
claws less curved, those on front feet longest ; fore foot with
third digit longest, second and fourth sub-equal and slightly
shorter, fifth extending a little beyond base of fourth, the thuml)
ludimentary but with rather evident, compressed nail ; hind
foot with third digit longest, second and fourth sub-etjual and
slightly shorter, fifth extending just beyond base of fourth, first
not quite to base of second ; pads both palmar and plantar as
in Sciurus vulgaris but relatively larger and more crowded ; palm
bare throughout, sole hairy behind tubercles. Tail cylindrical
at base, inconspicuously flattened beyond middle, where hairs
are about 15 mm. long. Mammse ; p 1 — 1 ; a 2 — 2 ; i2 — 2 = 10.

Colour. — Back and sides usually cream-buft' but occasionally
more yellowish, in some specimens approaching the bufi" yellow
of Ridgway ; sides inconspicuously " lined " with black ; entii-e
dorsal surface from nape to rump vermiculated with black, the
dark and light areas along middle of back usually well enough
defined to produce an effect of obscure light mottling, the spots
about 5 mm. in diameter ; crown and upper half of cheeks
grizzled, the light element paler than on body, the dark relatively
more evident ; an ill-defined eye-ring, whitish or yellowish accord-
ing to general colouring of body ; muzzle sometimes with a
rusty tinge ; sides of muzzle and lower half of cheeks clear
whitish or bufi'y, continuous with the similarly coloured pale
area covering throat and fore part of chest and inner side of fore
leg ; belly washed with a buff' usually somewhat more yellow
than that of sides ; feet like ground colour of back ; tail without
definite colour pattern, the upper surface grizzled, essentially
like back, somewhat darker at tip, tlie pencil with light margin ;
under surface of tail a nearly clear dull buif.

SLiill. — The skull is smaller than that of Sciurus vulgaris,

92G

RODENTIA

and its genei'al form differs notably in the ciineate outline due
to the narrowness of anterior zygomatic region, the almost
uniformly convex dorsal profile, and the slight deflection of basi-
cranial axis, so that upper third or fourth of foramen magnum
is above level of alveolar line. Dorsal profile convex from front
of nasals to lambdoid ridge, sometimes with a slight flattening
in interorbital region and a faint posterior concavity ; ventral
profile similar, though less strongly convex ; posterior truncation
of brain-case nearly vertical, but occipital condyles projecting

Fig. 188.
Citellus citellus. Nat. size.

sufficiently to be just visible from above. Brain-case very
broadly ovate when viewed from above, its greatest Avidth
equal to length, its posterior width obviously exceeding than
that at postorbital constriction ; surface less rounded than in
Sciurus vulgaris, and lyrate figure formed by parietal ridges
much narrower ; lambdoid crest essentially as in S. vid(jaris, but
constriction just in front of occipital region less evident. Occiput
low, its general form as in Sciurus vulgaris, but contrast between
its height and that of middle of brain-case not especially marked ;
protuberances on posterior surface of occiput less noticeable than

CITELLUS 927

in S. vnlgari.% and exposed portion of petrosal noticeably smaller ;
paroccipital processes rather large, their extremities reaching
beyond level of lower edge of condyle and often to that of
lower edge of bulla ; basioccipital essentially like that of Sciuriis
vulgaris in general form, but anterior portion of outer margin
raised into a conspicuous plate-like process applied to inner side
of bulla ; auditoi-y bulla essentially as in Sciurns vidgaris but more
evenly though somewhat less inflated. Interorbital region much
longer than broad, its least width less than that of postorbital
constriction, its surface nearly flat except for the curve of which
it forms a part ; edges of orbits slightly raised above the general
level, the supi'aorbital notch inconspicuous ; postorbital processes
relatively shorter than in Sciurus vulgaris, the space lying
between process and surface of brain-case narrower ; orbit
surrounded by bone through about two-thirds of its circum-
ference. Zygomata diverging gradually and evenly, nowhere
parallel, the greatest zygomatic breadth at or just in front of
glenoid level ; jugal projecting posteriorly as in Sciurus vulgaris
though less noticeably, its upper border without angle at
posterior limit of orbit ; anterior zygomatic root with conspicuous
projecting ridge extending along its entire anterior border ;
anteorbital foramen sub-circular in outlme, its position somewhat
more anterior in relation to tooth-row than in Sciurus vulgaris,
its entire lower and outer border thickened and lip-like.
Rostrum forming about one-third length of skull, its base not
noticeably broader than anterior region, the least depth behind
incisors about equal to width at same level ; nasals relatively
longer than in Sciurus vulgaris, extending about to lachrymal
level, the posterior border variable, usually truncate ; nasal
branches of premaxillaries broad but not expanded, rarely
extending behind nasals. Palate less concave both longitudinally
and laterally than in Sciurus vulgaris ; a low but evident median
ridge extending from incisive foramina to spine or blunt pro-
jection at posterior border ; incisive foramina about as large as
in S. vulgaris, but more distinct from each other, and .anterior
extremity not gradually narrowed to a fine point ; mesopteiygoid
fossa about three times as long as wide, narrowest near middle ;
ectopterygoid rather well developed, forming a wide flattened
area nearly as large as mesopterygoid space. Mandil^le less
robust than that of Sciurus vulgaris, the ramus in region near
symphysis scarcely deeper than wide ; coronoid process longer
and more curved than in S. vulgaris, its anterior border much
more convex ; articular process slender, deeply concave on outer
side ; angular process with anterior limit less well defined.

Teeth. — Upper incisor less robust than that of Sciurus
vulgaris, but course of shaft very distinctly marked on side of
rostrum from alveolus to anteorbital foramen ; cross section of
shaft nearly semicircular in outline, the flattened side turned
inward ; enamel extending from antcu'O-internal extremity nearly

928

half-way along anterior and outer cnr\'ature, its surface smooth,
yellowish white. Lower incisor with root extending slightly
beyond m.^ ; section of shaft and arrangement of enamel
essentially as in upper tooth, but outer border flattened behind
enamel plate, thus distorting the semicircular outline. Cheek-
teeth agreeing with those of Sciurus vulgaris in general plan of
enamel folding but differing in numerous details, principally the
result of a tendency toward greater height of tubercles and main
ridges. Anterior upper premolar simple, terete, its crown with
high, median, obliquely transverse ridge and small anterior
and posterior depression, the area of crown about one-half that
of succeeding tooth. Upper cheek-teeth with inner tubercle
much narrower and higher than in *S^. vulgaris, the length of
base scarcely equal to height, the width of base decidedly greater
than length ; main ridges (second and fourth) simple but high,

extending to summit of inner
tubercle and forming, together
with the tubercle, a narrowly
U-shaped figure, the most con-
spicuous feature of crown when
not worn away ; anterior ridge
very narrow, at extreme edge of
crown, separated from base of
second ridge by a conspicuous
furrow, its outer extremity form-
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rounded inner termination lying
on anterior surface of base of
main tubercle ; third ridge re-
presented by a minute cusp
between outer extremities of (J-
shaped figure, the space between
the limbs of the U occupied
]iy a smooth depression ; posterior margin of crown with a ridge
terminating internally like anterior ridge, but not extending to
outer margin. The principal variations of this pattern are : in
pm^, second limb of [J, with its outer cusp somewhat reduced ;
in m^, second limb of U obsolete and distorted, not extending to
outer border of tooth, the posterior half of crown occupied by a
shallow, basin-shaped area usually bearing a low but evident
cusp on its inner border. Lower molariform teeth with the
same elements as in Sciurus vulgaris, but crowns higher and
more compressed, the central depression deeper and narrower,
the cusps high and conspicuous, particularly the antero-external
and antero-internal, which are connected by a well-developed
ridge having much the appearance of a common base to the two
cusps ; inner side of bases of two outer cusps joined by a low
ridge, beyond which lies a small but rather deep depression.

Measure luents. — Averaije and extremes of four adults from

Fig. 189.
Cltellus citellus. Cheek-teeth, x

CITELIiUS 929

Hii'schberg, Bohemia: head and body, 200 (195-215) ; tail, 6G
(60-73); hind foot, 36 "6 (35 "4-38 '4). Average and extremes
of six adults from Znaim, Milhren, Austria- Hungary : head and
body, 209-1 (200-220); tail, 63-7 (58-70); hind foot, 36-4
(35 "6-37). For cranial measvirements see Table, p. 930.

Specimens examined. — Thirty-nine, from the following localities : —

Adsteia-Hungary : Haida, Arva, Bohemia, 1 ; Hirschberg, Bohemia, 5 ;
Znaim, Miihren, 7 ; Csallokoz-Somorja, western Hungary, 4 ; central
Hungary, 1; Sarvar, Eisenburg, Hungary, 1 (U.S.N.M.); Monos Petrie,
Bihar, 2 ; Csehtelek, Bihar, 6.

RouMANiA : Kustendje, 2 ; Dobrudscha, 3 (B.M., U.S.N.M. and Genoa) ;
Moldavia, 3.

Budgabia: Bogoroff, Sofia, 1; Sofia, 2 (Andersen).

Turkey: Ob-Meidan, 1.

2(5,2 9.

Hirschberg, Bohemia,
Austria-Hungary.

Lord Lilford (p).

8. 9. 10. 1-4.

d, 3 9.

Znaim, Miihren.

Lord Lilford (p).

8. 9. 10. 1-4.

4.

Csallokdz-Somorja, Press-
burg, 400 ft.

Budapest Museum (e)

. 94. 3. 1. 33-36.

2 9.

Monos Petrie, Bihar.

Hon. N. C. Roths-
child (c & P).

9. 7. 23. 1-2.

2(5,4 9.

Csehtelek, Bihar.

Hon. Mrs. Chas.
Rothschild (c & p).

10. 9. 14. 2-7.

2.

Kustendje, Roumania.

E. Cavendish Taylor

(c & P).
Purchased (Pruliere).

5. 5. 6. 16-17.

1.

Dobrudscha.

86. 4. 2. 3.

1.

Moldavia.

Dr. E. Riippell.

42. 9. 4. 1.

1.

Moldavia.

Purchased (Lead-
beater).

43. 9. 11. 2.

1.

Moldavia.

Dr. J. E. Gray (p).

123. a.

9.

Bogoroff, Sofia, Bulgaria.

Dr. A. Richards

(G & P).

2. 10. 5. 1.

CITELLUS SUSLICA Gueldenstaedt.

1770. Mus snslica Gueldenstaedt, Nov. Comm. Acad. Sci. Imp. Petrop.,

XIV, pt. I, p. 389 (Voronezh, Russia).
1770. [Aliis citellus] var. guttatiis Pallas, Nov. Comm. Acad. Sci. Imp.

Petrop., XIV, pt. I, p. 506, pi. 21, fig. 2 (Doubtfully a name).
1792. \_Arctomijs'] citellus & leucopictus Donndorif, Zool. Beytrjige, i, p. 486

(Renaming of guttatus).
1845. Sp[ermophilus'] guttulatus Schinz. Synop. Mamm., ii, p. 70 (Renaming

of guttatus).
1910. Citellus guttatus Trouessart, Faune Mamm. d' Europe, p. 128.

Type locality. — Voronezh, Chernigof, Russia.
Geographical distribution. — Central Russia and eastern
Hungary.

Diagnosis. — Slightly smaller than Citellus citellus, and tail
relatively shorter ; back marked with sharply defined whitish
spots about 4 mm. in diameter ; skull smaller than that of
C. citellus, the rostrum and palate narrower and incisiv{! foramina
much reduced in size.

Colour. — Dorsal area from nape to base of tail a tawny russet
faintly clouded by blackish hair tips, and thickly spotted with

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MARMOTA 931

buffy white, the spots roundish, vai^ying from 2 to 5 mm. in
diameter, those on middle of back smallest and most distinct,
those at sides larger and tending to become confluent. Sides
pale cream-bufF with faint dark shading, this colour passing
rather abruptly into bright buff of chest, belly, and inner
surface of hind legs. Chin, throat, and under side of neck as
far as fore legs like sides but still paler, rather sharply con-
trasted with colour of belly. Muzzle, cheeks, eye-ring (about
2 mm. wide) and ill defined area extending behind eye nearly to
ear, cream-buff. Crown and stripe 5 mm. wide below eye russet
sprinkled with cream-buff. Feet pale cream-bufl*; heel and
ankle clear russet. Tail a dull butfy russet below, fringed with
cream-buff, a grizzle of blackish and cream-bufF above.

Shdl and teeth. — The only skull examined is imperfect. It
is smaller than that of C. ritellus, and apjiarently less deep at
middle, though dorsal profile is similarly convex throughout.
Rostrum narrower and not so deep as in C. ciielliis, but not
peculiar in form. Incisive foramina reduced to mere slits barely
5 mm. in width, and so short that their length is scarcely equal
to diameter of alveolus of upper incisor. Palate and interorbital
region narrower than in C. citeUus, but interorbital breadth
greater than breadth of rostrum. Mandible smaller and less
robust than that of the related species but not peculiar in form.
The teeth do not differ appreciably from those of C. citeUus.

Measurements. — Adult female from Galicia, Hungary (from
well-made skin) : head and body, 200 ; tail, 40 ; hind foot, 32 ;
ear, 7. For cranial measurements see Table opposite.

Specimen examined. — One, from Galicia, Hungary.

1. Galicia; Hungary. Purchased (Pruliere). 86.4.2.2.

Genus MARMOTA Blumenbach.

1777. Glis Erxleben, Syst. Regni Anim., i, p. 358 (part). Not Glis Brisson,
1762.

1779. Marmota Blumenbach, Handb. der Naturgesch., i, p. 79 {alpina =

marmota).

1780. Arctomys Schreber, Saugethiere, pi. ccvii [alpina = marmota).
1780. Lagomys Storr, Prodr. Meth. Mamm., p. 39 (Renaming of Arctomys).
1811. Lipura Illiger, Prodr. Syst. Mam. et Avium, p. 95 {liudsonius =

mo7iax).

1857. Arctomys Blasius, Saugethiere Deutschlands, p. 278.

1904. Marmota Trouessart, Catal. Mamm. Viv. Foss., Suppl., p. 343.
Frisch (Natursystem der vierfiiss. Thiere, 1775, p. 9), cited as
authority, but this writer not binary (see Thomas and Miller,
Ann. and Mag. Nat. Hist., 7th ser., xvi, pp. 461-464, October,
1905).

Ty2)e specific. — Martiiota alpina 151ume:abach = 7lf«.s' maniiota
Linnajus.

Geographical distribution. — Northern Hemisphere from the

3 0 2

032 RODENTIA

Alps eastward through Asia and North America to the Atlantic
coast of the United States.

Characters. — Strictly terrestrial Sciiiridse, including the largest
members of the family ; form robust, Meline rather than Sciurine
in general aspect ; tail short and bushy, scarcely or not flat-
tened ; cheek pouches rudimentary or absent ; cranial and dental
characters essentially as in CiieUns, but skull with rostrum
nearly as deep as brain-case, and with heavy postorbital processes
standing out nearly at right angles with main axis of skull ;
anterior upper premolar always large and well developed.

Remarks. — About two dozen species of Marmota are now
known, the majority of them occurring in Central Asia. One
of these extends its range westward into eastern Hungary ;
while a single species is peculiar to the Alps and Carpathians.

KEY TO THE EUROPEAN SPECIES OP MARMOTA.

Colours noticeably varied, the head blackish, with
contrasted pale muzzle, the back much less yel-
lowish than sides, the tail conspicuously black
on terminal third ; skull with posterior border
of postorbital process decidedly in front of nar-
rowest portion of postorbital constriction (Alps
and Carpathians) M. mannota,']). 932.

Colours not varied, the entire animal a yellowish
brown overlaid on upper parts with black, the
head slightly darker than body, without con-
trasted j)ale muzzle, the tail inconspicuously
blackish at tip ; skull with posterior border of
postorbital process nearly over narrowest portion
of postorbital constriction (eastern Hungary) ... M. bobak, p. 937.

MARMOTA MARMOTA Limifeus.

1758. [Mus] marmota Linnaeus, Syst. Nat., i, 10th ed., p. 60.

1779. [Marwote] alpina Blumenbach, Handb. der Naturgesch., i, p. 80
(Substitute for Marmota).

1801. A[rctomys] m[annota] tigrina Bechstein, Gemeinn. Naturgesch.
Deutschlands, i, 2d ed., p. 1029.

1801. A[rcto)nys] m[a?-7nota] alba Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 1030.

1801. Alrctomys] 7n[ar77iota] nigra Bechstem, Gemeinn. Naturgesch.
Deutschlands, i, 2d ed., p. 1030.

1857. Arctomys marmota Blasius, Siiugethiere Deutschlands, p. 280.

1904. \_Marmota~\ marmotta Trouessart, Catal. Mamm. Viv. Foss., SuppL,
p. .343.

1910. Marmota marmota Trouessart, Paune IMamm. d'Europe, p. 129.

Type locality. — Alps.

Geographical distribution. — Alps and Carpathians. Frequently
referred to as occurring in the Pyrenees, but this probably an
error.*

Diagnosis. — Size medium (head and body about 550, tail

* See Trutat, Bull. Soc. d'Hist. Nat. de Toulouse, xi, p. 110, 1877.

MARMOTA 933

about 150, hind foot aljout 93 mm. ; condylobasal length of fully
adult skulls. 92 to 98 mm.) ; colours noticeably varied, the head
usually blackish with contrasted pale muzzle and cheeks, the
back blackish or greyish, the shoulders, sides and underparts
tawny or yellowish, the tail with extensive black terminal area ;
skull somewhat elongate, the postorbital processes standing so
far forward that their- hinder border is decidedly in front of
narrowest portion of postorbital constriction ; anterior face of
incisors deep orange.

External cliaracfers. — Form heavy and badger-like, the legs
short and muscular, the tail about one-third as long as head and
body, densely haired but not flattened, the head short, with
inconspicuous ears and rather broad, rounded muzzle. Ear
shorter than adjacent fur, its general outline rounded, but with
a slight backward projection above ; when laid forward the
extremity extends less than half way to eye. Muzzle pad bare
at middle, its outline not sharply defined above, the naked area
extending downward as a narrow line across upper lip, and out-
lined on each side by a deep groove running downward from
corresponding nostril. Feet robust, with long, slightly curved
fossorial claws, those of fore feet somewhat the better developed ;
fore foot with no trace of pollex or its nail,* the third digit
decidedly longer than the others, the fourth, second and fifth
successively shorter ; palm naked, with three confluent pads at
base of digits, and two larger, rounded pads occupying posterior
half of palm, that on inner side somewhat projecting ; hind foot
with third digit slightly longer than the sub-equal third and
fourth, the fifth and first successively much shorter ; hallux well
developed, extending to middle of first phahinx of second digit,
its claw like those of other toes but slightly smaller ; sole naked
except at heel ; four sub-equal, partly confluent pads at base of
digits, immediately behind these two smaller tubercles, that of
inner side smallest. Mamma? : p 1 — 1 ; a 2 — 2 ; i 2 — 2 = 10.

Colour. — Upper parts a grizzle of black and light buff", the
exact proportions of the two sufficiently variable to produce
considerable individual differences in the general effect, but top
of head usually blackish finely ticked with whitish buff" in
evident contrast with less dark neck, and back behind shoulders
usually darker than region in front ; underfur of back slaty
black at base, the tips of the hairs forming a buffy or greyish
band about 7 mm. wide ; when this band is conspicuously grey
it imparts to dark area of back a peculiar metallic sheen or
lustre ; muzzle nearly to level of front of eye whitish or buffy in
abrupt contrast with dark crown ; cheeks a coarse grizzle of
blackish and buff"y white ; underparts and inner surface of legs
a clear yellowish tawny-buff', this colour suff"using outer surface
of legs and sides of body, also a conspicuous area extending
upward immediately behind shoulder and partly separating dark

* Verified in twenty skins.

934

region of neck from that of back ; feet butt', soinewliat lighter
than outer surface of legs ; t-iil black or blackisli brown, sutiused
to a varying degree with buft', the light colour most conspicuous
at and near base.

SJiuU. — Apart from its much greater size the skull of Marmota
marvwta resembles that of Cltellus citellus in most of its more
essential features. The depth at front of brain-case and in
interorbital region is relatively less, and in adult individuals
there is a well developed sagittal crest on posterior half of brain-
case, with the resvilt that dorsal profile while equally convex in

Fig. 190.
Marmota tnarmota. Slightly reduced.

front is noticeably flattened posteriorly as compax*ed with the
smaller animal. Ventral profile with no special peculiarities.
Outline of brain-case somewhat obscured by the enlarged
squamosals, the area of which, when skull is viewed from above,
considerably exceeds that of parietals ; constriction in front of
lambdoid region relatively wider and more evident than in
Citelhis citellus. Lambdoid crest high, sometimes projecting
backward sufficiently to conceal occipital region in dorsal view.
Occiput lower relatively to its width than in C. citellus, the arch
slightly pointed at middle and essentially as high as middle

KIG. 191.
Marmota marmuta. Slightly reduced.

Fig. 192.
Marmota maniwta. .Slightly reduced.

93G

of brain-case ; paroccipital process robust, not very long, its
extremity projecting silghtly beyond level oi lower edge of
condyle ; basioccipital broad and sliort, its lateral processes very
distinct ; auditory bulla moderately large, evenly inflated, the
region below meatus constricted to form a short but evident
neck. Interorbital region broad, squarish, the median portion
depressed ; postorbital processes heavy, standing out abruptly,
not much bent downward, separated from brain-case by a wide
space ; orbit about two-thirds surrounded by bone. Nasals
abruptly bent downward anteriorly, considerably narrowed
posteriorly, so that their combined breadth at level of posterior
border of premaxillaries is only a little more than half greatest
anterior breadth. Incisive foramina extending back slightly
into maxillaries, their outer borders nearly parallel. In other
respects the skull so closely resembles that of Citellus citellus,
due allowance being made for its greater size and angularity, as
to require no detailed description.

Teeth. — In general form the teeth bear a strong resemblance
to those of Citellus citellus. Upper incisor with anterior face
slightly flattened, somewhat distorting the otherwise evenly
semicircular outline of cross-section, the enamel deep yellowish

brown, its surface marked by about
four faintly indicated longitudinal
ridges ; enamel of lower incisor
minutely pitted. Enamel pattern
of upper cheek-teeth essentially as in
Citellus citellus, but inner tubercles
lower and less compressed, the length
of base slightly greater than height
of tubercle, transverse ridges form-
ing a V-shaped rather than U-shaped
figure when worn, and anterior and
posterior ridges continuous at their
inner extremities with anterior and
posterior bases of inner tubercle.
Other details in which the upper
cheek-teeth differ from those of C.
citellus are as follows : no indication
of minute secondary cusps or ridges
between main cusps of outer side ; small premolar with
crown area barely half that of succeeding tooth, its anterior
and posterior depression tending to join on inner side to form
a narrow semicircular concavity ; depression on posterior half
of m^ flattened rather than basin-shaped. Lower cheek-teeth
differing from those of Citellus citellus in their relatively much
lower, more robust cusps and larger crushing surfaces, and the
relatively greater importance of ridge joining inner bases of
outer cusps, this ridge so high that it begins to wear almost as
soon as the cusps themselves, forming a conspicuous feature of

Marmota tiiannota.
Cheek-teeth, X 2.

MARMOTA

937

the enamel pattern ; premolar with a small hut e\ idcnt cinqulum
cusp on anterior hoi-der.

Mcasiiremenis. -Adult male from Barceloiniette, Basses- Alpes,
Franco: head and body, 577 ; tail, 153 ; hind foot, 94 ; ear, 30.
Adult male and female from Canton St. Gallen, Switzerland :
head and body, 540 and 530 ; tail, 159 and 150 ; hind foot, 90
and 87. Two adult males from Grisons, Switzerland : head and
body, 503 and 530; tail, 130 and 160; hind foot, 90 and 95.
For cranial measurements see Table, p. 939.

Sj^ecimens examined. — Forty-seven, from the following localities: —

France: Near Barcelonnette, Basses-Alpes, 8 (B.j\I. and Mottaz).

Switzerland : Salanfe (at foot of Dent du Midi), Valais, 1 (IMottaz) ;
Meiringen, Bern, 2 ; Engstlen Alp, Unterwalden, 1 ; Engelberg, Unter-
walden, 1 (U.S.N.M.) ; Schaufigg, Grisons, 3 ; Canton Grisons (no exact
locality), 4 (U.S.N.M.) ; Rheinwaldhorn, Grisons, 1.

Italy : Vagna Valley, near Site. Rosa, Novara, 1 (Genoa) ; Aosta
Valley, Turin, 20 (Turin) : Argentera, Cuneo, 3 (Turin) ; Maritime Alps, 1
(Genoa).

Austria-Hungary : Felkaerthal, Tatra IMountains, Hungary, 1.

2(^,2 9. Barcelonnette, Basses-Alpes,

France. (C. Mottaz.)
Skull. Meiringen, Bern, Switzer-
land. [Kocserman.)
1. Engelberg, Unterwalden.

{Koeserman.)
S, 2 9. Schaufigg, Grisons.

{Zollikofer.)
S. Rheinwaldhorn, Grisons.

{Zollikofer.)
1. Felkaerthal, Tatra Moun-
tains ; Hungary.

MARMOTA BOBAK Miiller.

177G. Mus bobak P. L. S. Miiller, Natursyst. Suppl. u. Regist. -Band, p. 40

(Poland).
1778. Mils arctomys Pallas, Nov. Sp. Quadr. Glir. Ord., p. 75 (Poland).
1780. Arctomys hobac Schreber, Siiugethiere, pi. ccix (description, iv,

p. 738, 1782). Renaming of Mus arctomys.
1811. Arctomys baibac Pallas, Zoogr. Rosso-Asiat., p. 155 (Poland).
1857. Arctomys bobac Blasius, Siiugethiere Deutschlands, p. 283.
1904. [Marinota] bobac Trouessart, Catal. IVIamm. Viv. Foss., Suppl., p. 343.
1910. Marmota bobac Trouessart, Faune Mamm. d'Europe, p. 129.

Type locality. — Poland.

Geographical distribution. — From Poland and eastern Hungary
(Galicia and Bukowina) eastward into Asia. Exact limits of
range not known.

Diagnosis. — Size essentially as in Maniiota uiarmota ; colours
uniform, the entire animal a yellowish brown overlaid on upper
parts with black, the head slightly darker than body, without
contrasted pale muzzle or cheeks, the tail blackish at extreme tip
only ; skull relatively broader than in M. marmota, and post-
orbital processes standing so far back that their hinder border is
nearly over narrowest portion of postorbital constriction ; teeth

0. Thomas (p).
Tomes Collection.

8. 8. 10. 145.

63-65.
7. 1. 1. 195.

Tomes Collection.

7. 1. 1. 131.

0. Thomas (p).

2. 8. 4. 31-33.

0. Thomas (p).

2. 8. 4. 30.

Poprad ]\Iuseum

(E).

7. 4. 19. 1.

938

RODENTIA

somewhat heavier than those of M. tnarmota, especially the lower
molars ; anterior face of incisors whitish or pale orange.

External characters. — Essentially as in Marmota marmota, but
thumb present as a small though distinct tubercle bearing an
evident appressed nail. Mammai : 2^ 2 — 2, a 2 — 2, i 1 — 1 = 10.

Colour. — ^Entire animal a light yellowish brown, the head
suffused with a darker brown, the back, sides of body and outer
surface of limbs clouded with blackish hair-tips, these most
numerous over posterior half of back, but nowhere obscuring the

Fig. 194.
Marmota bobak. Slightly reduced.

ground colour ; feet a clear yellowish brown ; underparts and
inner surface of limbs like back but slightly darker and more
tawny, the hairs without blackish tips ; tail essentially similar,
the extreme tip dark brown or blackish ; underfur everywhere
dull yellowish, the hairs with slaty bases.

Skull and teeth. — The skull, though of about the same length
as that of Marmota marmota, is broader and more robust in
general form, and all ridges for muscular attachment are more
prominent. Nasals less abruptly bent downward anteriorly and
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RODENTIA

posterior border of premaxillary fully three-fourths greatest
combined breadth anteriorly, f )rbit largei- than in the related
species, the postorbital process situated so far back that its
tip lies over a point decidedly behind middle of temporal
fossa instead of over middle, and its posterior border nearly
overhangs deepest part of postorbital constriction and anterior
border of squamosal. Brain-case noticeably broader in proportion
to its length, and foramen magnum wider relatively to its height.
Auditory bulla? larger than in Marmota marmoia, the area of
each bulla appearing greater than that of basioccifjital and con-
dyles instead of about equal to it. Palate relatively narrower
posteriorly, and incisive foramina wider in proportion to their
length than in M. marmota. Mandible more I'obust than in the
related animal, especially in front ; sigmoid flexure longer and
less concave.

Teeth essentially as in Marmota marmota as regards form, but
anterior surface of incisors pale yellowish instead of deep orange,
and cheek-teeth more robust, a character especially noticeable in
second and third lower molars.*

Measurements. — Male and female from Russia (skins) : head
and body, 600 and 530 ; tail, 190 and 160 ; hind foot, 90 and 89.
For cranial measurements see Table^ p. 939.

Specimens examined. — Two, both from Russia.

Family PETAURISTID^.

1855. Pteromyinse Brandt, M^m. Acad. Imp. Sci., St. Petersburg, 6th ser.,

Sci. Nat., VII, p. 151.
1857. Sciurina Blasius, Saugethiere Deutschlauds, p. 266 (part).
1879. Pteromidse Anderson, Anat. and Zool. Researches, Expeds. Yunnan,

p. 278.

1891. Sciurinfe Flower and Lydekker, Mammals, living and extinct, p. 450
(part).

Geographical distribution. — Wooded portions of Northern
Hemisphere south to the Malay region and the southern United
States. In Europe confined to Scandinavia and northern Russia.

Characters. — Like the Sciuridse, but with fore and hind limbs
connected by a broad fold of skin extending to wrists and ankles,
supported anteriorly by a stiff cartilaginous process growing from
wrist, the membrane serving as a parachute by means of which
gliding flight is effected.

Remarks. — The Pefaiiristidse are so sharply differentiated from
the Sciuridse, that it seems preferable to recognize the two groups
as families. Eight genera are known,| one of which is repre-
sented in Europe.

* This difference between the teeth of M. marmota and M. bobak is of
much importance in determining the identity of subfossil remains.

t See Thomas, Ann. and Mag. Nat. Hist., 8th ser., i, pp. 1-8, January,
1908.

SCIUROPTKRUS 941

Genus SCIUROPTERUS F. Cuvier.

1825. Sciuropterus F. Cuvier, Dents des Mammiferes, p. 255.
1857. Pteromys Blasius, Saugethiere Deutschlands, p. 268.

Type species. — Sciurus volans Liniiieus = Pteromys russicus
Tiedemann.

Geographical distribution. — As in the family.

Characters. — No membrane extending between tail and hind
leg ; structure of molars essentially as in Sciurus, the enamel not
thrown conspicuously into wrinkles, the transverse ridges well
developed ; posterior upper premolar about as large as first molar,
never decidedly smaller.

ItemarTis. — The genus Sciuropterus as thus defined contains
about thirty species, most of them Malayan. It is divided into
four sub-genera, to one of which (Glaucomys) belong all the
American members of the family. The single European repre-
sentative of the group is the type of the sub-genus Sciuropterus,
the three or four other known members of which occur in central
and northern Asia and in Japan.

SCIUROPTERUS RUSSICUS Tiedemann.

1758. [Sciurus] volans Linuseus, Syst. Nat., i, 10th ed., p. 64 (Sweden).

Not Mus volans Linnseus, I.e., p. 63.
1808. P[teromys'] russicus Tiedemann, Zoologie, i, p. 451 (Russia).
1822. Pteromys sibiricus Desmarest, Mammologie, ii, p. 342 (Substitute

for volans).
1843. Pt[eromys'] vulgaris Wagner, Schreber's Saugthiere, Suppl., in, p. 228

(Substitute for volans).
1857. Pteromys volans Blasius, Saugethiere Deutschlands, p. 269.
1903. Sciuroptertis russicus Allen, Bull. Amer. Mus. Nat. Hist., xix, p. 132.

March 31, 1903.
1910. Sciuropterus russicus Trouessart, Faunc Mamm. d'Europe, p. 116.

Type locality. — Russia.

Geographical distribution. — Wooded portions of northern
Siberia and Russia, southward nearly to the boundary of north-
eastern Germany, eastward into northern Scandinavia.

Diagnosis. — Size median (head and body about 170 mm., tail
about 110, hind foot about 33, condylobasal length of skull
about 37) ; colour of upper parts a light silvery or buffy grey ;
skull with nasals abruptly constricted just behind middle, not
narrowing gradually backward ; auditory bulla very large, its
greatest antero-posterior diameter equal to distance between
bulla and anterior portion of first molar.

External characters. — Form not essentially different from that
of Sciurus vulgaris, but appearing broadened and flattened on
account of the more depressed tail and the wide flying membrane.
Head short and round ; eyes xevy large ; ear low,' rounded,
almost concealed in the fur, the outer margin faintly concave

942 RODKNTIA

above, both surfaces clothed with fine short hairs which never
form tuft at tip. Feet relatively shorter and broader than in
S. vuhjaris, the digits much less graduated, particularly those of
hind foot, the claws rather short, strongly curved, very acutely
pointed, those of hind foot nearly concealed by tufts of curved
hairs lying over them ; front foot with fourth digit longest, third,
fifth and second successively shorter, supporting cartilage about
twice as long as foot, thumb an inconspicuous tubercle with
rudimentary flattened nail ; hind foot with three middle digits
sub-equal and longest, first extending to base of (daw of second,
fifth a little beyond base of first phalanx of fourth ; palmar and
plantar tubercles bare, arranged as in S. vulgaris, their size
relatively smaller ; palms thinly haired between pads, soles with
dense growth of long woolly hairs especially conspicuous along
inner side and adding much to apparent width of foot. Fur of a
peculiar silky texture, noticeably softer and more dense than in
,S^. vulgaris, this particularly marked in the broad, much flattened
tail ; hairs along anterior edge of membrane (in region of
supporting cartilage) noticeably stiffened.

Colour.- — Upper parts a uniform pale, silvery grey, with a
faint buffy cast, the slaty under colour appearing irregularly at
surface when hairs are disarranged, particularly near outer edge
of lateral membrane, and on feet ; everywhere there is a very
faint suggestion of dark " lining," produced by the presence of
black-tipped longer hairs ; cheeks paler and less buffy than
crown ; a narrow blackish eye-ring ; underparts and inner
surface of limbs dull buffy white, everywhere inconspicuously
sprinkled with blackish hairs ; tail more buffy than body (nearly
the cream-buft' of Ridgway), the upper and lower surfaces and
the tip much clouded with dusky, the sides almost clear.

Skull. — As compared with that of Sciurus vulgaris the skull
is much smaller, but at the same time with all lines more
exaggerated, it might almost be said distorted, and contrasts
more abrupt. Profiles essentially as in Sciurus vulgaris, but
nasals abruptly curved downward at tip, occiput slightly concave
owing to the more posterior position of condyles and paroccipital
processes due to enlargement of bullfe, and ventral profile more
nearly parallel with dorsal profile, its posterior portion carried
further downward by the large bullfe. Brain-case squarish in
general outline when viewed from above, its posterior margin
broadly and evenly rounded, its width posteriorly about double
that of postorbital constriction ; surface essentially smooth,
though a faint ridge may usually be traced diagonally across
each side of parietal, the ridges curving inward posteriorly and
meeting in median line somewhat in front of lambdoid region ;
in old individuals a tubercular projection is developed on edge of
tem])oral fossa at antero-external corner of parietal, much as in
Lepus ; lambdoid crest low and rounded ; \iewed fx'om the side
the brain-case is more depressed than in Sciurus vulgaris, so that

SCIUROPTERUS

943

less of the surface of parietal is visible ; occiput very low, its
height above lower lip of foramen magnum about one-half
mastoid breadth, its dorsal and lateral margins forming a flattened
curve much less than a semicircle in extent ; foramen magnum
decidedly wider than high ; surface of occipital with very obscure
swellings between foramen magnum and lambdoid ridge ; par-
occipital processes very slender ; basioccipital rather narrow and
long, its width anteriorly about one-half median length, its
lateral ridges obsolete, not tending to form processes as in Sciurus
vulgaris and Citellus ; auditory bullae sub-circular in general
outline, their form essentially as in Sciurus vulgaris, but their

Fig. 19;').
Sciuropterus russicun. Nat. size.

size actually as well as relatively greater, the antero-posterior
diameter equal to distance from front of bulla to front of j^i, the
lateral diameter to outer margin of meatus fully 1^ times least
width of basioccipital ; anterior upper margin of meatus developed
into a conspicuous forward-curved flange ; jjetrosal tending to be
exposed in region above meatus instead of on occipital surface.
Interorbital region decidedly longer than broad, its surface
noticeably concave laterally, so that bases of ])ostorbital processes
are evidently above level of median region ; notch at anterior
base of postorbital process large, always open ; postorbital
processes long and slender, strongly curved downward, the orbit

9i4 KODENTIA

actually as well as relatively largei* than iu Sciuriis vulgaris, and
less well defined posteriorly owing to absence of distinct anterior
concavity and median projection on upper border of zygoma,
its orifice directed more upward. Zygoma no more abruptly
spreading than in Sciurus vulgaris, but the arches appearing
wider by contrast with the narrower interorbital region, and also
on account of the much more abrupt angle at which the anterior
surface of zygomatic root meets outer surface of rostrum ; median
portion of arch splayed so strongly outward as to be almost
horizontal ; jugal projecting posteriorly behind zygomatic process
of squamosal, its upper border without angular median projection,
its anterior extremity running upward nearly to lachrymal ;
anteorbital foramen small, nearly circular, scarcely in front
of level of anterior premolar, hidden in lateral view Vjy the
conspicuously projecting process formed by lower portion of its
outer border. Rostrum relatively somewhat shorter than in
Sciurus vulgaris, its base so narrow that its sides are nearly
parallel, the least depth behind incisors somewhat greater than
width in same region ; nasals rather short, abruptly narrowed
posteriorly, their slightly eniarginate hinder border lying at level
of posterior base of zygomatic root, the moderately expanded
nasal branches of premaxillaries scarcely extending behind them.
Palate relatively about as wide as in Sciurus vulgaris ; incisive
foramina large but lying almost entirely within premaxillaries,
the posterior border of which is therefore relatively much further
back (under middle of zygomatic root) than in Sciurus vulgaris,
the greatest length of foramina exceeding least width of rostrum
and equal to somewhat more than half diastema ; region behind
notch back of m-* conspicuously thickened ; mesopterygoid space
about as long as in Sciurus vulgaris, but conspicuously wider
anteriorly than posteriorly. Mandible with anterior portion of
ramus less deepened and more curved than in Sciurus vulgaris ;
posterior region so deep and short that greatest depth (exclusive
of coronoid) is nearly equal to distance from condyle to front of
mj, while in >S^. vulgaris the same depth barely equals distance
from condyle to back of m^^ ; coronoid process low and short,
strongly curved backward ; angular process large, its broad apex
curving abruptly outward, its lower border greatly developed
and forming almost a separate lobe bent strongly inward.

Teeth. — Incisors less compressed than those of Sciurus vulgaris,
the roots forming no evident protuberances, the anterior surface
smooth, deep orange in colour. Anterior upper premolar terete,
its crown simple, about one-fourth or one-fifth that of succeeding
tooth in area. Upper molariform teeth resembling those of
Sciurus vulgaris in their relative sizes and general structure ;
anterior border of crown more elevated, its outer extremity
appearing as a distinct cusp nearly as high as the two main
cusps when tooth is viewed from the side ; main transverse
ridges relatively higher and narrower, the second, except in m^.

SCIUKOPTERUS

945

cut l)y a deep posU'rior re -entrant fold, so tliat its inner
extremity stands as a nearly free subterete cusp; inner cusp
relatively higher, obscurely trilobed owing to the presence on
palatal border of two well marked depressions extending from
apex nearly to base of crown (these become obsolete as the tooth
wears away) ; in nt^, m? and m^, a short process extends outward
from base of inner cusp into space between tirst and second
transverse ridges, while in pm:^, ni^
and )ir a similar but less developed
process lies in space between second
and third ridges ; third and fourth
ridges of ni-^ smaller than in the
other teeth, but well developed, so
that there is no indication of a pos-
terior basin-shaped area on surface of
crown. Mandibular teeth like those
of Sciurus vulgaris in relative sizes
and in general form except that j/;.,
is longer and more narrowed pos-
teriorly. The pattern of enamel
folding, while essentially like that of
S. vulgaris in its main features, is
more complicated, owing to the greater development of all the
smaller cusps and folds ; main cusps relatively higher and more
slender, their bases extending further inward toward middle of
crown, this together with the greater development of cusp and
ridge coiniecting inner portion of bases of outer main cusps
greatly reducing area of median crushing surface, which is
scarcely basin-shaped and which occupies less than half area of
crown ; antero-external ridge well developed in the three molars,
terminating in a distinct cusp-like elevation at middle of anterior
border of crown ; a deep depression at anterior base of postero-
external cusp ; a similar but larger depression behind base of
postero-internal cusp of m.^, the cingulum rising back of it so
as almost to form a supplemental cusp.

3Ieasiiremeuts. — Adult male from Finland : head and body,
170; tail, 109; hind foot, 35; ear, 19. A second adult from
the same region : tail, 128 ; hind foot, 37. For cranial measure-
ments see Table, p. 940.

Specimens examined. — Two from Finland (B.M. and U.S.N.M.) ; also
one from Russia, and a stuffed specimen from unknown locality.

Sci'uropterus russicus.
Cheek-teeth, x 5.

1. Finland.
1. Russia.
1 st. Europe.

Dr. H. Schulman (p).
Purchased (Parzudaki).
Purchased (Brandt).

1. 6. <J. 1.
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42. 5. 16. 11.

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CASTOR

947

Family CASTORID^.

1821. Castoridn; Gray, London Med. Ropos., xv, p. 302, April 1, 1S21.

Geographical distribution. — Forested portions of the Northern
Hemisphere, south to the Mediterranean region and the southern
United States.

Characters. — Essentially as in the Sciuridx, but skull without
postorbital processes, and cheek-teeth rootless, with re-entrant
enamel folds ; premolars -}- ; angular portion of mandible short,
broadly rounded ; size large ; form heavy ; tail broad, flattened
from above downwards, its surface scaly ; habits aquatic.

BemarJea. — Though several extinct members of the group are
known the family Cantoridse is represented by only one living-
genus, widely distributed in the northern portion of both the
Old and New Worlds.

Genus CASTOR Linnteus.

1758. Castor Linnfeus, Syst. Nat., i, 10th ed., p. 58 {Jlbcr by tautouyray).
ISOG. Fiber Dumeril, Zoologie Analytique, p. 18, diagnosis on p. I'J.
(Substitute for Castor.) Not Fiber Cuvier, 1800.

Tt/2)e species. — Castor fiber Linnteus.

Geographical distribution. — As in the family.

Characters. — General characters as in the family ; u[)pei"
molars sub-equal, each with one internal and three external
enamel folds.

BemarJis. — The genus Castor is the only living representative
of the family. It contains two closely related species, one
peculiar to North America, the other to the northern portions of
the Old World.

CASTOR FIBER Linnreus.

1758. [Castor] fiber Linnffius, Syst. Nat., i, 10th ed., p. 58 (Sweden).

1792. C[astor]f[iber] albtts Kerr, Anim. Kingd., p. 222.

1792. Clastor] f[iber] solitarius Kerr, Anim. Kingd., p. 224.

1801. Clastor^fliber] variegatus Bechstein, Gemeinn. Naturgesch. Dcutsch-

lands, I, 2d ed., p. 913 (Europe).
1801. C[astor] /[ibcr] fulvus Bechstein, Gemeinn. Naturgesch. Dcutsch-

lands, I, 2d ed., p. 913 (Europe).
1803. Castor gallin- Geoffrey, Catal. INIamm. du ]Mus. Nat. d'Hist. Nat.,

Paris, p. 168 (Banks of the Rhone, France).

1822. Castor niger Desmarest, Maniiualogie, pt. ii, p. 278 (no exact

locality).
1822. Castor varius Desmarest, Mammalogie, pt. ii, p. 278 (northern and

central Europe).
1822. Castor flavus Desmarest, Mammalogie, pt. ii, p. 278 (no exact

locality).
1829. [Castor fiber] |3. gallicus Fischer, Synops. ]\Iamm., p. 287 (Substitute

for galliiv).

3 p 3

948

RODENTIA

1833. Castor proprius Billberg, Linn. Samf., p. 3i footnote (Substitute for
fiber).

1857. Castor fiber Blasius, Saugethiere Deutschlands, p. 405.

1897. Castor fiber CoUett, Bergens Museums Aarbog, pp. 3-127. Habits in
Norway.

1907. Castor albicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch. Freunde,
Berlin, p. 216, October, 1907. (Dessau, Anhalt, Germany.) See
Lonnberg, Arkiv for Zoologi, v, no. 6, pp. 1-16, 1909.

1907. ? Castor vistulinus Matschie, Sitz. - Ber. Gesellsch. Naturforsch.
Freunde, Berlin, p. 219, October, 1907 (westei-n Poland).

1910. Castor fiber Trouessart, Faune Mamm. d'Europe, p. 130.

Tijpe locality. — Sweden.

Geographical distribiition. — Formerly the entire forested region
of Europe, west to Great Britain ; now exterminated eve j"y where
except in portions of Scandinavia, and along the courses of some
of the larger rivers of central Europe, as the Rhone, Elbe, and
Danube.

Diagnosis. — General features as in the family and genus ;
posterior extremity of nasals lying at or slightly behind level
of middle of orbit and decidedly behind termination of na.sal
branches of premaxillaries ; least depth of rostrum behind incisors
not equal to distance from gnathion to anteorbital foramen.

External characters. — General form heavy and thick-set, the
head large and rounded, the eyes and ears small, the legs short,
the feet large ; the tail somewhat less than half as long as
head and body, its width about one-third its length. Head
short, deep and rounded, the eye small, about midway between
ear and nostril ; ear low and rovmded, densely haired on both
surfaces, scarcely appearing above the surrounding fur ; meatus
small but apparently without any specially developed valves ;
muzzle pad mostly naked, but with fur encroaching on its
posterior border and along inner margin of nostril ; below pad
the lip is less densely haired than elsewhere, and the hairs tend
to converge along median line ; whiskers very coarse and bristly,
the longest extending a little beyond eye when laid back. Front
foot with no special peculiarities, the digits all well-developed,
with strong, little curved claws, the thumb shorter than the
other digits but armed with a large claw somewhat more curved
than the others ; palmar tubercles five, the three at bases of
median digits large, ill-defined, that at base of thumb not so
large as the others but better defined, its outer surface tending
to become smooth and horny, the posterior pad largest, covering
about one-third of entire palmar area ; surface of pads (except
as otherwise described) coarsely reticulate. Hind foot very
large and broad, the digits united by a web extending to base
of claws ; second and third digits equal and longest, their
claws heavy, scarcely concave on lower surface, considerably
larger than those of other toes ; first, fourth, and fifth toes
successively shorter than second and third, their claws smaller

CASTOR 949

and more curved, tliat of fourtli digit with a conspicuous, horny,
laterally-conijjressed supplement fully as large as claw itself
spi'inging from ball of finger beneath claw ; entire sole naked,
tlie surface coarsely reticulate ; three elongate, ill-defined pads
at base of median digits, a larger and better-defined tubercle
behind base of hallux ; entire sole between hallucal tubercle and
heel thickened and pad-like. Tail furred at extreme base, the
scales of naked portion wider than long, 4-G-sided, about 100
in longitudinal series, forty in circumference, those on lower
side somewhat larger and better defined than those on upper
side. Fur consisting of dense soft under portion, the hairs of
which are about 25 mm. in length, and an abundant growth of
rather coarse stiff longer hairs (50 to 60 mm.) nearly concealing
the underfur.

Colour. — General colour a peculiar clayey buff produced by
the longer hairs, and very uniform throughout, the nnderparts
a little less yellowish ; underfur light smoke-grey at base,
darkening to hair-brown or bister at tips, this colour rarely
showing thi-ough at surface ; muzzle, chin, and lower part of
cheeks greyish but not forming any decided contrast with
surrounding parts ; feet washed with a drab brown.

SJiull. — In general aspect the skull is low, robust and heavily-
built, though not con.spicuously ridged and furrowed, and not
noticeably widened ; the broadly expanded malar is a conspicuous
feature in lateral view. Dorsal profile moderately and irregularly
convex, usually with a flattened or slightly concave area between
orbits and another just in front of lambdoid region ; occiput
squarely truncate or somewhat overhanging, the condyles barely
or not visible when skull is viewed from above ; ventral profile,
owing to depth of rostrum and shallowness of brain- case not
diverging conspicuously in general direction from dorsal profile
except for the projection formed by the tooth-row and by the
auditory bulla. Brain-case rather narrowly ovate in general out-
line, its breadth over roots of zygomata contained about 1^ times
in length ; meatal tubes of auditory bulkw standing out con-
spicuously just in front of mastoid protuberances and rising to
level of dorsal surface of brain-case ; sagittal crest moderately
developed in adults, highest posteriorly, dividing anteriorly into
two low ridges curving forward and outward to rudimentaiy
postorbital processes ; lambdoid crest well developed, sloping
backward ; occiput low and broad, somewhat quadrate in outline
when viewed from behind, scarcely exceeded in height by middle
portion of brain-case, its median height slightly more than half
its greatest width, its surface with no special features ; par-
occipital process short and robust, not extending to level of
lower lip of foramen magnum ; floor of brain-case with deep pit
occujiying entire area between bullae, its edges well defined
throughout, its posterior margin formed by lower lip of foramen
magnum ; auditory bulla flask-sha^jed, the meatal tube about as

950

Fig. 197.
Castor fiber. X *.

CASTOR 951

long as transverse diameter of inflated portion of bulla, con-
spicuously ridged postero-externalJy, the main axis of flask
oblique but nearly upright, slojaing outward at an angle of only
about 30° ; portion of bulla appearing on lower surface of skull
about Ih times as long as wide, evenly inflated, the width at
middle about et(ual to that of transverse diameter of basioccipital
pit measured inside the rim ; bulla separated posteriorly and
postero-externally from paroccipital and mastoid processes by
a broad irregular groove. Interorbital region short and wide,
imperfectly marked ofi" from brain-case by the rudimentary post-
orbital processes. Zygomata heavy, gradually spreading, the
greatest zygomatic breadth about at glenoid level ; malar large,
greatly expanded at middle, its upper edge with projection
marking posterior limit of orbit, its depth through this projection
rather greater than diameter of orbit, its anterior extremity
iirmly ankylosed with lachrymal ; anteorbital foramen minute,
slit-like or rounded, vertical, hidden in lateral view by the
oblique ridge which forms its outer margin and extends down-
ward toward alveolus of hrst cheek-tooth. Rostrum heavy and
deep, its least depth nearly equal to depth of brain-case at
middle, but less than distance from anteorbital foramen to
gnathion ; region between incisors and lower rim of nares con-
spicuously thickened ; nasals gradually narrowing from befoi'C
backward, their greatest combined breadth usually much less
than half length, their posterior border extending about to level
of middle of orbit, and decidedly behind ends of nasals branches
of premaxillaries ; incisive foramina slit-like, parallel sided,
situated somewhat nearer to incisor than to cheek-teeth, their
greatest length about one-third that of diastema. Palate narrow,
its width between anterior cheek-teeth about one-half that of
alveolus, its width between posterior cheek-teeth about twice
that of alveolus (the relative width of palate greater in immature
individuals) ; posterior boi'der slightly behind level of vP, its
median sjDine well developed ; mesopterygoid space about two-
thirds as long as palate, its width posteriorly nearly three-
quarters its length, fully half its lateral border formed by the
long, robust, distally thickened hamular. Mandible very robust,
the ramus much thickened to accommodate the constantly grow-
ing roots of the cheek-teeth on outer side of large incisor-shaft,
the symphysis extending backward in adults to level of posterior
boi'der of first cheek-tooth ; articular process very short, the
surface of the condyle scarcely extending beyond a line joining
tip of high, abruptly recurved coronoid process with hinder
extremity of evenly rounded-oflf angular process.

Teeth. — Upper incisor heavy, its course scarcely indicated
on outer surface of rostrum, its root lying behind anteorbital
foramen ; shaft about as deep as wide, the antei'ior face slightly
curved, a little longer than sub-equal outer and inner faces, the
posterior border abruptly rounded ; enamel dark yellowish brown
in coloui", essentially smooth but with veiy obscure longitudinal

952

\\'rinklc3 ; lower incisor with root extending slightly beyond
W3, which is deflected outward to allow its passage ; section
of shaft much as in upper tooth but antero-posterior diameter
relatively greater and inner border more flattened. Cheek-
teeth large, their roots forming conspicuous capsule-like pro-
jections in orbit, and low but evident protuberances on lower
portion of outer side of lower jaw, both upper and lower root
capsules becoming less noticeable with advancing age ; outline
of crowns squarish, tending to be broader than long above,
longer than broad in the mandibular teeth, the outer border of
upjDer teeth and inner border of lower teeth straight, slightly

crenulate, the opposite borders dis-
tinctly bilobed. The pattern of
enamel folding is essentially the
same in all the teeth : three narrow
re-entrant folds on one side, a single
wider fold on the other ; in the
maxillary teeth the three folds are
on the outer side, while in the
mandibular teeth they are on the
inner side ; inner fold in maxillarv
teeth nearly straight, extending
obliquely forward to come in con-
tact with extremity of first outer
fold, or frequently in pm* to curve
slightly behind it ; outer fold in
mandibular teeth curving backward
between extremities of second and
third inner folds, with both of which
it is nearly or quite in contact. In
the maxillary teeth the first outer fold extends obliquely forward
half-way across crown, or in jjjm* occasionally about to inner
border, the second outer fold curves noticeably backward and
extends nearly to inner side of crown, and the third outer fold
extends directly inward or somewhat obliquely backward, its
length equal to or a little less than that of first fold, except
when the latter assumes its lengthened form. In the mandi-
bular teeth the first and third inner folds extend nearly across
crown, while the second is often distinctly shorter, all three are
somewhat wavy in outline, the first the least so.

Measurements. — Adult female from the Rhone, near Aries,
France : head and body about 820 ; tail about 380 ; bare portion
of tail (in dry specimen), 310 X 120; hind foot, 170; ear, 35.
Immature male from the same region : head and body about 650 ;
tail about 250 ; bare portion of tail (in dry specimen), 225 x 1 10 ;
hind foot, 160. For cranial measurements see Table opposite

Fig. 198.
Castor fiber. Cheek-teeth.

Specimens examined. — Six, from the following localities :—
NoBWAY : Froland, Kristiansand, 1.
England : Norfolk, 1 (skull).

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RODKNTIA

France : Rhone, near Avles, 2 ; near Aviguon, Vaucluse, 1.
Gebmany: Elbe River, 1 (U.S.N.M.).
Austria-Hungary: Danube, 1 (skull).

RemarJcs. — The material examined does not indicate the
existence of more than one form of beaver in Europe. Castor
fiber is nearly related to the American beaver, C. canadensis.
In the American animal the least depth of rostrum behind
incisors usually equals or slightly exceeds the distance from
gnathion to anteorbital foramen, and the nasals terminate
posteriorly at lachrymal level, slightly behind nasal branches of
premaxillaries.

t?, skeleton. Froland, Kristiansand, Dr. R. Collett (p). 97.10.14.1.
Norway.

Skeleton. Norfolk, England (from W. J. Angove (c). S5. 8. 4. 1.
peat-bog).

6. Aries, Bouches du Rhone, Lord Lilford (p). 94. 5. 30. 1.

France. (Dr. P. Siepi.)

?. Aries, Bouches du Rhone. Dr. P. Siepi (c). 5. 3. 9. 1.

1, Avignon, Vaucluse. Purchased (Dey- 85. 5. 2C. 1.

roUe).

Skull. Danube, Austria-Hungary. Purchased (Brandt). 875. b.

UNGULATA

955

Order UNGULATA.

1801. IJiujulata Bechstein, Gemcinn. Naturgcsch. Deutschlands, i, 2nd
ed., p. 182.

Geofiraplncal distrihution. — Essentially throughout all of the
larger land masses of the world, Australia excepted.

CT(fl7-ac<('r.s\— Terrestrial, herbivorous or omnivorous, placental
mammals with broad-crowned, tuberculated or ridged molars,
digitigrade feet, and hoofed digits.

BemarJcs. — The exact limit of the order Ungidata is still
a matter of controversy. The group is here considered as
equivalent to the Diplartlira. Thus restricted it contains two
main groujDs, the Artiodactyla, in which the main axis of the
foot lies between the third and fourth digits, producing the well-
known cloven hoof, and the Perissodactyla, in which the main
axis of the foot passes through the middle of the third digit.
All the European members of the order now existing in the
wild state belong to the sub-order Artiodactyla. The genus
Equufi, representing the Perissodactyla, probably disappeared as
a natural member of the fauna of western Europe during paleo-
lithic times. The Artiodactyla are at present the most widely
distributed and abundantly represented of the order. About
eight families, eighty genera and six hundred species are
currently recognized. Three families and nine genera are repre-
sented among the species now occurring in western Europe.

The Ungidata of Europe are so inadequately represented in the
British and other Mu.seums that it has been found impossible to
treat the group in the same manner as the preceding orders. No
general monographic study of any genus has been possible. The
descriptions have consequently been reduced to diagnosis, and
some of the recently named forms are entei'ed at second hand.

ARTIFICIAL KEY TO THE EUROPEAN FAMILIES OP
ABTIODACTYLA.

Upper incisors present ; lower canine not incisor-like ;

horns absent (Pigs) Suich-, p. 95G.

Upper incisors absent ; lower canine incisor-like ; horns

present, at least in males.

Horns, consisting of an outer sheath supported by a

bony core growing from the frontal bone, present

at all times in adult males and usually in females

(Cattle, sheep, goats, &c.) Bovidn; p. 9SG.

Horns (antlers), consisting of a solid bony outgrowth
from the frontal bone, periodically shed and re-
newed, often absent in females (Deer) Cervichr, p. 9G2.

956

UNGULATA

Family SUID^.

1821. SuidH' Gray, London IMed. Repos., xv, p. 30G, April 1, 1821.

Geograjpltical distributlou. — Wanner and temperate portions
of the Old World from Japan and the Philippines to Ireland
(now extinct in the British Islands), south throughout Africa
(including Madagascar) and in the Malay region to New Guinea.

Characters. — Snout elongated, the abruptly truncate muzzle
terminating in a vertically flattened, expanded pad svipported by
a well developed free ossicle (not shown in figs. 199 and 200);
incisors rooted, the upper teeth normally present though some-
times lost in old age ; upper canine curving outward and upward
at side of muzzle ; molars bunodont ; stomach essentially simple ;
feet with four well-developed toes ; no horns or antlers.

Remarks. — The family Suidse contains at least six well-
defined living genera. Only one of these, the typical Sua, occurs
in Europe.

Genus SUS Linno?us.

1758. Sus Linnteus, Syst. Nat., i, 10th ed., p. 49.

176G. Apcr Pallas, Miscellanea Zoologica, p. IG (Substitute for Sus).

1815. Apcr Rafinesque, Analyse de la Nature, p. 56 (Substitute for Sus).

1857. Sus Blasius, Siiugethiere Deutschlands, p. 508.

1868. Scrofa Gray, Proc. Zool. Soc, London, p. 38 (Domestic Pig).

Type species. — Sus scrofa Linna?us (by tautonymy).

GeograpMcal distrihution.- — Same as that of the family Suidse,
except that in Africa it is confined to the Mediterranean region.

Characters. — Dental formula : i |Ef, c JE^, pjm 4E5, m ^Eg = 44 ;
incisors and premolars tending to remain functional throughout
life, though the small i^, pvi^ and pru^ are not infrequently lost ;
bunodont structure of molars showing no special modification ;
canines small in females, moderately large and conspicuously
trenchant in males ; skull elongate, high and narrow, typically
suine and without con.spicuous modifications.

Bemai-Jcs. — The genus Sus as now understood contains about
thirty recognized living species, mostly of the Indo-Malayan
region. Two appear to be readily distinguished among the
members of the group occurring in western Europe.

KEY TO THE EUROPEAN SPECIES OF SUS.

Upper length of skull in adult males usually more

than 350 mm. ; vi^ with third transverse ridge

well developed (Distribution general) S. scrofa, p. 957.

Upper length of skull in adult males usually less

than 300 mm. ; 7n^ with third transverse ridge

obsolete (Sardinia) S. meridionalis, p. 960.

sus

957

sus SCEOFA Linn;eus.

1758. [Sus] scrofa Linnseus, Syst. Nat. i, 10th ed., p. 49.

1785. [Siis] setosus Boddaert, Elenchus Animalium, i, p. 157 (Substitute

for scrofa).
1785. [Sus setosus] a aper Boddaert, Elenchus Animalium, i, p. 157

(Special name for the wild boar).
1811. Sus europ^us Pallas, Zoogr. Rosso-Asiat., i, p. 265 (Substitute for

scrofa).
1857. Sus scrofa Blasius, Siiugethiere Deutschlands, p. 510.
1882. [Sus scrofa] var. ccltica Strobel, Atti Soc. Ital. Sci. Nat., IMilauo,

XXV, p. 79, June, 1882 (Renaming of true scrofa).
1910. Sus scrofa Trouessart, Faune, Mamm. d'Europe, p. 225.

Ti/pe lucalifi/. — Germany.

Geographical distribution. — Southei"n and central Europe,

XEBZ/-^

FIG. 109.
Sua scrofa. X l-

formerly west to Ireland and north to .southern Norway and
Sweden ; now restricted to that portion of the Continent lying

9n8

UNGULATA

south of the Baltic, and occurring in forested regions only ;
eastern limit of range unknown.

Diagnosis. — Size rather large, the length of head and body
about 1 • 5 m. in full-grown males, somewhat less in females,
upper length of skull in adult males about 350 mm., upper tooth-
row including canine 143-152 mm. ; posterior upper molar large,
its third transverse ridge evident ; general colour of adults
brown, in diifei'ent individuals tending to become blackish,
greyish or reddish, the nature of these variations not under-
s tood ; face, cheeks and throat grizzled by sprinkling of whitish

VlG. 200.
SuK xcrofa. X i.

hairs, but these not tending to produce definite markings ; new-
born young brown with conspicuous blackish stripes ; bristles
along median line of neck lengthened but not forming a con-
spicuous crest ; underfur everywhere dense and woolly.

Measurements. — Adult male from Province of Burgos, Spain :
head and body about 1300; tail, 170; longest hairs at tip of
tail, 200 ; hind foot (with hoof), 250 ; length of hind hoof along
inner border, 50 ; length of fore hoof along inner border, 70 ;
ear from ci'own, 115 ; width of ear, 90; length of hairs at upper
edge of ear', 30. For cranial measurements see Table, p. 961.

sus 959

Siiecimens examined. — Sixteen, from the followiDg localities: —
Prance : No exact locality, 1.

Germany : Harz Mountains, 1 ykull (tJ.S.N.M.) ; Waldloinigen,
Odenwald, N. Baden, 1 ; Wiirtemberg, 2 (skulls) ; no exact locality, 2.

A.:

Fig. 201.
Sus scro/a. Cheek teetli, iiat. sizf.

Italy: San Rossore, Tuscany, 3 (Turin).

Spain : Pinares de Quintanar, Burgos, 3 (B.M. and U.S.N.M.) ; Almonte,
Seville, 1 ; Goto Donana, Huelva, 2.

960

UNGULATA

Bemarks. — Though it seems probable tliat more than one
geographical race of wild boar may occur in western Europe the
material examined is entirely inadequate to the formation of any
definite opinion on the subject.

[Since this account of Sks scrofa was in type three new forms
have been recognized by Thomas : —

Sus attila Thomas, Abstr. Proc. Zool. Soc. London, Ko. 105,
p. 1, March 12, 1912. Type locality, Koloszvar, Austria-
Hungary. Distribution, Hungary and Russia. Much larger
than Sus scrofa (upper length of skull, 452 mm. ; height of
skull at occiput, mandible included, 271 mm.). Based on an
adult male, No. 12. 1, 23. 1, presented by Miss Sarolta von
AVertheimstein.

Sus scrofa castilianus Thomas, Abstr. Proc. Zool. 8oc. London,
No. 105, p. 1, March 12, 1912. Type locality. Province of
Burgos, Spain. Distribution, northern Spain. Smaller than
true Scrofa (upper length of skull, 353 mm. ; height at
occiput, 198).

Sus scrofa. hseticus Thomas, Abstr. Proc. Zool. Soc. London,
No. 105, p. 2, March 12, 1912. Type locality. Goto Donana,
Huelva, Spain. Distribution, southern Spain. Smaller than
castilianus (upper length of skull, 324 mm. ; height at occiput,
208).]

1 (juv.). France. Purchased (Lefebvre). 43. 12. 29. 12.

6. Waldleinigen, Oden- H.R.H. Grand Duke 92. 8. 3. 1.

wald, N. Baden, Louis of Hesse

Germany. (c & p).

6 skull. Wurtemberg. Dr. A. Giinther (c). 59. 9. 6. 100.

6 skeleton. Germany. Zoological Society's 713. m.

Museum.

6 skull. Germany. Purchased. 62. 3. 20. 6.

c5. Province of Burgos. Hon. N. C. Rothschild 11. 10. 5. 3.

(Rev. S. Gonzalez.) (p).

(Type of S. s. castilianus Thomas.)

2?. Quintanar de la Sierra, S. & N. Gonzalez (c). 8.7.7.32-33.

Burgos.

9. Almonte, Seville, Spain. Lord Lilford (p). 95.9.4.16.

(A. Ruiz.)

6. Goto Donana, Huelva. A, Chapman (c & p). 8. 3. 8. 12.

(Type of S. s. hutticus Thomas.)

6 skull. Goto Donana. A. Chapman (c & p). 8. 3. 8. 13.

SUS MERIDIONALIS Major.

1881. Sus scrofa meridionalis Major, Atti Soc. Tosc. Sci. Nat., Pisa, Proc,

Verb., Ill, p. 119 (Sardinia ; the wild pig). Fully described in
Vol. VI, fasc. 2, pp. 346-362, 1883.

1882. ? Sus scrofa var. sarclous Strobel, Atti Soc. Ital. Sci. Nat., Milano,

XXV, p. 221 (Sardinia ; the domestic pig).
1910. Sus scrofa sardous Trouessart, Fauna Mamm. d'Europe, p. 226 (the
wild pig).

Type locality. — Sardinia.
Geographical distrihulion. — Sardinia.

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962 UNGULATA

Dluijiums. — Like Sns scrofa but not attaining so great size,
the upper length- of skull in adult male about 300 mm., upper
tooth-row including canine 114 to 128 mm.; posterior molar
both above and below with third transverse ridge much reduced ;
colour in both adult and young essentially as in Sus scrofa.

Measureiuents. — For cranial measurements see Table, p. 961.

Specimens examined. — Five skulls (Genoa).

Bemarhs. — The wild boar of Sardinia appears to be a well-
defined local form.

Family CERVID^.

1820. Cervidfl' Gray, London Med. Repos., xv, p. 307, April 1, 1821.

Geographical distrihution.- — Essentially the entire mainland of
North and South America ; in the Old World from the Arctic
region south to extreme north-western Africa, the Malay
Archipelago and the Philippine Islands.

Characters. — Artiodactyles with frontal appendages usually
occurring in males and sometimes in females, always, when
present, in the form of solid, periodically shed and renewed
antlers growing from permanent bases or pedicles on the frontal
bones ; molars usually (always in European members of the
family) brachyodont ; lateral digits of both fore and hind feet
usually present.

Bemarlcs. — About sixteen genera of living deer are known, a
number which will probably be increased by more detailed study.
Three of these, Cervus, Alces and Bangifer, are common to the
northern portion of both Eastern and Western Hemispheres,
four or five are peculiar to America, and the rest are confined to
the Old World. Five occur in Europe.

KEY TO THE EUROPEAN GENERA OF CERVIDJE.

Vomer higb, dividing posterior nares into two chambers ;
no marked contrast in size between anterior pre-
molar and vi- ; widtb of upper molars about one-
fourtb that of palate ; antlers normally present
in both sexes, the base of pedicle situated con-
spicuously behind level of orbital cavity ; lateral
hoofs functional ; main hoofs very short and broad,
the outline of the entire sole subcircular ; muzzle
entirely hairy ; antlers beginning to develop within
a few weeks of birth (Reindeer) Bangifer, p. 979.

Vomer low, not dividing posterior nares into two
chambers; a marked contrast in size between
anterior premolar and m- ; width of upper molars
about one-third that of palate; antlers normally
■ present in males only, the base of pedicle extending
over posterior portion of orbital cavity ; lateral
hoofs not functional ; main hoofs narrow, the out-
line of the entire sole noticeably longer than wide ;
muzzle not entirely hairy ; antlers not beginning to
develop until about a year after birth.

cERVus 963

Rostrum lengthened and nasal shortened, so that
distance from front of nasal to front of pre-
maxillary is about equal to that from back of
nasal to back of occiput ; antlers conspicuously
palmate, the shaft nearly horizontal ; muzzle
hairy except for a small bare spot between nostrils ;

young not spotted (Elk) Alecs, p. 97G.

Rostrum and nasal normal, the distance from front
of nasal to front of premaxillary much less than
half that from back of nasal to back of occiput ;
antlers terete or slightly palmate, the shaft ascend-
ing ; muzzle naked ; young spotted with white.
Maxillary canines present in both sexes ; antlers
terete, spreading, the brow tine, bez tine and
trez tine usually present in fully developed

individuals (Red Deer) Cerviis, p. 963.

Maxillary canines absent, antlers either not terete
or not spreading, the complement of tines not
entire.
Auditory bullse evenly inflated, their surface with-
out conspicuous ridges ; antlers spreading,
without bez tine, the distal portion narrowly
palmate; tail well developed (Fallow deer)... Dama, -p. 970.
Auditory bulke collapsed, their surface con-
spicuously ridged ; antlers erect, terete, with-
out brow tine ; tail reduced to an incon-
spicuous papilla (Roe Deer) Capyeolus, -p. 913.

Genus CERVUS Linnajus.

1758. Cervus Linnreus, Syst. Nat., r, 10th ed., p. 60.

1827. Elaplms Hamilton Smith, Griffith's Guvier, Animal Kingdom, v,

p. 307 (Substitute for Cervus).
1857. Cervus Blasius, Saugethiere Deutschlands, p. 438 (part).
1899. Eucervus Acloque, Faune de France, IMamm., p. 71 (not of Gray,

1866). Substitute for Cervus.

Type species. — Cervus elaphus Linmeus (by tautonymy).

Geograpldcal distribution. — North temperate region of both
Hemispheres ; in the Old World west to the Atlantic coast of
Norway, the British Islands and north-western Africa.

Characters. — Plesiometacarpalian deer of large size with
narrow, elongate hoofs ; maxillary canines present in both sexes ;
lower incisors (fig. 203) distinctly though not extremely differen-
tiated in size and form ; lachrymal vacuity widely open, the pit
well developed, moderate ; vomer low posteriorly, showing no
tendency to divide the posterior nares into two chambers ;
antlers large, spreading, terete, not beginning to grow until
about a year after birth, when fully developed with five or more
tines including the brow tine ; base of pedicle extending
conspicuously over posterior portion of oi^ital cavity ; tail well
developed, moderate ; muzzle naked ; young spotted with white,
their colour very different from that of adults.

BemarJcs. — The genus Cervus as thus defined contains the
Red Deer group and its immediate allies. The exact limits of
the genus are not definitely known ; but there can be no doubt

3 Q 2

964

UNGULATA

that the name Cerviis is cvirrently appHed to several gi'oups
which shovild be recognized as distinct.

It is not possible to attemj^t any revision of the European
members of this genus, since the requisite material, if it exists,
cannot now be brought together in one place. So far as can be
judged from the few specimens seen and from the descriptions
recently published, there is a distinct small form confined to
Sardinia, and a wide ranging, rather plastic continental animal
occurring in the forested regions, where not exterminated, from
Spain and Ireland eastward, and from Scotland and west-central
Norway to the Mediterranean coast. Apparently the Spanish,
British and Norwegian forms are smaller than those inhabiting
central Europe, while the largest specimens occur in eastern
Hungary. Although there is no probability that this course is
final I am treating all the European forms as geographical races
of Cervns claplms.

CERVUS ELAPHUS Linnreus.
(Syuonymy under subspecies.)

Geogrnj)]nral distrihution. — Forested regions (where not ex-
terminated) from Spain and Ireland eastward, and from Sardinia

Fig. 202.
Cervus elaphns.

Xi.

OBRVUS 065

and the MediteiraucMU coast to northern Scotland and central
Sweden ; an isolated i*ace on the coast of central Norway ;
absent from the peninsula of Italy ; eastern limit of range not
known.

Diwpums. — Height at shoulder about 1370 mm. or less;
upper length of skull in adult male usually about 330 to 370 mm. ;
tail rather long, pointed ; ear more than half as long as head ;
general colour reddish brown in summer, greyish brown in winter,

KlG. -J 13.
Cervus claphus. Incisiform teeth. Nat. .si/e.

the miderparts always somewhat paler (drab), but never strongly
contrasted ; a dark dorsal line sometimes present ; no conspicuous
whitish markings except occasionally the speculum, which may
however be indistinct ; antlers terete throughout except for the
flattening which often occurs in region of " cup " in old individuals ;
in fidly developed antlers there are normally present a brow tine,
bez tine and more than 5 points.

CERVUS ELAPHUS OERMANicus Desmarest.

1822. Cervus elaphiis gcrmanicus Desmarest, Mammalogie, ii, p. i'6i

(Germany).
1822. ? Cervus elaphus albus Desmarest, IMammalogie, ii, p. 435 {nomcn

nudum).
1845. Cervus elaphus albifrons Reichcnbach, VoUstiindigste Naturgesch.

des In- und Auslauds, Siiugeth., iii, pi. iii bis, fig. 26 (no exact

locality; see p. 18).
1857. Cervus elaphus Blasius, Siiugethiere Deutschlands, p. 439.
1874. Cervus elaphus, varius Fitzinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, lxix, pt. i, p. 574 (Germany).
1874. Cervus elaphus, albus Fitzinger, Sitzungsber. kais. Akad. Wissenscb.

Wien, Math.-Naturwiss. Classe, lxix, pt. i, p. 575 (Germany).
1898. ? Cervus elaphus maral Lydekker, Deer of All Lands, p. 79 (part ;

specimens from the Galician Carpathians, Hungary).
1903. Cervus vulgaris Botezat, Morphol. Jahrb., xxxii, p. 115, November 17,

1903 (Renaming of elaphus).

96G UNGULATA

1903. ? C[crvus} vulgaris campcstris'Botcziii, ]\Iorpho]. Jahrb., xxxii, p. 154,
November 17, 1903 (Lowiaads and beech forests on lower slopes
of Carpathian IMountains, Bukowina, Austria-Hungary). Not
Ccrvus campestris F. Cuvier, 1817.

1903. ? C[crvus] vulgaris montanus Botezat, IMorphol. Jahrb., xxxii, p. 155,
November 17, 1903 (Carpathian ]Mountains, Bukowina, Austria-
Hungary).

1906. Ccrvus claplius germanicus Lonnberg, Arkiv fdr Zoologi, iii, No. 9,

p. 14, January 22, 1906.

1907. ? Ccrvus balticus Matschie, Das Waidwerk in Wort und Bild, xvi,

p. 186, March 15, 1907 (Forest near Liebemiihl, Ostpreussen,

Germany).
1907. ? Cervus albicus INIatschic, Das Waidwerk in Wort und Bild, xvi,

p. 186, March 15, 1907 (Muskau, Oberlausitz, Silesia, Cxermany).
1907. ? Ccrvus rlicnanus Matschie, Das Waidwerk in Wort und Bild, xvi,

p. 186, March 15,1907 (Viernheim, Hessen-Darmstadt, Germany).
1907. ? Cervus bajovaricus Matschie, Das Waidwerk in Wort und Bild,

XVI, p. i86, March 15, 1907 (Rohner, Konigssee, Oberbaycrn,

Germany).
1910. Ccrvus claplms gcrvianicus Trouessart, Faune Mamm. d'Europe,

p. 228.

Type locality. — Germany.

Geographical distribution. — Continental Europe, limits of range
not known ; animals of this general type occur from the Mediter-
ranean region to the Baltic and from western Fi-ance to eastern
Hungary, though they probably represent more than one geo-
graphical race.

. Diagnosis. — Size large, as in C. elaplms elaphus ; caudal disk
conspicuously lighter than flanks and usually with a definite
black or blackish border.

Measurouents. — For cranial measurements see Table, p. 982.

Specimens examined. — Nine, from the following localities : —

France : No exact locality, 2.

Germany : Gohrde, Hanover, 1 ; south Germany, 1 ; no exact locality, 1.

Austria-Hungary : Bohemia, 2 ; Carpathians, 1.

Italy : Campo Carnico, Cadore, 1 (Turin).

BemarJi-s. — The German Red Deer has been divided by
Matschie into the following species, based primarily on peculiarities
of the antlers. I am unable to express any opinion as to the
validity of these forms.

Cervus halticus.— Antlers with beam evenly and gradually
concave on inner border ; points of all tines directed inward.
Type locality : Liebemiihl, Ostpreussen. Additional specimens
mentioned from Auer, Ostpreussen, Cladow-West, Ostpreussen
(near the Baltic divide), Hartingswalde, Ostpreussen, Kommusin,
Ostpreussen, Ramuck, Ostpreussen, Schwalgendorf, Ostpreussen,
and Klein-Ottlau, Westpreussen.

Cervus albicus. — Antlers with beam bent abruptly inward at
level of trez tine ; points of inner tines directed inward, those of
outer prongs directed upward. Type locality : Muskau, Ober-

CERVUS 967

liuisit/,, Silesia. Range: middle Germany soulli of the Baltic
divide, west to the Weser divide, and south to the Main divide.

Cervus rlic^ianns. — Antlers with beam bent abruptly inward
at level of trez tine ; points of all tines directed upward. TyjM!
locality: Viernheim, Hessen - Darmstadt. Range: western
Germany from the Harz Mountains to the Danube basin.

Cervus hajonaricus. — Antlers as in C. rJiennnus, but with inner
tines directed inward and backward. Type locality : Rohner,
Konigssee, Oberbayern. Distribution : Oberbayern.

One or more forms of Red Deer probably distinct frcjm
C. daphus (jeriiKtnirits occur in eastern Hungary. They ha\e
been referred to C. elaplius maral (type locality, " the Persian
IVIountains ") by Lydekker, and have been described as new under
the names campi'stris and montanus by Bozetat.

? & yg. St. France. Turchasecl (Lefebvre). 43. 12. 29. 5 & 14.

6 frontlet Gohrde, Hanover. J. E. Hartiug (i-). 88. 6. 12. 1.

(hornless). [H.M. Emperor
William I.)
9 skull. S. Germany. Dr. A. Giinther (c). 59. 9. 6. 103.

6 antlers. Germany. 689. p.

2 S antlers. Bohemia. Col. J. Evans (p). 89. 11. 20. 1-2.

6 antlers. Carpathians. H.H. Prince Heinrich 96. 10. 10. 1.

von Liechtenstein (p).

Cervus elaphus elaphus Limiiwus.

1758. {Cervus] elaphus Linnscus, Syst. Nat., i, 10th ed., p. 07.
1906. Cervus elaplms Lonnberg, Arkiv for Zoologi, in, No. 9, p. 9,
January 22, 1906.

Tijpc Joealiti/. — Southern Sweden.

Gcoijraphlcal distribution. — ■" The [range] .... extended in
bygone days probably over the greater part of Cuitaland. At
present the red deer is in Sweden confined to southern Skania,
and there chiefly found on a few large estates, Hiickeberga,
Of \ edskloster, Borringe, Sofdeborg, Snogeholm, Skabersjo, etc.
The number is quite small, perhaps not more than about 100 in
all " (Lcinnberg).

Diagnosis. — Size large ; caudal disk not conspicuously lighter
than flanks and never with a definite black border.

licmarJi-s. — I have not seen this animal. From Lunnberg's
account it appears to be sufhciently different from the stag of
central Europe to merit recognition by name.

Cervus elaphus atlanticus Lonnberg.

1906. Cervus elaphus atlanticus Lonnberg, Arkiv for Zoologi, in, No. 9, p. 9,

January 22, 1906.
1910. Cervus elaphus atlanticus Trouessart, Faune j\Ianim. d'Europe, j). 228.

Twpe localiti/. — Hitteren Island, Trondhjem, Norway.
Geographical distribution. — West coast of Norway from
Stavanger Fjord north to about latitude 65°.

968 UNGULATA

Diagnosis. — Size smaller and colour paler than in true Cerviis
elaphus ; caudal disk with distinct dark border.

Measurements. — For cranial measurements see Table, p. 982.

Specimens examined. — Two, one from Frcdo, Norway, the other from
&loppen, Nordfjord, Norway (bofch U.S.N.M.).

BemarJcs.— Though sharply differentiated from the Swedish
and central European forms, the Norwegian red deer closely
resembles that of Scotland.''

Cervus elaphus scoticus Lonnberg.

1906. Cervus elaphus scoticus Lonnberg, Arkiv for Zoologie, iii. No. 9,

p. 11, January 22, 1906.
1910. Cervus elaphus scoticus Trouessart, Paune Mamm. d'Europe, p. 228.

Type localiii/. — Glenquoich Forest, Inverness-shire, Scotland.

Geographical distribution. — Great Britain ; limits of range
not known.

Characters. — Like Cervus elaphus atlantirus but colour darker
and less grey.

Measurements. — For cranial measurements see Table, p. 982.

Specimens examined. — Eighteen, from the following localities : Balma-
caan, Inverness, Scotland, 2 ; Island of Jura, Scotland, 5 ; Fort William,
Lochaber, Scotland, 1 ; Loch Sunart, Argyllshire, 1 ; Jura, Ardgour, Argyll-
shire, 2 ; Northumberland, 1 ; Woburn Abbey, Bedfordshire, 1 ; Exmoor,
Devon, 1; England, no exact locality, 4.

BemarTcs. — The status of the British form of Red Deer is
not well understood. The cranial characters mentioned by
Lonnberg, as distinguisbing the animal from atlanticus, are
inconstant, but there appears to be an appreciable difference
in colour between the two races.

6, 9. Balmacaan, Inver- Bradley Martin (r). 9. 1. 15. 1-2.

ness, Scotland.
2 c5 antlers. Jura, Ardgour, Ar- F. Hamilton-Leigh (p). 11.2.21. 1-2

gyllshire.
6 skull with- Loch Sunart, Argyll- Geu. Hamilton (p). 86. 6. 10. 1.
out antlers. shire.

9. Fort William. W. Jones (p). 8. 2. 10. 1.

3 skulls and Island of Jura. Henry Evans (p). 96. 12. 21. 1-5.

2 pairs of
antlers.
<$ stuffed. Alnwick Chase, Duke of Northumber- 63.11.16.5.

Northumberland. land (p).
6 stuffed. Woburn, Bedford- Duke of Bedford (p). 97. 4. 3. 3.
shire.

* For discussion of the meaning of this resemblance see Stejneger,
Smithsonian Miscellaneous Collections, xlviii, pp. 162-469, May 4, 1907,
and Bergens Museums Aarbog, siv, 1908.

CERVUS

969

9 skuU. Exmoor, Devon. R. A. Sanders (p). 6. 2. 26. 1.

i antlers. England. 689. b.

<5 antlers. England. Purchased (Lead- 46.11.20.17.

beater).

6 skull. England. Purchased (Baker). 47. 12. 11. 16.

6 antlers. (From a peat-bog.) Jabez Allies (p). 49. 3. 6. 1.

Cervus elaphus hispanicus Hilzheimer.

1909. C[crt'7/s] claj^hus hisjmnicns Hilzheimer, Archiv fiir Rasseu- und

Gesellschafts-Biologie, 1909, p. 313. (South-western Spain?)

Type in Stuttgart Museum.
1911. ? Cervus cla2>hus bolivari Cabrera, Bol. Real Soc. Espan. Hist.

Nat. XI, p. 559, December, 1911 (El Pardo, Madrid, Spain). Type

in Madrid Museum.

Ti/jje locality. — Spain, exact locality not known.*

Geographical (listrihution. — Iberian Peninsula, limits of range
unknown.

Diagnosis. — Size apparently less than in Cervus elaphus
scoticus ; colour decidedly more greyish ; skull narrower,
particularly in interorbital and palatal regions.

Measurements. — For cranial measurements see Table, p. 982.

Bemarks. — The Red Deer of Spain are divided by Cabrera
into a smaller southern race (Impanicus) and a larger central
and northern race (bolivari). At present the evidence seems
inconclusive, though the cranial measurements (p. 982) tend
to indicate the presence of two forms.

Specimens examined. — Four, from Goto Donana, Huelva, Spain, and
two from Pinares de Quintanar, Burgos, Spain (U.S.N.M.).

6 9. Goto Donana, Huelva, Lord Lilford (p). 95. 9. 4. 14-15.

Spain. {A. Ruiz).
6. Goto Donana. A. Chapman (c & p). 8. 3. 8. 14.

6 skull. Goto Donana. A. Chapman (c & p). 8. 3. 8. 15.

Cervus elaphus corsicanus Erxleben.

1777. [Cervus'] corsicanus Erxleben, Syst. Regni Anim., i, p. 304 (Corsica).
1822. Clcrvus] mediterrancus Blainville, Journ. de Phys. Chem. Hist.

Nat., xciv, p. 262 (Corsica).
1855. \_Cervus elaphus'] j3 minor Wagner, Schreber's Saugth., Suppl., v,

p. 854 (substitute for corsicanus and mediterrancus).
1910. Cervus elaphus corsicanus Trouessart, Fauue Mamm. d'Europe,

p. 229.

Type locality. — Corsica.

Geographical distribution. — Corsicaf and Sardinia.

Diagnosis. — Size about as in Cervus elaphus hispanicus (height

* Dr. Hilzheimer informs mo that the only clue to the history of the
type is the fact that a pair of roebuck antlers in the Stuttgart collection
was taken by the same collector in Estremadura.

t Polybius (xii, cap. in) states that the stag is not native to Corsica.
(The Histories of Polybius translated by Evelyn S. Shuckburgh, ii, p. 80).

970 UNGULATA

at .shoulder of mounted uuile in Turin ]\[useuui iSOO nnn.), but
genei'fil colour darker than in any of the small continental forms.
Measurements. — For cranial measurements see Table, p. 984.

Specimens examined. — Four, all from Corsica (Turin).

Genus DAMA Hamilton Smith.

1827. Datna Hamilton Smith, Griffith's Cuvier, Anim. Kingd., v, p. 80G.

1844. Platyccros Wagner, Schreber's Siiugth., Suppl., iv, p. 347.

1855. Dactyloccros Wagner, Schreber's Siiugth., Suppl. v, p. 352 (Substitute

for Dama and Plafyceros).
1857. Ccrvus Blasius, Saugethiere Deutschlands, p. 39 (part).
1893. Machlis Zittel, Handb. Palaeont. iv, p. 402 (published as a synonym

of Dama with Kaup as authority).
1898. Pahnatus Lydeiiker, Deer of all Lands, p. 125. Wrongly attributed

to Giebel, Saugethiere, p. 351 (a group name used in the plural :

" Palniati mit schaufelformigem Gewieh ").

Type species. — Cervus dama Linnseus.

Geographical distribution. — Mediterranean region of southern
Europe and western Asia ; occurs in a condition of semi-
domestication as far north as the British Islands and southern
Scandinavia.

Characters. — Like Cervus but with skull .shorter and relati^'ely
much broader, orbits larger, maxillary canines absent, cheek-
teeth, particularly vi", more brachydont, lower incisors (fig. 205)
excessively differentiated in both size and form ; antlers with
brow tine and trez tine present, and a distinct though narrow
palmation in region of surroyals.*

Bemarlcs. — The Fallow Deer form a sharply defined group
generically distinct from Cervus. Two species are known, one of
which occurs in Europe.

DAMA DAMA Linmvus.

1758. [Cerrtts] dama Linnaeus, Syst. Nat., i, 10th ed., p. 67.

1798. Ccrvus platyceros Cuvier, Tabl. Klem. de I'Hist. Nat. des Anim.,

p. 160 (Renaming of dama).
1816. L (sic) [ervus} mauricus F. Cuvier, Bull, des Sci. Soc. Philomath.,

1816, p. 72 (locality not known).

* The extinct genus Alec Blumenbach (Handb. d. Naturgesch., 6th ed.,
p. 697, 1799, type Alee gigantcus Blumenbach from the peat bogs of
Ireland = " C(?rf?<s megaceros" Auct. or " Megaceros hibcrnicus'' Auct.),
though usually regarded as related to Dama, or even as identical with it
(see Lydekker, Deer of all Lands, pp. 126-127, 1898, for statement of
this extremely radical view), differs widely from all known genera of recent
deer iu the combination of plesiometacarpalian foot, high vomer nearly
dividing posterior nares into two cavities, absence of maxillary canines and
excessively reduced lachrymal vacuity (see Lonnberg, Arkiv for Zoologi,
III, No. 14, August, 1906).

971

Akad.
p. 555

Akad.
p. 555

1829. [Ccrvtis dama] 13. Icuavthiops Fischer, Syiiops. j\Iamm., p. 448

(Germany).
1829. \_Ccrvus dama] y. maura Fischer, Syuops. Mamm., p. 448 (Renaming

of maiiriciis).
1843. Dama vtdgaris Gray, List Spec. Mamm. Brit. IMus., p. 181

(Renaming of dama).
1857. Ccrv2is dama Blasius, Saugethiero Deutschlands, p. 453.
1874. Damaplafi/ccros,nigcrFitzmger,^it'^ung&her:. kais. Akad. Wissensch.

Wien, ]Math.-Naturvviss. Classe, lxix, pt. i, p. 553 (Renaming of

mauricus) .
1874. Dama platyceros, varius Fitzinger, Sitzungsher. kais.

Wissensch. Wien, Math.-Naturwiss. Classe, lxix, pt. i,

(Germany).
1874. Dama i-tlatyceros, alhus Fitzinger, Sitzungsher. kais.

Wissensch. Wien, Math.-Naturwiss. Classe, lxix, pt. i,

(Renaming of Iciiaithiops).
1885. \_Dama'] dama Lataste, Actes Soc. Linn, de Bordeaux, xxxix, p. 288.
1910. Dama dama Trouessart, Faune Mamm. d'Europe, p. 229.

Type loeaUii/. — Sweden (introduced).

Geographical distrilmtion. — Mediterranean region of Europe
and Asia Minor ; semi-domesticated in central Europe as far
north as England and southern Sweden.

Diagnosis. — Height at shoulder about 1 mm. ; back and sides
in summer fawn colour with numerous
whitish spots, in winter uniform greyish
(melanism and albinism not infre-
quent) ; underparts and inner surface
of ear whitish at all seasons ; blackish
dorsal line continued to end of tail ;
spots at each side of dorsal line
distinct, not tending to become con-
riuent ; antlers with brow tine large,

Dama damn. X \.

Dama davia. Incisifonn teeth.
Nat. size.

and with I'ogion of trez tine showing no tendency to become
flattened.

Measurements. — Adult male: head and body, 15-10; tail,

972

UNCtULATA

190 ; pencil, 100 ; hind foot (with hoof), 4.35 ; (!;ii- from cruwii,
165 ; width of ear at middle, 90. For cranial measurements see
Table, p. 983.

Specimens examined. — Sixty-seven, from the following localities: —
England : As enumerated below, 62.
Sweden : No exact locality, 1 (U.S.N.M.).
Germany : No exact locality, 1.
Locality unknown : Three (U.S.N.M.)

6 St. Woburu, Bedford- Duke of Bedford (r). 9G. 9. 24. 1.

shire, England.

6 St. Tring Park, Hert- Hon. Walter Eoths- 98. 10. 18. 1.

fordshire. child (p).

6 st. England. Purchased (Lead-

beater).

6 St. England. Purchased (Baker). 46. 10. 23. 11.

2 6 St. England. No history.

3 pairs antlers. England. 693. a. b. d.

antlers. NannanPark, J. E. Harting (p). 88.6.12.3.

Merionethshire.

46 antlers. New Forest. Mrs. Smyth (p). 50. 2. 5. 1-46.

6 skull. England. Purchased. 50. 11. 13. 15.

9 skull. England. 693. k.

2 (5 2 ? skuUs. Lidth de Jeude Coll. 67. 4. 12. 234,

235, 236, 241.

6 St. Italy. Purchased (Lefebvrc). 43. 12. 29. 4.

Genus CAPREOLUS Gray.

1821. Capreohis Gray, London ]Med. Repos., xv, p. 307, April 1, 1821.
1837. Caprca Ogilby, Proc. Zool. Soc. London, 1836, p. 135, June 27, 1837.
1857. Capreolus Blasius, Saugethiere Deutschlands, p. 457.

Type apecies. — Cerims capreolus Linnaeus.

Geographical distribution. — Central and southern Europe from
Great Britain and southern Scandinavia eastward ; in Asia east
to the Pacific coast.

Characters. — Telemetacarpalian deer of small size with narrow,
elongate hoofs, maxillary canines normally absent,* lower incisors
distinctly though not excessively difFerentiiated in size and form,
lachrymal vacuity large, the pit reduced to a shallow, inconspicuous
depression ; vomer low posteriorly, showing no tendency to divide
the posterior nares into two chambers ; antler small,! lirst appear-
ing about a year after birth, erect, terete, without brow tine,
the prongs normally three, the pedicle appearing to arise further
back than in Cervus, but its anterior base extending over orbital
cavity ; tail reduced to an inconspicuous papilla ; muzzle naked •
young with .spotted coat A'ery different from pelage of adult.

* For account of their occasional occurrence see Kolliker, Wiirzburger
Naturwiss. Zeitschr., vi, p. 82, 1866.

t Sometimes present in females (see Sherrcu, Field, London, cxiv,
p. 751, October 23, 1909).

CAPREOLUS

973

Memarks. — About a dozen forms of Capreolus have been
described. One species, represented by four geographical races,
occurs in Europe.

CAPREOLUS CAPREOLUS Linnoeus.
(Synonymy under subspecies.)

Geograpltical (lisfrilmtion. — From Great Britain eastward into
Asia (eastern limit of range not known), and from the Mediter-
ranean coast north to Scotland and central Sweden.

Diagnosis. — Size small (condylobasal length of skull in adult
male about 150 to 160 mm.); ear slender, uniform greyish or
reddish on outer side ; antlers not nor-
mally attaining special massiveness ;
teeth strictly brachyodont, the height of
crown of middle upper premolar less
than diameter of crown in line of
tooth-row ; summer and winter pelages
strikingly different, the former reddish,

Fig. 20G. Fig. 207.

Capreolv^ capreolus. Capreolm capreolus.

X 1 . Incisiform teeth. Nat. size.

the latter grey with well developed white speculum ; lips and
sides of muzzle always blackish, the chin and front of muzzle
at each side of naked area white in con.spicuous contrast.

Memarls. — In westei'n Europe Capreohs capreolus is repre-
sented by four well-defined local races,

KEY TO THE EUROPEAN FORMS OF CAPREOLUS
CAPREOLUS.

General colour of face distinctly darker than that

of body (Great Britain) C. caim'olw; thotu, p. 975.

General colour of face not darker tban that of body.

Pale throat patcb and neck patch in winter

pelage sharply defined whitish (South-eastern

Europe) C'. capreohm transsylmni-

cux, p. 975.
Pale throat patch and neck patch in winter
pelage obscure, yellowish or greyish.
General colour of winter pelage decidedly

yellowish (Southern Scandinavia) C. caprcolux capreolux,

p. 974.
General colour of winter pelage a coarsely
grizzled grey without decided yellowish
tinge (Spain) C. capreoluf: cami'.', t^. 975,

974

UNGULATA

1758.

1792.
1829.
1843.
1844,
1845,

1857.
1860,

1874.

1874.

1910.

Capreolus "capreolus capreolus Linnoeus.

[Ccrvus] capreolus Linnaeus, Syst. Nat. i, 10th ed., p. 08 (Sweden).
C[ervus\ capreolus albus Kerr, Anim. Kingd., p. 302 (Franche Comt6,

France).
? Capreolus dorcas Burnett, Quart. Journ. Sci. Lit. Art, 1829, pt. 2,

p. 353 (nomen nudum).
Capreolus caprwa Gray, List. Spec. Mamm. Brit. Mus., p. 17G

(Renaming of capreolus).
C[apreolus'] europrrus Sundevall, Ofversigt af Kongl. Vetensk.-Akad.

Fiirhandl., Stockholm, i, p. 184 (Renaming of capreolus).
C[ervus'] capreolus plumbeus Reichenbach, Vollstandigste Natur-

gesch. der In- und Auslands, Saugeth., iii, pi. iii bis, fig. 53

(Germany).
Capreolus capreolus Blasius, Saugethiere Deutschlands, p. 457.
Capreolus vulgaris Fitzinger, Wissensch.-pop. Naturgesch. der
192 (Renaming of capreolus).
niger Fitzinger, Sitzungsber. kais. Akad.
Math.-Naturwiss. Classe, lxx, Abth. i, p. 247

Saugethiere, iv, p.
Capreolus vulgaris,

Wissensch. Wien

(Germany)
Capreolus viilgaris, varius Fitzinger, Sitzungsber. kais. Akad,

Wissensch. Wien, Math.-Naturwiss. Classe, lxx, Abth. i, p. 247

(Germany).
Capreolus capreolus Trouessart, Faune Mamm. d'Europe, p. 238.

Tijpe localiti/. — Sweden.

Geographical distribution. — Originally confined to southern
Sweden but now somewhat extended artificially to the north
and west.

Diagnosis. — Throat patch indistinct ; general colour in winter
pelage grey decidedly tinged with yellow.

Measurements. — For cranial measurements see Table, p. 983,

c5st.

France.

2 <5, 1 ?.

Ferrieres,

Seine-

Hon. N. C. Roths-

10. 11. 17. 1.*

et-Marne.

child (p).

11. 11. 18. 1.*

12. 1. 17. 1.*

c?.

Armandvilliers,

Hon. N. C. Roths-

11. 12. 5. 1.*

Seine-et-Marne.

child (p).

9.

Skabersjo, Sweden.

Dr. Einar Lonn-

10. 10. 18. 1.

berg (p).

S st. (melano).

Westphalia.

Purchased

76. 5. 4. 1-3.*

9, 1 separate

(Gerrard).

skull.

3 skulls.

S. Germany

Dr.A. Giinther(c).

59.9.6.107-109.*

15 pairs of

Bavaria.

F. N. A. Fleisch-

11. 9. 13. 1-15.*

antlers.

mann (p).

6 head st.

Austria.

Col. R. W. Shipway

98. 10. 12. 1.*

2 9.

Csehtelek,

Bihar

Hon. IMrs. Charles

10. 12. 4. 1.*

Comitat,

Hun-

Rothschild (p).

11. 1. 14. 11.*

7 skulls.

gary.

Lidth de Jeude

67. 4. 12. 225-

Collection.

231.*

* Not determined by the author. — 0. T.

CAPREOLUS 975

Capreolus CAPREOLUS TRANSSYLVANicus Matschie.

1907. Capreolus transsylvanicus Matschie, Das Weidwerk in Wort und
Bild, XVI, p. 224, March 15, 1907.

Type locality. — Bana, Roumania.

Geographical distribution. — Eastern Europe, westward to th(^
Italian Alps.

Diagnosis. — Pale throat patch sharply defined ; general colour
in winter pelage clear grizzled grey.

Measurements. — For cranial measurements see Table, p. 984.

Specimens examined. — Four, from Padola, Cadore, Italy (B.M. and
U.S.N.M.).

Remarhs. — In its decidedly grey colour this form differs
noticeably from true capreolus and agrees with the description
and figure of transsylvanicus. It is distinguishable from the
equally grey ccunis by the sharply defined white throat patch.

2 9 Padola, Cadore, Venetian Alps. Turin ]\Iuseum (e.) 9. 1. 18. 3-4.

Capreolus capreolus canus Miller.

1910. Capreolus capreolus canus Miller, Ann. and Mag. Nat. Hist., 8th ser.,
VI, p. 460, November, 1910. Type in British Museum.

Type locality. — Quintanar de la Sierra, Burgos, Spain.

Geographical clistrihution. — Iberian Peninsula.

Diagnosis. — Like C. capreolus capreolus in respect to the
obscurely defined pale neck patch, but general colour in winter
pelage a coarsely grizzled grey without decided yellowish tinge.

Measurements. — Type and a second adult male from the type
locality : head and body, 1220 and 970 ; hind foot, 305 and
330; hind foot including hoof, 355 and 370; ear, 120 and — .
For cranial measurements see Table, p. 983.

Specimens examined. — Eight, all from Pinares de Quintanar de la
Sierra, Burgos, Spain (B.M. and U.S.N.M.).

3 (J, 2 ?. Pinares de Quintanar S. & N. Gonzalez (c). 8. 7. 7. 27-31.
de la Sierra, Burgos, (8. 7. 7. 28. Type of subspecies.)

Spain.

Capreolus capreolus tiiotti Lunnbei-g.

1910. Capreolus capreolus tJiotti Liinnberg, Ann. and Mag. Nat. Hist.,
8th sor., VI, p. 297, September, 1910. Type in British Museum.

Type locality. — Arndilly, Craig EUachie, Morayshire, Scot-
land*.

Geographical distribution. — Great Britain.

* The locality of the type specimen was accidentally recorded as
Aberfeldy in the original description.- 0. T.

976

UNGULATA

Diagnosis. — Differs from C. cap-eolus caj)reoluft in general
darker colour, this particularly noticeable in the face, which is
darker than body, a peculiarity not occurring in other European
races.

Measurements. — Head and body (of type), 1150 mm.; hind
foot, 380; ear, 127. For cranial measurements see Table, p. 983.

S head st.

6, ?.

S skeleton.

? skull.

, 6, 9.

S, 9 heads st.
6,9.

6 St.
Frontlet and

antlers.
2 frontlets
and antlers.

6 St.

6 St.

Nairn, Scotland.

Nairn.

Nairn.

Nairn.

Arndilly, Craig Ella-
chie, Morayshire.

Poltalloch, Argyll-
shire.

Thornhill, Dum-
friesshire.

Scotland.

Scotland.

Scotland.

Earl Cawdor (p). 93. 1. 3. 1.

Earl Cawdor (p). 8. 8. 18. 1-2.

Earl- Cawdor (p). 85.10.6.1.

Earl Cawdor (p). 85. 10. 6. 2.

W. S. Menzies (p). 8. 11. 22. 1-2.

(8. 11. 22. 1. Ty]je of subspecies.)

Col. E. D. Malcolm 7. 6. 6. 1-2.

H. S. Gladstone (p). 11. 2. 22. 1-2.

Earl of Derby (p).
Gen. Hardwicke (p).

GO. k.
G88. d.

G88. a-b.

Purchased (R. Ward). 97. 12. 11. 2.
Whatcombe, Bland- J. C. Mansel Pleydell 97. 8. 21. 1.
ford, Dorset. (p).

Genus ALCES Gray.

1821. Alces Gray, London Med. R«po3., xv, p. 807, April 1, 1821.

1841. Alcelaphus Gloger, Hand- u. Hilfsbuch der Naturgesch, i, p. 143

(Substitute for Alces). Not of Blainville, 1816,
1857. Alces Blasius, Saugethiere Deutschlands, p. 434.
1902. Paralces Allen, Bull. Amer. Mus. Nat. Hist., xvi, p. IGO, July 1,

1902 (Substitute for .4 Zees assumed to be a homonym of Alee

Blumenbach, 1799).

Ti/pe species. — Cervus alces Linnpeus.

Geographical distribution. — Northern forested portions of both
hemispheres ; in the Old World west to Norway and south to
eastern Germany.

Characters. — Telemetacarpalian deer of largest size with
narrow, elongate hoofs, maxillary canines absent in both sexes,*
lower incisors (fig. 209) scarcely differentiated in form and not
much contrasted in size, lachrymal vacuity widely open, the
pit well developed, small, vomer low posteriorly, showing no
tendency to divide the posterior nares into two cavities, pre-
maxillary region of skull greatly lengthened and nasal region
shortened, so that distance from front of nasal to front of pre-
maxillary is about equal to that from back of nasal to back of
occiput ; antlers conspicuously palmate, present in male only ;

* Sometimes present in a rudimentary condition (see Lonnberg, Zool.
Anzeiger, xxviii, pp. 448-449, January 17, 1905).

977

Fig. :iOti.
Alces alces. X i.

3 R

978

UNGDLATA

general form heavy, the shoulders high ; upper lip conspicuously
produced in front ; muzzle hairy except for a very small median
bare spot ; throat of male with pendant flap of skin ; young not
spotted with white, their colour essentially like that of adults.

Remarls. — The genus Alces contains four currently recognized
species or geographical forms, two in America and two in the
Old World. Only one occurs in Eurojje.

ALCES ALCES Linnseus.

1758. [Cervus] alces Linnaeus, Syst. Nat., i, 10th ed., p. 66.

1829. ? Alces europ<-vus Burnett, Quart. Journ. Sci. Lit. Art, 1829, 353

(nonien Jiudum).
1836. Alces machlis Ogilby, Proc. Zool. Soc. London, p. 135 (Renaming of

alces).

1842. Alces antiquorum Riippell, Museum Senckeubergianum, iii, p. 183

(Renaming of alces).

1843. Alces palmahis Gray, List Spec. Mamm. Brit. Mus., p. 182 (Re-

naming of alces).
1857. Alces palviatus Blasius, Stiugethiere Deutschlands, p. 434.
1860. Alces jubata Fitzinger, Wisseuch.-pop. Naturgesch. der Saugethiere,

IV, p. 86 (Renaming of alces).
1898. Alces alces Lydekker, Deer of all Lands, p. 54.
1910. Alces alces Trouessart, Faune Mamm. d'Europe, p. 270.

Geogra'pliical distribution. — Forested portions of northern and
central Europe ; now confined, in the region west of Russia, to the
wilder parts of the Scandinavian Peninsula and eastern Germany.

Alces alces.

Fig. 209.
Incisiforni teeth.

Nat. size.

Diagnosis. — General characters as in the genus Alces ; height
at shoulder about two meters ; general colour brown, lighter and

RANGIFEK 979

approaching wood-brown on head, back, and legs, darker, ahnost
blackish on tail, throat, and underside of body.

Measurements. — Adult male from Stenkjix'r, Trondhjem,
Norway : head and body about 2900 ; tail, 95 ; hind foot, 855 ;
ear, 330 ; height at shoulder, 1900. For cranial measui'ements
see Table, p. 984.

Specimens examined. — Twelve, from the following localities : —
Norway: Stenkjrer, Trondhjem, Norway, 1 (U.S.N.M.). No exact
locality, 1 (U.S.N.M.).

Sweden : No exact locality, 9 (B.M. and U.S.N.M.). Udoholm 1.

Single antler. Udoholm, Sweden. Earl of Selkirk (p). 703. d.

6 antlers. Sweden. College of Surgeons (p). 703. e.

(From the Leverian Museum.)
7 pairs of Sweden. Purchased. 3. 11. 21. 1-7.

antlers.

Genus RANGIFER Hamilton Smith.

1827. Rangifer Hamilton Smith, Griffith's Cuvier, Animal Kingdom, v,

p. 304.
1827. Tarandus Billberg, Synopsis Faunae Scandinavise, i, p. 22.
1838. Procerus Serres, Essai sur les Cavernes a Ossements, 3rd ed., p. 143.
1840. Procervus Blainville, Comptes Rendus, Acad. Sci., Paris, xi, p. 392

(Substitute for Procerus).
1845. Achlis Reichenbach, VoUstandigste Naturgesch. des In- und Auslands,

Saugeth., in, p. 12 (Alternative for Tarandus).

Type species. — Cervus tarandus Linnaeus.

Geographical distribution. — Northern forests and barren
grounds of both hemispheres ; in the Old World north to
Spitzbergen ; Novaya Zemlaya 1

Characters. — Telemetacarpalian deer of medium size, with
lateral hoofs functional and main hoofs so broadened that the
outline of the two together is nearly circular, maxillary canine
present in both sexes; lower incisors (fig. 211) relatively smaller
than in other recent deer, slightly differentiated in both size and
form ; lachrymal vacuity large, the pit shallow and ill-defined ;
vomer high posteriorly, completely dividing posterior nares into
two chambers ; skull normal in general form, but orbital
cavities not pushing back under bases of horn pedicles ; antlers
usually present in both sexes, beginning to develop within four
or five weeks after birth, the beam slender and curved, slightly
palmate distally ; muzzle entirely hairy ; young not spotted with
white, their colour essentially like that of adults.

Bemarhs. — About sixteen forms of living Reindeer have been
described, most of them from North America. At least three
occur in western Europe or have done so within very recent
times.

3 R 2

980

UNGULATA

KEY TO THE EUROPEAN SPECIES OP RANGIFER.

Size small, upper length of skull less than

225 mm. (Spitzbergen) R. platyrhynclms, p.

Size large, upper length of skull about 270-

300 mm.

Upper length of skull in adult male 270-

290 mm. ; upper tooth-row about 95 mm.

(treeless alpine portions of Norway and

formerly of Sweden, now rare except in

captivity) R. tarandus, p. 980.

Upper length of skull in adult male about
300 mm. ; upper tooth-row about 85 mm.
(wooded portions of northern Pinnland
and formerly of northern Sweden) R. fennicus, jj. 981.

985.

RANGIFER TARANDUS Linnteus.

1758. [Cervus] tarandus Linnteus, Syst. Nat., i, 10th ed., p. 67.

1789. \_Cervus tarandits] a rangifer Gmelin, Syst. Nat., i, 13th ed., p. 177.

1827. Tarandtis lapiMnum Billberg, Synopsis Fatmse Scandinavise, i, p. 22

(Renaming of C. tarandus).
1842. Tarafulus horealis Riippell, Museum Senckenbergianum, iii, p, 183

(Renaming of C. tarandus).

Fig. 2J0.
Rangifer tarandux. x i.

981

1852. Tarandus furcifer Baird, Rep. Comm. Patents, 1851, ii, Agric.,p. 109

(Renaming of tarandus).
1902. [^Rangifcr tarandus] var. cylindricornis Camerano, Mem. Reale Accad.

Sci. Torino, 2nd ser., li, p. 167 (Renaming of true tarandus).
1910. Rangifer tarandus Trouessart, Faune Mamm. d'Europe, p. 231.

Type locality. — Alpine region of Swedish Lapland.

Geographical distrihution. — Formerly the entire alpine region
of the Scandinavian Peninsula ; now confined in the wild state
to two widely separated districts in Norway ; west Finmarken in
the north, and the main high mountain
region in the south. Extensively domes-
ticated throughout Scandinavian Lapland.

Diagnosis. — Size rather large (height
of adult male at shoulder about 1150 mm. ;
upper length of skull 270-290 mm.) ;
skull Avith nasal bones broad and little
arched ; teeth relatively large (upper tooth-
row 94-98 ; lower tooth-row 101-104 mm.) ;
general colour a greyish or drab brown
above, buffy whitish beneath and on
muzzle ; a darker longitudinal area on
side of body ; tail buffy white with dark
median line.

Measurements.^A.Avi\t male from Heimdalen, Norway : head
and body, 2000±, tail, 150 ; hind foot, 450± ; ear, 100±. For
cranial measurements see Table, p. 984.

Specimens exa.mined. — Nine, from the following localities : —
Norway : Heimdalen, Norway, 1 (U.S.N.M.), wild ; no exact locality,
1 (U.S.N.M.), tame ; B.M. specimens as enumerated below.

BemnrJcs. — The material examined is too limited to form the
basis of any discussion of the various forms of wild and tame
reindeer.

Fig. 211.
Ramjifer tarandus. In-
cisifoi'm teeth. Nat. size.

(5 St.

(J & 9 antlers.

(5 St.
& skeleton.

9 St.

& skeleton.

9.

9 (ad. & yg.).

LoQrdalMts.,Sogne Sir "W. J. Ingram 87.9.20.1.
Fjord, Norway. (c & p).

Sir W. J. Ingram 87. 9. 20. 2-3.

(c & p).
Chas. Ingram (c & p). 79. 10. 9. 1.

Loerdal Mts
Fillefjeld, Norway
Fillefjeld, Norway

Norway.
Norway.

Sir W. J. Ingram 81. 9. 28. 2.

(c & p).

Chas. Ingram (c & p). 75. 10. 30. 1.

J. C. Ingram (c & p). 83. 7. 28. 1-2.

RANGIFER FENNICUS Lonnberg.

1909. Rangifer tarandus fennicus Lijnnberg, Arkiv for Zoologi, vi, No. 4,
p. 10, July 14, 1909.

Type locality. — Torne, Lappmark (" thus probably in Enon-
teY\^,'^ fide Lonnberg), Finnland.

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RANGIFER 985

GeograjjJiical distribution. — Now probably confined to the
wooded portions of Finland, east to the Kola Peninsula, and
nearly extinct. Formerly ranging westward into the wooded
portion of northern Sweden where it is now probably represented
by the large woodland race of tame reindeer found in certain
districts.

Diagnosis. — Size greater than in Hangifer tarandus (height of

adult male at shoulder about 1200 mm. ; upper length of skull

'about 300 mm.); skull with nasal bones narrow and highly

arched ; teeth relatively small (upper tooth-row about 85 mm. ;

lower tooth-row about 90 mm.).

Measurements. — For cranial measurements see Table opposite.

Remarhs. — This animal, which I know only from Lonnberg's
description and figures, appears to be specifically distinct from
Rang if er tarandus.

RANGIFER PLATYRHYNCHUS Vrolik.

1829. Cervus (Tarandus) platyrhynchus Vrolik, Nieuwe Verliandcl. van 't

Kroninke Nederl. Inst., Eerste Klasse, pt. 2, p. 160 (Spitzbergen,

see Vrolik, I.e., p. 239).
1862. Cerviis tarandus forma spetsbergcnsis Andersen, Ofversigt af Kongl.

Vetensk.-Akad. Forhandl., Stockholm, xix, p. 457, October 8, 1862.
1866. [Rangifer arcticus] var. spitzhergcnsis Murray, Gcogr. Distrib.

Mamm., p. 334, described on p. 155.
1902. Rangifcr spitzbcrgensis Camerano, Mem. Realc Accad. Sci. Torino,

2d ser., li, pp. 159-240, 1902.
1910. Eangifer spitzbergejisis Trouessart, Faune Mamm. d'Europe, p. 232.

Type locality. — Spitzbergen.

GevgrajjMcal distrihution. — Spitzbergen.

Diagnosis. — Size much less than in Rangifer tarandus and
R. fennicus (height at shoulder of mounted adult male in Turin
Museum, 680 mm. ; upper length of skull less than 225 mm.) ;
skull with nasal bones broad and little arched, more widened
anteriorly than in the continental species ; teeth relatively as
large as in R. tarandus.

Measurements. — For cranial measurements see Table opposite.

Specimens examined. — Two mounted individuals and six skulls
(U.S.N.M. and Turin).

RemarJis. — ^The Spitzbergen Reindeer is sharply and com-
pletely differentiated from both of the species occurring in
continental Europe.

9(?6 ungulata

Family BOVID^.

1821. Bovich- Gray, London Med. Repos., xv, p. 308, April 1, 1821.

Geocjrajili'tcal distribution. — Africa ; southern Europe (within
historic times north to the Baltic and west to Great Britain) ;
Asia, including Japan and the larger Malay Islands ; North
America from Greenland to northern Mexico (recently exter-
minated over much of its former range).

Characters. — Artiodactyles with frontal appendages usually
occurring in both sexes, always, when present, in the form of
permanent horns supported by bon}^ cores arising from the frontal
bones ; molars usually (always in European members of the
family) hypsodont ; lateral digits rudimentary or absent, usually
represented by the hoofs alone.

BemarJcs. — The family Bovidse is richer in both genera and
species than any other group of existing ungulates. About fifty
genera are usually recognized, three of which, Ovls, Capra and
Bupicapra, now occur naturally in western Europe.*

Genus OVIS Linnajus.

1758. Ovis Linna3us, Syst. Nat., i, 10th ed., p. 70.

1762. Aries Brisson, Regn. Anim. in CI. ix distrib., 2nd ed., p. 12.

1776. Musimon Pallas, Spicil. Zoologica, ii, fasc. 11, p. 8 {M. asiaticus).

1798. Mtisvion Scbrank, Fauna Boica, i, p. 81 (Substitute for Ovis).

1795. Aries Link, Zool. Beytrage, i, pt. ii, p. 96 (part).

1816. Aminon Blainville, Bull. Soc. Philomathique, Paris, p. 76, ]\Iay, 1816

(Alternative name for Ovis).
1857. Ovis Blasius, Saugetbiere Deutschlands, p. 466.

Type species. — Ovis aries Linnfeus (by tautonymy).

Geo(irap)hical distribution. — Holarctic region from Cyprus and
Asia Minor eastward across central and northern Asia, and in
western North America from Alaska to northern Mexico ; one
species isolated in the Mediterranean portion of Europe.

Characters. — Bovidde of small size and rather heavy form, the
tail normally short, the pelage dense, not tending to special
elongation on chin, throat and neck ; feet with glands between
the hoofs ; skull conspicuously wedge-shaped in general outline
from wide orbital region to narrow rostrum ; occipital region
abruptly bent downward, the upper surface concave or nearly
flat, the portion behind zygomata frequently almost tubular in
general form ; lachrymal pit present ; nasal branch of premaxillary
not wedged between nasal and maxillary ; horns always present
in males, usually absent or rudimentary in females, very robust

* Tbe genera Bos and Bison bave become extinct witbin historic times ;
a species of Buhalus, introduced in tbe seventeenth century, exists in a
wild or semi-feral state in parts of Italy.

ovis 987

at base, the under surface concave, the general form usually aii
outwardly directed spiral ; teeth strongly hypsodont, the upjser
molars without supplementary pillars in re-entrant angles, the
outer side of upper premolars with both terminal and median
ridges well developed ; lower incisors with crowns elevated, the
width of z'l when unworn not more than height.

Remarks. — The genus Ods contains about forty described
species, mostly Asiatic. Only one is definitely known to occur
wild in Europe.

OVIS MUSIMON Pallas.

1811. JEgoccros mtisimon Pallas, Zoogr. Rosso-Asiat., i, p. 230 (Sardinia).
1829. [Ovis musimon'] b var. occidentalis Brandt and Ratzeburg, Gctreue

Darstelluug und Beschreibung der Thiere, p. 55 (Corsica).
1841. Ovis niusmon Bonaparte, Iconogr. Faun. Ital., Indice distrib.
1857. Ovis musivion Blasius, Saugetbiere Deutscblands, p. 471.
1905. ? Ovis viatscJiiei Duerst, Martin Wilckens Grundziige der Naturgescb.

der Haustiere, 2nd ed., p. 180 (Corsica and Sardinia).
1910. Ovis musimon Trouessart, Faune Mamm. d'Europe, p. 242.

Type localitjj. — Sardinia.

Geograpliical distribution. — Corsica and Sardinia.

Diagnosis. — Size decidedly less than in ordinary domestic
sheep (head and body of rams about 1300; condylobasal length
of skull about 225 mm.) ; general colour of back and sides
reddish brown ; a blackish median stripe on neck and shoulders,
this spreading as a dark shade over sides behind shoulders ; a
conspicuous greyish white patch on posterior half of sides ; under-
parts of body and inner surface of legs dull whitish ; under
surface of neck with conspicuous black area, this continued
broadly down forearm and as a narrow stripe nearly to hoof ;
colour of side sepai^ated from that of belly by a black strij^e
beginning at axilla and extending down outer side of hind leg to
heel ; tail black above, whitish below ; skull small, with shallow
indistinct lachrymal pits ; horns occasionally present in female,*
those of male curved in a single plane or with tip bent either
outward or inward.

Measurements. — Adult males from Sardinia and Corsica
(skins) : head and body, 1270 and 1300 ; tail, 55 and 60 ; hind
foot, 220 and 240 ; ear, 70 and 70. For cranial measurements
see Table, p. 996.

Specimens examiwcf/. —Sixteen from Sardinia (B.M., U.S.N.M., and
Genoa) and two from Corsica (B.M. and U.S.N.M.).

Remarks. — Two species of Ovis are recognized by Duerst as
occurring together in both Corsica and Sardinia. Their characters
are said to be as follows : —

* See Lvdckker, Field, London, ex, pp. 147, 197, and Bernard, ibid.,
p. 147.

988 UNGULATA

Ocis masimon : foxy red with brown stripe on back, white
spots (Flecken) on head and flanks ; horns Hght coloured, curved
in a single plane.

Ovis matscJdei : more brownish grey in general colour, the
face ash-grey rarely marked with white ; horns dark brown,
curved in a spiral with tips directly strongly outward.

It seems not improbable that the Corsican form (occidentalis)
will prove to be distinct from true Ovis musimon. The single
male skin examined shows no appreciable peculiarities ; but the
females are said to be usually if not always horned, a condition
which rarely or never occurs in the Sardinian race.

C?St.

Gennargentu
Sardinia.

Mts.,

Ford G. Barclay
(c & P).

95. 4. 4. 1.

6, 9.

Gennargentu Mts.

(P. Bonomi.)

E. N. Buxton (p).

95. 4. 16. 6-7.

yg-

Sardinia.

A. B. B a y 1 e y -
Worthington (c & p).

5. 3. 6. 2.

S St.

Zoological Society.

53. 8. 29. 49.

2?.

Zoological Society.

61. 3. 24. 2.

62. 12. 22. 2.

6 skeleton.

Zoological Society.

60. 4. 23. 1.

6 skull.

(W. Ewer.)

Zoological Society.

55. 12. 26. 162,

6 horns.

Corsica.

Dr. C.I. Forsyth Major
(c).

9. 1. 11. 1.

Genus CAPRA Linnaeus.

1758. Gapra Linnaeus, Syst. Nat., i, 10th ed., p. 68 {hirctis, by tautonymy).

1762. Hircus Brisson, Regu. Anim. in Classis ix distrib., 2nd ed., p. 12

{Hircus Brisson = Capra hircus Linnaeus).

1766. Ibex Pallas, Spicil. Zoologica, ii, fasc. 11, p. 31 (sibiricus).

1795. Aries Link, Zool. Beytrage, i, pt. 2, p. 96 (Substitute for Capra).

1798. Tragus Schrank, Fauna Boica, i, p. 80 (Substitute for Capra).

1811. Mgoceros Pallas, Zoogr. Rosso-Asiat., i, p. 224.

1857. Capra Blasius, Saugethiere Deutschlands, p. 474.

Type species. — Capra hircus Linnaeus.

Geographical distribution . — Mediterranean region of Europe
(Portugal, Spain, Pyrenees, Alps and Grecian Archipelago) east-
ward through the Caucasus, Asia Minor and north-eastern
Africa to central Asia.

Characters. — Like Ovis externally but hind feet without
glands, males bearded, and horns never forming an outward or
inward spiral ; skull without lachrymal pit ; upper extremity of
premaxillary deeply wedged between nasal and maxillary ;
brain-case conspicuously convex above ; outer side of upper
premolars with terminal and median vertical ridges obsolete.

Remarks. — About thirty-five species are currently referred to
the genus Capra. Half a dozen of these occur in the Mediter-
ranean region of Europe.

989

CAPRA IBEX Linnseus.

1758. [Capra] ibex Linnseus, Syst. Nat., i, lOtli ed., p. 68.

1786, Capra alpina Girtanner, Observ. et Mem. sur la Phys., I'Hist. Nat.

et les Arts, xxviii, p. 224, March, 1786.
1857. Capra ibex Blasius, Saugethiere Deutschlands, p. 475.
1906. I[bex'] ibex Camerano, Mem. Accad. Reale Sci. Nat., Torino 1905-

1906, p. 284.
1910. Capra ibex Trouessart, Faune IMamm. d'Europe, p. 236.
1912. Capra ibex graicus Matschie, Deutsche Jagu. Zeitung, lix, no. 8,
April, 1912.

Tj/pe locality. — Swiss Alps.

GeograpMcal distribution. — Formerly throughout the Alps ;
now confined to a few localities in Piedmont, Italy.

Diagnosis. — Size larger than in ordinary domestic goats (head
and body of males about 1350 mm.) ; general colour a dusky grey,
darker on chin and upper part of throat and on belly ; legs
blackish below and along anterior surface above, this dark area
not noticeably contrasted or sharply defined ; tail like body at
base, blackish at tip ; horns very large, curved backward in a
single plane, conspicuously cross-ribbed on anterior surface, but
antero-internal border not elevated.

Measurements. — Adult male (mounted) : head and body, 1350 ;
tail, 155 ; hind foot, 310 ; ear, 85 ; height at shoulder, 760.
Adult female (skin) : head and body, 1000 ; tail, 30; hind foot,
290. For cranial measurements see Table, p. 996.*

Specimens examined. — Fifteen (B.M., U.S.N.M., and Turin).

S st. Italian side of Monte Purchased (Rowland 97. 12. 11. 1.

Rosa. Ward).

5 St. Alps. Purchased (Parreys). 41. 596.

6 St. Alps. Mrs. A. G. Campbell 650. 2.

6 skull. Alps. Messrs. Murray and 1835.

Brockedon (p).

6 horns. Chamonix. Purchased (Faery). 45. 12. 16. 1.

2 6 horns. 650. a. b.

? skull. Purchased (Warwick). 57. 3. 18. 2.

CAPRA PYRENAICA Schinz.
(Synonymy under subspecies.)

Geographical distribution. — Formerly throughout the moun-
tainous districts of the Iberian Peninsula and the Pyrenees.
Now extinct except in isolated colonies on the south side of the

* In the elaborate Tables of cranial measurements published by
Camerano, Ricerche intorno alio Stambecco delle Alpi (Mem. Accad. Reale
Sci. Nat., Torino, 2nd ser., lv, pp. 283-358, 1906; ibid., lvi, pp. 1-70,
1907), there are relatively few dimensions given which are comparable with
those here used. Twenty males, presumably adult, furnish the following
averages and extremes : basilar length, 252 • 1 (242-265) ; nasal, 92 • 1 (84-100) ;
maxillary tooth-row, 68*1 (62-76).

990 UNGULATA

Pyrenees, in the Serra do Gerez, Portugal, the Sierra de Gredos,
Sierra Morena, Sierra de Ronda, Sierra Nevada, Sierra Martes,
and Sierra de Card 6, Spain.*

Diagnosis. — Horns curved upward, outward and backward,
distinctly compressed laterally, the anterior sui'face without
evident transverse ridges, the antero-external border raised and
rib-like ; colour paler than in Capra ibex, and dark areas sharply
defined. f

HemarJis. — Four local races of Capra pjyrenaica are recognized
by Cabrera in his recent paper on " The Subspecies of the Spanish
Ibex." I have been unable to form any personal opinion as to
the validity of these forms. The following brief synopsis is based
entirely on the accounts published by Cabrera and Bocage.

Capra pyrexaica pyrenaica Schinz.

1838. Capra pyrenaica Schinz, Neue Denkschr. Allg. Schweiz. Gesellsch.

Naturwiss., Neuchatel, ii, p. 9.
1857. Capra pyrenaica Blasius, Saugethiere Deutschlands, p. 480.

1910. Capra pyrenaica Trouessart, Faune Mamm. d'Europe, p. 237.

1911. Cap^-a pyrenaica pyrenaica Cabrera, Proc. Zool. Soc, London, p. 966,

December, 1911.

Tijpje locality. — Spanish Pyrenees.

Geographical distribution. — Formerly the Pyrenees and eastern
part of Cantabrian chain. Now restricted to the region of
Mt. Perdido, Huesca, Spain.

Diagnosis. — Dark markings maximum for the species, the
lateral stripe broad, the black of withers extending downward
nearly or quite to that of upper part of leg ; horns widely
spreading, the rib on antero-internal border abruptly defined.

i St. &? skull. Pyrenees. Purchased (Boubee). 48. 2. 5. 4-5.

9 head st. Spain. Capt. J. Marriott (c&p). 0. 10. 28. 1.

6 St. (imm.). Sir R. Owen (p). 50. 9. 4. 1.

1 yg. Zoological Society. 68. 9. 12. 12.

* For map of present and past distribution, see Cabrera, Proc. Zool.
Soc, London, 1911, p. 965, December, 1911.

t " The s]pecies, as a whole, may be described as a pale brown animal
with the outer side of the limbs black, a black band on the lower part of
the flanks, and a short black mane, continued along the back by a narrow
stripe. The forehead and the beard are blackish or very dark brown, and
the belly and inner part of the limbs white. In winter jjelage there is a
whitish underfur, quite absent in summer, when the ground colour is
browner and the black areas become more abruptly definite. The females
lack at all seasons the mane and the black markings of the head and body,
presenting only a blackish tint on the anterior face of the limbs, and it is
the same with the young males, in which the black areas appear in the
second or third year, becoming larger and darker as the animal grows
older " (Cabrera, Proc. Zool. Soc, London, 1911, p. 967, December, 1911).

991

Capra pyrenaica lusitanica Fran(;a.

1909. Capra lusitanica Fran(;a, Bull. Soc. Portugaise Sci. Nat., ii, p. 144.
Based on the " Cabra-Montez da Serra do Gerez " of Bocage, Mem.
Acad. Real das Sciencias de Lisboa (Sci. Math. Phys. Nat.), N.S.,
II, pt. I, pp. 1-20, pis. 1-2. Articles separately paged. Bocage
wrongly cited as authority for the name lusitanica. Type in
Lisbon INIuseum.

1911. Capra pyrenaica, peculiar subspecies, Cabrera, Proc. Zool. Soc,
London, p. 966, December, 1911.

Type localiii/. — Serra do Gerez, Minho, Portugal.

Geographical distribution. — Formerly the mountains of Galicia
and northern Portugal. Now confined to the Serra do Gerez or
extinct.

Diagnosis. — ^Horns apparently less spreading than in the other
races.

Capra pyrenaica viCTORiiE Cabrera.

1911. Capra pyrenaica victorix Cabrera, Proc. Zool. Soc, London, p. 975,
December, 1911. Type in Madrid Museum.

Type locality. — Madrigal de la Vera, southern slope of Sierra
de Gredos, Province of Caceres, Spain.

Diagnosis. — "An intermediate form, in size and in the extent
of the black markings, between C. p. pjyrenaica and C. p. liispanica,
rather browner than liispanica in summer coat, and with [widely
spreading] horns similar in size to that race, but comparatively
broader and flatter."

Measurements. — Type, adult male, mounted (from Cabrera) :
head and body, 1355; tail, 130; hind foot with hoofs, 385; ear,
120; height at shoulder, 700. For cranial measurements see
Table, p. 997.

Capra pyrenaica hispanica Schimper.

1848. Cap-a hispanica Schimper, Comptes-Eend. Acad. Sci., Paris, xxvi,
p. 318, March, 1848.

1910. Capra pyrenaica hispanica Trouessart, Faune Mamm. d'Europe,

p. 237.

1911. Capra pyrenaica hispanica Cabrera, Proc. Zool. Soc, London, p. 966,

December, 1911.

Type locality. — Sierra Nevada (" Picacho de Veleta et Mula-
hacen "), Spain.

Geographical distribution. — Mountains of southern and eastern
Spain (Sierra de Honda, Sierra Nevada, Sierra Morena, Sieri'a
Martes and Sierra de Cardo).

Diagnosis. — Dark markings minimum for the species, the
lateral stripe very narrow, the black of withers strictly confined
to median line ; nasal bones more abruptly narrowed than in true

992 UNGULATA

pyrenaica ; horns widely spreading, less compressed than in
victorise, the antero-internal rib less abruptly defined than in
pyrenaica.

Measurements. — -Adult male (from Cabrera) : head and body,
1190 ; hind foot with hoofs, 305.

6 st. S. Spain. Purchased (Parzudaki). 55. 11. 2G. 14.

CAPRA .SGAGRUS Erxleben.

1777. [Capra] xgagrus Erxleben, Syst. Regni Anim., i, p. 260 (Caucasus).

1857. Capra xgagrus Blasius, Saugethiere Deutschlands, p. 485.

1858. ? JEgocerus ]pictus Erhard, Fauna der Cycladen, Wirbelthiere, p. 32

(Syra, Cyclades, Greece).
1888. ? Capra dorcas Reichenow, Zool. Jahrb., Syst., iii, p. 594, May 31,

1888. Island of Joura (Gyaros), Greece.
1899. ? C[apra] xgagrus var. jourensis Ivrea, Proc. Zool. Soc, London,

p. 599. Nomen nudum.
1S99. ? Capra cretensis Lorenz-Liburnau, Wissensch. Mitth. aus Bosnien

und der Hercegovina, vi, p. 865 (Crete).
1910. Capra legagrus picta, C. xgagrus dorcas and C. legagrus cretetisis

Trouessart, Farms Mamm. d'Europe, pp. 288-239.

Wild representatives of the domestic goat occur on several of
the islands of the Grecian Archipelago. Three local forms have
been described, the status of which is very imperfectly understood.
The Museum contains no material bearing on the question.

Genus RUPICAPRA Blainville.

1816. Rtipicap^-a Blainville, Bull. Soc. Philomathique, Paris, p. 75, IMay,
1816.

1840. Capella Keyserling and Blasius, Wirbelthiere Europas, p. iv.

1841. Cemas Gloger, Gemeinn. Hand- u. Hilfsbuch der Naturgesch., i, p. 153

(not of Oken, 1816).
1857. Capella Blasius, Saugethiere Deutschlands, p. 487.

Type species. — Capra rupicapra Linnaeus.

Geographical distribution. — Mountains of the Mediterranean
region from the Asturias and Pyrenees eastward to the Caucasus.

Characters. — General form less robust than in Ovis and
Capra ; skull with occipital region moderately bent downward,
not distinctly tubular posteriorly ; lachrymal pit absent ; nasal
branch of premaxillary not in contact with nasal ; horns present
and well developed in both sexes, rising almost perpendicularly,
their tips hooked abruptly backward, downward and slightly
outward ; teeth essentially as in Ovis, but crowns of incisors less
elongated, and outer side of upper premolars with terminal ridges
well developed, but median ridge obsolete.

Bemarks. — Four members of the genus Rupicapra are now
recognized, a number which will probably be increased when,
adequate material is brought together.

EUPICAPRA

993

RUPICAPRA RUPICAPRA LilUlRMlS.

1758. [Capra] 7-upicapra LiniiiEus, Syst. Nat., i, 10th ed., p. 68 (Switzerland).
1829. ? Rupicapra hamulicornis Burnett, Quart. Journ. Sci. Lit. Art.,

1829, pt. II, p. 353 (nomen nudum).
1843. Rupicapra tragus Gray, List Spec. IMamm. Brit. Mus., p. 167

(Kenaming of rupicapra).
1845. R[upicap7-a] capella Bonaparte, Atti della sesta Riunione degli

Scienziati Itaiiani, Milano, 1844, p. 337 (Renaming of rupicapra).
1847. [Capra rupicapra'] a sylvatica Sundevall, Kongl. Vetensk.-Akad.

Handl., Stockholm, 1845, p. 284 (Wooded portions of Alps).
1847. [Capra rupicapra'] /3 alpina Sundevall, Kongl. Vetensk.-Akad.

Handl., Stockholm, 1845, p. 284 (Higher portions of Alps).
1857. Capella rupicapra Blasius, Siiugethiere Deutschlands, p. 488.
1870. Rupicapra europca Cornalia, Fauna d'ltalia, pt. i, p. 53 (Substitute

for rupicapra ; " Guv." cited as authority).
1897. [Rupicapra] clorcas Schulze, Abhandl. u. Vortrage Gesammtb. Natur-

wissensch., iv, No. 10, p. 9 (Substitute iox rupicapra) .
1910. Rupicapra rupicapra Trouessart, Faune Mamm. d'Europo, p. 235.

Type locality. — Switzerland.

Geographical distribution. — Alps and Apennines, eastward
through Tirol, the Carpathians and northern portion of Balkan
Peninsula to the Caucasus.

Diagnosis. — General colour tawny brown in summer, blackish
brown in winter, the sides usually lighter than underparts and

Rupicapra rupicapra.
X i.

Fig. 213.

Rupicapra rupicapra.
Incisiform teeth (nat. size).

legs ; a dark dorsal line sometimes present ; throat with con-
spicuous buffy whitish area extending slightly behind interramial
region ; neck never conspicuously paler on ventral and dorsal
surfaces than on sides ; a dark line along median dorsal region ;

3 s

994

UNGULATA

horns essentially perpendicular, the bases of their cores wide
enough apart to allow for the presence of a noticeable flattened
or double concave area on surface of frontal between them.

Measurements. — Adult male (mounted) : head and bod}', 1100 ;
tail, 40 ; hind foot, 330 ; ear from crown, 110 ; height at shoulder,
690. For cranial measurements see Table, p. 997.

Specimens examined. — Twenty-three, from the following localities : —
Italy: Maratime Alps, 1 (Genoa) ; Val Formazza, Novara, 1 (Genoa).
Switzerland : No exact locality, 9 (B.M. and U.S.N.M.).
Austria-Hungary: Tirol, 5 (B.M. and U.S.N.M.) ; Transj'lvania ;
S. Carpathians, 2.
No History : Five.

6 St.

Tyrol.

H.R.H. Crown Prince 81. 9. 21. 1.
Rudolph of Austria

ad. and yg.

Tyrol.

Prince Rudolph of

78. 6. 20. 1-2.

sljeletons.

Austria (p).

2 6 skulls.

Alps.

Gen. Hardwicke (p).

631. a. b.

3 frontlets

Alps.

Dr. J. E. Gray (p).

46.10.13.31-33.

and horns.

6 skull.

Alps.

J. Gould (p).

46. 7. 7. 2.

6 skull.

Alps.

Dr. A. Giinther (c).

59. 9. 6. 102.

4 6 skulls

—

631. d. e. m. n.

and frontlets.

6 st. and

Hatszeg,

S. Carpa-

C. G. Danford (c).

86. 12. 27. 1.

skeleton.

thians,
vania.

, Transyl-

Foetus in al.

Hatszeg.

C. G. Danford (p).

94. 6. 26. 2.

Skeleton.

—

Purchased (Brandt).

631. g.

RUPICAPRA ORNATA Neumann.

1899. Bupicapra ornata Neumann, Ann. Mus. Civ. Stor. Nat. Genova,

XX, p. 347, December 20, 1899. Type in Genoa Museum.
1910. Bupicapi-a ornata Trouessart, Faune Mamm. d'Europe, p. 235.

Type locality. — Barrea, near Alfedena, Province of Aquila,
Italy.

Geographical distribution. — Abruzzi Mountains, Italy.

Diagnosis. — General colour as in bupicapra rupncapra, but
light throat-patch extending downward nearly to brisket, and
dorsal surface of neck almost or quite as pale, the two pale areas
separated along side of neck by a dark line extending from base
of ear downward and forward to join its fellow of opposite side in
region just above brisket ; horns slanting much more evidently
backward than in B. rupicapra, and area of upper surface of
brain-case somewhat reduced.

Measurements. — Adult male from the type locality (skin) :
head and body, 1200; tail, 30; pencil, 120; ear, 110±. For
cranial measurements see Table, p. 997.

Specimens examined. — Three, all from the tyx^e locality (B.M. and
Genoa).

6. Abruzzi Mts., Italy. Genoa Museum (e). 4. 2. 29, 1.

EUPICAPRA 995

RUPICAPRA PYRENAICA Bonaparte.

18i5. Rliqncapra] pyrenaica Bonaparte, Atti della sesta Uiuiiioiio dcgli

Scieuziati italiaui, Milano, 1844, p. 337.
1878. Antilope rupicapra var. Riitimeyer in Trutat, Bull. Soc. Hist. Nat.

Toulouse, xii, p. 117.
1910. Rupicapra rupicapra Trouessart, Faune Manim. d'Europe, p. 235

(Part ; in synonymy only).
1910. Rupicapra rupicapra pyrcnaica Cabrera, Proc. Zool. Soc. Loudon,

p. 998. December, 1910.

Type locality. — Pyrenees.

Geographical distribution. — Pyrenees.

Diagnosis. — Colour of winter pelage distinctly less dark than
in Rupicapra rupicapra ; markings on neck essentially as in
B. ornata ; horns about perpendicular, placed more closely
together than in the other species, the very nari-ow region of
frontal between bases of horn cores single concave ; teeth weak.

Measurements. — -Adult male from the Pyrenees (no exact
locality), from skin : head and body, 1140 ; tail, 40; hind foot,
344 ; ear, 95. For cranial measurements see Table, p. 997.

Specimens examined. — Three, all from the Pyrenees, no exact locality
(B.M. and U.S.N.M.).

Bemarks. — This veiy distinct species was accurately described,
though not recognized by name, by Riitimeyer and Trutat, in
1878.* Since then, until mentioned by Cabrera in 1910 (in
description of B. parva), it appears to have been completely
overlooked."!"

1 st. Pyrenees. Purchased (Boubee). 48. 2. 5. 7.

RUPICAPRA PARVA Cabrera.

1910. Rupicapra rupicapra, parva Cabrera,, Proc. Zool. Soc. Lordon, p. 999,
December, 1910. Type in Madrid Museum.

Type locality. — Picos de Europa, Santander, Spain.

Geograpliical distribution. — Cantabrian Mountains, Spain.

Diagnosis. — Described as smaller and darker than Bupicapra
pyrenaica, and with throat-patch very ill-defined, scarcely paler
than general colour of body.

Measurements. — Type (adult female) : height at shouldei-,
570 mm. ; length of horn along anterior curve, 146 (Cabrera).

6 St. Picos de Europa, Cantabrian J. W. R. Lee (c & p). 97. 1. 7. 3.
Alps.

* Bull. Soc. Hist. Nat. Toulouse, xii, pp. 115-117.

t See Lydekker, Great Game of P^urope, Asia and America, p. 185, 1901.
Trouessart places the name pyrenaica in the synonymy of rupicapra.

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APPENDIX.

Additional material shows the distinctness of the Sardinian
Grocidura mentioned on pp. 101 and 111-112. The animal has
been described as : —

CROCIDURA ICHNUSiE Festa.

1912. Crocidura ichmisx Festa, Boll. Mus. Zool. Anat. Comp., Torino,
XXVII, No. 648, p. 1, March 9, 1912. Type in Turin Museum.

Ty2)e locality. — Piscina, Lanusei, Sardinia.

Geographical distribution. — Sardinia.

Characters. — Like Grocidura sicula, but unicuspid teeth both
above and below fully as large as in G. russula, and third upper
unicuspid more conspicuously exceeding second than in either of
the related species. Brain-case apparently as deep in proportion
to its width as in G. russula.

Measurements. — ^Type (adult male) : head and body, 64 ; tail,
35; hind foot, 12. Average and extremes of five adults from
Ovile Seardu and Zinnigas : head and body, 65 '2 (60-69) ; tail,
36-8 (33-6-39); hind foot, 12 (11 •8-12-2). Immature female
from Assuni : head and body, 61 ; tail, 38; hind foot, 11-6.
Cranial measurements of type (teeth moderately worn), and of
immature female from Assuni (basal suture not closed, teeth not
worn): condylobasal length, 18 '8 and 18-6; zygomatic breadth,
6 - 2 and 6 - 0 ; lachrymal breadth, 4 • 0 and 4 - 2 ; breadth of
brain-case, 9 - 2 and 8 • 8 ; depth of brain-case (median), 4 • 8 and
4 - 6 ; mandible, 10*4 and 1 0 • 0 ; maxillary tooth-row, 8 • 8 and 9 • 0 ;
mandibular tooth-row, 8 • 0 and 8 • 2.

Specimens examined. — Seven, from the following localities in Sardinia:
Piscina, Lanusei, 1 (Turin; type); Ovile Seardu, 3 (Genoa); Zinnigas, 2
(Genoa) ; Assuni, 1 (U.S.N.M.).

The cranial measurements of Evotomys glareolus norvegicus
were accidentally omitte*! from Table, pp. 652-653, and are as
follows : —

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1000 APPENDIX

Genus Spalax (pp. 887-896). The living forms recognized
by Mehely (Species Generis Spalax. A Foldi Kutyak Fajai
Szarmazas- es Rendszertani Tekintetbeu, Budapest, 1909) as
occurring in Europe west of Russia, are as follows :* —

Sub-genus Mesospalax (p. 22). New sub-genus to contain the
species montlcola and hunyaricus. No type designated.

Spalax monticola inoniicola Nehring. Type locality : Kupres,
Bosnia. f (Type in Agricultural High School, Berlin.)

Spalax monticola turcicus Mehely (p. 25). Type locality :
vicinity of Makri-Koi, Constantinople, Turkey. (Type in
Bulgarian Royal Museum.)

Spalax monticola dolbrogese Miller.

Spalax monticola liellenicus Mehely (p. 29). Type locality :
Lamia, Thessaly, Greece. (Type in Hungarian National Museum.)

Spalax monticola liercegovinensis Mehely (p. 30). Type
locality : Ulog-Obruja, Herzegovina.f (Type in Agricultural
High School, Berlin.)

Spalax monticola syrmiensis Mehely (p. 30). Type locality :
0-Pazua, Szerem, Slavonia, Austria-Hungary. (Type in Agram
Museum.)

Spalax monticola serhicus Mehely (p. 31). Type locality :
Servia. (Type in Hungarian National Museum.)

Spalax fiungaricus hungaricus Nehring.

Spalax Jmngaricus transsiilvanicus Mehely (p. 31). Type
locality : Transylvania. (Type in Hungarian National Museum.)

Sub-genus Macrospalax (p. 23). New name for sub-genus
SjJalax.

Spalax grstcus grsecus Nehring. Type and paratype said to
have been taken in Greece ; two other specimens from the vicinity
of Czernowitz, Bukowina, Austria-Hungary.

Spalax istricus Mehely (p. 34). Type locality : Barza,
Roumania. (Type in Hungarian National Museum.)

Spalax polonicus Mehely (p. 35). Type locality : vicinity of
Lemberg, Galicia, Austria-Hungary. (Tj-pe in Lemberg Museum.)
As synonyms are cited : Podolian Marmot Pennant, Synopsis,
p. 271, 1771, and Glis zemni Erxleben, Syst. Regni Anim. i,
p. 370, 1777.

* Types of the new forms are apparently not designated ; but the lists
of material examined furnish indications of the specimens on which the
descriptions are mainly based.

t In the original description of monticola Nehring mentions two
specimens : (a) from Bosnia, and (6) from Herzegovina. Neither is desig-
nated as type. As a is young and b adult, and Nehring expressly notes
that the characters of the species are particularly well shown by b, I
assumed (p. 894) that 5 should be regarded as the t}^e. Mehely states
(p. 121) that a is the type of monticola. He bases his description of
liercegovinensis on Nehring's specimen b exclusively.

INDEX

[Priticijyal references are indicated by the use of heavy tyi)e.'\

abramus, Pipistrellus, 213
abramus, Vesperugo, 213
Acauthomys, 883
acervator, Mus, 878
acervifex, Mus, 878
Achlis, 979
Acomys, 888

,, miuous, 883
Acosminthus, 883
Acromys, 863
Adelonycteris, 225
sedilis, Vespertilio, 184
ffigagrus, Capra, 992
yEgoceros, 988
Aeorestes, 167

sestiva, Mustela erminea, 389
afra, Genetta, 451
africana, Mustela, 412
africanus, Putorius, 412
africanus, Putorius nivalis, 412
agilis, Micromys, 844
agilis, Mus (Apodemus) minutus, 844
agilis, Mus miuutus, 844
agilis, Vespertilio, 220
agrarius, Apodemus, 836
agrarius, Mus, 836
agrarius, Mus (Apodemus), 836
agrestis, Arvicola, 668, 671, 672, 673,

675
agrestis, Microtus, 662, 668, 671, 683
agrestis, Microtus agrestis, 668
agrestis, Mus, 668
Agricola, 659
agrius, Felis, 470
agrius, Felis ocreata, 470
alba, Arctomys marmota, 932
alba, Canis vulpes, 381
alba, Hystrix cristata, 543
alba, Lepus timidus, 526
alba, Lynx vulgaris, 471
alba, Mustela foina, 375
alba, jMustela lutreola, 415
alba, Talpa europsea, 4
albicans, Mus musculus, 869
albicans, Vesperugo kuhlii, 216
albicus, Castor, 948

albicus, Cervus, 966

albida, Talpa europeea, 4

albifrons, Cervus elaphus, 965

albigularis, Vesperus (Marsipolajmus),
238

albipes, Sorex, 88

albirostris, Mus agrarius, 836

albiventris, Crocidura aranea, 101

albiventris, Sorex fodiens, 70

albolimbatus, Vespertilio, 215

albo-maculata, Talpa ouropaea, 4

albo-uotatus, Sciurus vulgaris, 905

albus, Castor fiber, 947

albus, Cervns capreolus, 974

albus, Cervus elaphus, 965

albus, Cricetus vulgaris, 602

albus, Dama platyceros, 971

albus, Lepus, 529

albus, Lepus timidus, 502

albus, Mus amphibius, 745

albus, Mus arvalis, 683

albus, Mus musculus, 869

albus, Mus sylvaticus, 803

albus, Mustela putorius, 423

albus, Rattus domesticus, 853

albus, Sciurus vulgaris, 905

albus, Sorex fodiens, 69

albus, Ursus arctos, 286

Alcelaphus, 976

Aloes, 976

„ aloes, 978

aloes, Alces, 978

aloes, Cervus, 978

alcythoe, Vespertilio, 215

alexaudrino rattus, ]\Ius, 853

alexandrinus, Epimys rattus, '854

alexandrinus, ]\Ius, 854

alexandrinus, Mus (Epimys) rattus,
854

algidus, Lepus, 526

algirus, Erinaceus, 130

algirus, Erinaceus algirus, 131

Alopex, 818

,, lagopus, 319

,, spitzbergenensis, 324

alopex, Canis, 330

1002

alpina, Capra, 989

alpina, Capra rupicapra, 993

alpina, Marmota, 932

alpina, Sciurus vulgaris, 910

alpinus, Arvicola, 716

alpinus, Lepus timidus, 526

alpinus, Myotis muriuus, 193

alpinus, Putouius, 415

alpinus, Rhinolophus hipposideros,

150
alpinus, Sciurus, 914
alpinus, Sciurus vulgaris, 914
alpinus, Sorex, 60, 62
alpinus, Sorex alpinus, 62
alpinus, Ursus, 286
alpinus, Ursus arctos, 287
alticola, Sorex araneus, 43
altivolans, Vespertilio, 245
Amblyotus, 225
americana, Arvicola, 730
Ammomys, 752
Amnion, 986
amori, Eliomys, 560
amori, Mioxus nitela, 560
amphibius, Arvicola, 725, 730, 738,

744-746, 749
amphibius, Arvicola amphibius, 730
amphibius, Mus, 730
amphibius, Sorex, 70
Amphisorex, 29, 65
anglicus, Muscardinus avellanarius,

584
angularis, Microtus, 706
angustifrons, Arvicola agrestis, 671
angustifrons, Lutra, 356
Animalivora, 134
annulatus, Ursus arctos, 286
anomalus, Neomys, 81
anomalus, Neomys fodiens, 81
An Otis, 887
antinorii, Sorex, 62
antipse, Crocidura, 95
antipai, Crocidura, 95
antiquorum, Alces, 978
Aper, 956

aper, Sus setosus, 957
Aphrontis, 898
Apodemus, 791

,, agrarius, 886

,, epimelas, 794
flavicoUis, 828

flavicoUis, 829

,, ,, wintoni, 831

,, fridariensis, 825

„ hebridensis, 824

„ hirtensis, 825

,, svlvaticus, 797

" „ callipides, 809

,, ,, creticus, 818

„ „ dichrurus, 810

,, ,. sylvaticus, 803

aquatica, Arvicola, 730

aquaticus, Hypudseus paludosus, 738

aquaticus, Lemmus, 730

aquaticus, Sorex, 69

Aquias, 137

aquilonius, Lepus, 508

aquilonius, Lepus medius, 508

aquitanius, Microtus (Chionomys) ni-
valis, 717

aquitanius, Microtus nivalis, 717

araneus, Crocidura, 101

araneus, Sorex, 31, 101

araneus, Sorex araneus, 35

arcalus, Meles, 352

arctica, Vulpes, 819

arcticus, Gulo, 434

Arctogale, 384

Arctomys, 931

arctomys, Mus, 937

arctos, Gulo, 434

arctos, Ursus, 285

arenarius, Cricetus, 593

arenicola, Arvicola, 708

arenicola, Microtus, 708

argenteogrisea, Lepus timidus, 502

argenteus, Canis lagopus, 319

argenteus, Ursus arctos, 286

argentoratensis, Arvicola, 745

Aries, 986, 988

Aristippe, 225, 238

aristippe, Vespertilio, 219

arundinaceus, Mus, 846

arvalis, Arvicola, 683

arvalis, Lemmus, 632, 668

arvalis, Microtus, 681, 683

arvalis, Microtus arvalis, 683

arvalis, Mus, 683

arvensis, Arvicola, 683

arvensis, Mus, 844

Arvicola, 723

„ amphibius, 725

,, ,, amphibius, 730

,, illyricus, 741

,, italicus, 740

„ musignani, 744

,, sapidus, 732

„ ,, sapidus, 738

„ „ tenebricus, 735

,, scherman, 744

,, ,, exitus, 746

,, ,, monticola, 749

,, ,, scherman, 745

,, terrestris, 738

Aspalax, 887

assimilis, Arvicola arvalis, 683

asturianus, Microtus, 693

ater, Arvicola arvalis, 683

ater, Hypudseus terrestris, 738

ater, Mus (Epimys) rattus, 853

ater, Mus rattus, 853

ater, Rattus domesticus, 853

atlanticus, Cervus elaphus, 967

atlanticus, Euryalus, 155

1003

atlanticus, Rhinolophus curyalc, 155
atratus, Amblyotus, 234
atticus, Cricetulus, 593
attila, Sus, 960

aureolus, Llustela putorius, 4?5
aiireolus, Putorius putorius, 425
aureus, Brachyotus mystacinus, 169
aureus, Canis, 315
aureus, Ursus arctos, 286
auritus, Plecotus, 256
auritus, Vespertilio, 256
austriacus, Vespertilio auritus, 257
avellanarius, IMus, 683
avellanarius, Muscardinus, 588
avellanarius, Myoxus, 583
avellanus, Myoxus, 577
avenarius, Mus, 844
azoreum, Nyctalus, 254
azoreum, Pterygistes, 254
azoricus, Mus, 871
azoricus, Mus musculus, 871

badius, Ursus, 286

baiticus, Sciurus, 923

b??ticus, Sciurus vulgaris, 923

bisticus, Sus scrofa, 960

baibac, Arctomys, 937

bailloni, Arvicola, 672

bailloni, IMicrotus agrestis, 672

bajovaricus, Cervus, 966

balcanicus, Canis aureus, 315

balearica, Crocidura, 112

balearica, Genetta genetta, 452

balticus, Cervus, 966

Barangia, 354

Barbastella, 263

„ barbastellus, 263

barbastella, Vespertilio, 263

Barbastellus, 263

barbastellus, Barbastella, 263

barbastellus, Synotus, 263

barbastellus, Vespertilio, 263
bechsteiuii, Myotis, 179
becbsteinii, Vespertilio, 179
bergensis, Sorex araneus, 41
bicolor, jMyodes, 641
bicolor, Neomys fodiens, 73
bicolor, Sorex, 73
bifer, Rhinolophus, 150
bihastatus, Rhinolophus, 149
blasii, Rhinolophus, 162
blasii, Vespertilio, 187
blasiusi, Rhinolophus, 162
bobac, Arctomys, 937
bobac, Marmota, 937
bobak, Marmota, 937
bobak, ]\Ius, 937
boccamela, Mustela, 405
boccamela, IMustela nivalis, 405
boccamela, Putorius (Ictis), 405
boccamela, Putorius nivalis, 405
bolivari, Cervus elaphus, 969

bonapartii, Vespertilio, 219

borealis, Felis, 471

borealis, Gulo, 434

borealis, Lemmus, 615

borealis, Lepus, 526

borealis, Tarandus, 980

borealis, Vespertilio, 234

borealis, Vesperugo, 234

boscai, Eptesicus, 226

boscai, Vespertilio, 226

Bovidffi, 986

brachyotos, Vespertilio, 204

Brachyotus, 167

breviauritus, Lepus medius, 528

brevimanus, Plecotus, 256

brevipes, Plecotus auritus, 257

brevirostris, Mus, 869

britannicus, Arvicola, 673

britannicus, Evotomys glareolus, 634

britannicus, Evotomys hercynicus,

634
britannicus, Meles meles, 349
brunnea, Sciurus vulgaris, 910
brunneus, Pitymys pyrenaicus, 772
brunneus, Ursus arctos, 286
budapestiensis, Myotis ciliata, 177
buffonii, Lemmus arvalis, 745
bunites, Martes, 380
bunites, Mustela foina, 380

cabrerte, Euryalus, 155

cabrerse, Microtus, 701, 703

cabrcrai, Microtus, 701

cabrerai, Rhinolophus euryale, 155

caeca, Talpa, 15

cserulea, Canis vulpes, 319

cresarius, Evotomys, 645

calcarata, Romicia, 215

callipides, Apodemus sylvaticus, 809

callipides, Micromys sylvaticus, 809

callipides, Mus sylvaticus, 809

campestris, Arvicola, 683

campestris, Cervus vulgaris, 966

campestris, Mus, 844

campestris, Mus (Apodemus) minu-

tus, 844
campestris, Mus minutus, 844
Campicola, 659
campicola, Lepus, 502
canaliculatus, 55
candidus, INIus sylvaticus, 803
candidus, Sorex araneus, 101
canese, Crocidura, 109
canescens, Cricetus cricetus, 603
canescens, Cricetus vulgaris, 603
canescens, Lepus, 526
caniceps, Eriuaceus, 120
canicularius, IMus, 878
caniculator, IMus, 878
CanidtB, 303

caninus, Erinaceus, 120
cauinus, Meles communis. 348

1004

Caiiis, 304

,, aureus, 315

,, lupus, 305

,, „ dietanus, 315

„ lupus, 813

„ ,, signatus, 814

cantabra, Crocidura, 99
cantabra, Crocidura inimula, 99
canus, Canis lupus, 313
canus, Capreolus capreolus, 975
canus, Mus ampbibius, 745
capaccinii, IMyotis, 187
capaccinii, Myotis (Leuconoe), 187
capaccinii, Vespertilio, 187
Capaccinius, 166
Capella, 992
capella, Rupicapra, 998
Capra, 988

„ aegagrus, 992

„ ibex, 989

„ pyrenaica, 989

„ „ hispanica, 991

,, ,, lusitanica, 991

„ ,, pyrenaica, 990

,, ,, Victoria, 991

caproea, Capreolus, 974
Caprea, 972
Capreolus, 972

,, capreolus, 973

„ ,, canus, 975

„ ,, capreolus, 974

tbotti, 975
„ ,, transsylvanicus,

975
capreolus, Capreolus, 973, 974
capreolus, Capreolus capreolus, 974
capreolus, Cervus, 974
Caprios, 21

capucinellus, Vespertilio, 184
capuciuus, Pitymys subterraneus, 760
carinatus, Sorex, 69
Carnivora, 283

carpatbicus, Sorex araneus, 43
carpetanus, Rbinolopbus, 159
caspius, Mus, 858
castaneus, Arvicola terrestris, 746
castaneus, Sorex araneus, 37
castaneus, Sorex tetragonurus, 37
castilianus, Sus scrofa, 960
Castor, 947

„ fiber, 947
Castoridse, 947
Cateorus, 225
Catolynx, 456
Catus, 456
catus, Felis, 462
caudata, Crocidura, 110
caudata, Talpa, 3
cellarius, Mus, 829
celtica, Sus scrofa, 957
celticus, IMus sylvaticus, 803
Cemas, 992

centralis, Pitymys ibericus, 782

Cervaria, 471

cervaria, Felis, 472

Cervidse, 962

Cervus, 963, 970
„ elaphus, 964
„ ,, atlanticus, 967

,, ,, corsicanus, 969

,, ,, elaphus, 967

„ ,, germanicus, 965

„ ,, hispanicus, 969

„ „ scoticus, 968

cestoni, Dinops, 277

cestonii, Dysopes, 277

cestonii, Nyctinomus, 277

Cbionobates, 495

Cbionomys, 712

Cbiroptera, 134

chrysothorax, Sorex, 101

ciliatus, Neomys fodiens, 73

ciliatus, Sorex, J3

ciliatus, Vespertilio, 177

cincticauda, Eliomys, 559

cinerea, Canis vulpes, 331

ciuerea, Sciurus vulgaris, 910

cinerea, Talpa europsa, 4

cinereo-maculatus, Mus musculus, 869

cinereus, Lepus timidus, 502

cinereus, Sciurus vulgaris, 906

cinereus, Sorex araneus, 101

cintrae, Crocidura russula, 108

Citellus, 924

citellus, 924
,, suslica, 929

citellus, Citellus, 924

citellus, Citellus (Citellus), 924

citellus, Mus, 924

citellus, Spermopbilus, 924

Citillus, 924

citillus, Mus, 924

clivosus, Rbinolopbus, 162

Cnephseus, 224

cnossius, Oryctolagus cuniculus, 491

Coelopbyllus, 137

coUaris, Sorex, 69

coUaris, Vespertilio, 169

coUinus, Lepus borealis, 526

coUinus, Lepus timidus, 527

Comastes, 167

communis, Barbastellus, 263

communis, Genetta, 451

communis, Meles, 848

communis, Plecotus, 256

communis, Vulpes, 830

concinnus, Sorex, 35

console!, Erinaceus europseus, 126

constrictus, Sorex, 101

contigua, Arvicola arvalis, 683

corilinum, Mus, 584

cornutus, Vespertilio, 256

coronatus, Lepus timidus, 502

coronatus, Sorex, 35

1005

corsicanus, Cervus, 'J69
corsicanus, Ccrvus elaphus, DG'J
corsicanus, Lepus, 507
corsicauus, Lepns europtcus, 507
corsicanus, Lcpus meditorraneus, 507
corsicanus, Putorius nivalis, 412
Corsira, 2'J
Craseomys, 624

crassicaudatus, Sorex vulgaris, 43
crassidens, Lemmus lemmus, 621
cretensis, Capra, y92
cretensis, Capra ffigagrus, 992
creticus, Apodemus sylvaticus, 813
creticus, Lepus, 512
creticus, Lepus europseus, 512
Cricetinae, 592
Cricetulus, 593

,, atticus, 598

Cricetus, 596

,, cricetus, 597

,, ,, canescens, 603

,, ,, cricetus, 602

,, ,, nehringi, 605

cricetus, Cricetus, 597, 602
cricetus, Cricetus cricetus, 602
cricetus, Mus, 602
cristata, Hystrix, 543
Crocidura, 86

,, balearica, 112

„ caneae, 109

„ caudata, 110

„ cyrnensis, 311

,, ichnusse, 998

,, leucodon, 88

,, mimula, 94

,, ,, cantabra, 99

„ „ iculisrna, 98

„ „ mimula, 95

,, russula, 99
,, ,, cintrse, 108

„ „ pulchra, 103

„ ,, russula, 101

,, sicula, 108

Crossopus, 65
crucigera, Canis, 331
crucigera, Vulpes vulpes, 331
cunicularius, Arvicola, 683
Cuniculus, 484
cuuiculus, Lepus, 490
cuniculus, Oryctolagus, 485, 490
cuniculus, Oryctolagus cuniculus, 490
cylindricornis, Raugifer tarandus, 981
Cynomionax, 418
cyrnensis, Crocidura, 111

dacius, Pitymys, 760
Dactyloceros, 970
dalmatinus, Canis, 315
Dama, 970

„ dama, 970
dama, Cervus, 970
dama, Dama, 970

dauubicus, Brinaceus, 127

danubicus, Erinaceus curopanis, 127

dasycarpos, Vcspertilio, 252

dasycneme, IMyotis, 189

dasycneme, Vespertilio, 189

dasj-pus, Vespertilio, 187

daubentonii, Barbastellus, 263

daubentouii, Myotis, 184

daubentonii, Sorex, 69

daubentonii, Vespertilio, 184

daubentoni, Myotis, 184

decumanus, Mus, 858

dentatus, Microtus, 703

depressa, Arvicola arvalis, 683

depressus, Pitymys, 779

Desman, 21

Desmana, 21

Desmaninee, 20

Desmanus, 21

destructor, Arvicola, 744

dicbrurus, jMuscuIus, 810

dicbrurus, Apodemus sylvaticus, 810

dichrurus, Mus sylvaticus, 810

Dieba, 304

dietanus, Canis lupus, 315

discolor, Vespertilio, 238

discolor, Vesperugo, 238

dolbrogese, Spalax, 889

dolbrogeae, Spalax mouticola, 1000

dombrowskii, Mustela (Ictis), 405

dombrowskii, Putorius (Ictis) nivalis,
405

domestica, Martes, 375

domesticus, Rattus, 853

dorcas, Capra, 992

dorcas, Capra segagrus, 992

dorcas, Capreolus, 974

dorcas, Rupicapra, 993

dorsalis, Herpestes ichneumon, 441

drueutius, Pitymys, 762

dry as, Eliomys, 570

dryas, Myoxus, 568

Drymomys, 863

Dryomys, 566, 863

duodecimcostatus, Arvicola, 784

duodecimcostatus, IMicrotus, 784

duodecimcostatus, Pitymys, 784

duplicata, Arvicola arvalis, 686

duplicatus, Microtus arvalis, 686

Duplicidentata, 483

Dyromys, 566

„ nitedula, 567
„ ,, intermedius, 569

„ ,, nitedula, 568

„ „ wingei, 570

„ robustus, 572

echinus, Erinaceus, 120
eggenhoffner, Rhinolophus, 150
Elaphus, 963

elaphus, Cervus, 964, 965, 967
elaphus, Cervus elaphus, 967

1006

Eliomys, 550, 566

„ gymnesicus, 558
,, lusitanicus, 560
,, pallidus, 559
,, quercinus, 551
„ sardus, 560

Elius, 572

emarginatus, Myotis, 177

emarginatus, Vespertilio, 177

Emballonuridse, 276

epimelas, Apodemus, 794

epimelas, Mus, 794

epimelas, Mus (Epimys), 794

Epimys, 848

„ norvegicus, 858

„ rattus, 849

,, ,, alexandrinus, 854

,, ,, rattus, 853

Eptesicus, 224

„ nilssonii, 234

,, serotinus, 226

,, sodalis, 231

equinus, Vespertilio, 142

eremita, Sorex, 69

Erinaceidse, 114

Erinaceus, 114

algirus, 130, 131
,, ,, algirus, 131

,, ,, vagans, 133

,, europseus, 115

„ ,, consolei, 126

„ ,, europseus, 120

„ „ hispanicus, 122

,, ,, italicus, 123

„ nesiotes, 129

,, roumanicus, 127

erinaceus, Hystrix, 120

erminea, Poetorius, 387

erminea, Mustela, 385, 387

erminea, Mustela erminea, 387

erminea, Putorius, 388

ermineus, Putorius (Ictis), 388

escalerai, Myotis, 174

esculentus, Glis, 577

etrusca, Pachyura, 83

etruscus, Sorex, 83

Euarctos, 285

Euarvicola, 659

Eucervaria, 471

Eucervus, 963

Eulagos, 495

Eulepus, 495

Eumustela, 384

eurouotus, Sorex araneus, 41

europsea, Hystrix cristata, 543

europsea, Lutreola, 415

europsea, Talpa, 3

europseus, Alces, 978

europseus, Canis Vulpes alopex, 331

europseus, Capreolus, 974

europseus, Erinaceus, 115, 120, 123

europseus, Erinaceus europseus, 120

europseus, Lepus, 498, 502, 508

europseus, Lepus europseus, 502

europseus, Macrotus, 250

europseus, Mclcs, 348

europaius, Sus, 957

europea, Rupicapra, 993

euryale, Rhinolophus, 155

Euryalus, 137

euryrhinus, Ursus, 286

Euvespertilio, 167

Euvesperugo, 203, 243

eversmanni, Myrmarctos, 287

Evotomys, 623

„ csesarius, 645

glareolus, 626
„ „ britannicus, 634

„ ,. glareolus, 632

„ ,, hallucalis, 643

„ ,, helveticus, 640

„ ,, istericus, 637

„ ,, nageri, 641

„ ,, norvegicus, 638

,, ,, suecicus, 636

„ ,, vasconise, 639

,, rufocanus, 648

rutilus, 646
,, skomerensis, 644

exilis, Arvicola scherman, 747

exilis, Sorex, 55

exitus, Arvicola scherman, 746

Exochurus, 167

exsul, Microtus agrestis, 669

fseroensis, Mus, 875

fseroensis, Mus musculus, 875

fagorum, Mustela martes, 375

falciger, Ursus, 286

fatioi, Pitymys, 763

fatioi, Pitymys multiplex, 763

favonicus, Myotis bechsteinii, 179

Pelidse, 455

Felis, 456

,, agrius, 470

,, sarda, 468

„ silvestris, 457

,, ,, grampia, 464

,, ,, silvestris, 462

,, ,, tartessia, 465

fennicus, Rangifer, 981

fennicus, Rangifer tarandus, 981

ferox, Catus, 462

ferruginea, Herpestes ichneumon, 441

ferrugineus, Vespertilio, 244

ferrum-equinum, Rhinolophus, 139,
142

ferrum-equinum, Rhinolophus fer-
rum-equinum, 142

ferrum-equinum, Vespertilio, 142, 149

ferus, Felis catus, 462

Fiber, 947

fiber, Castor, 947

fimbriatus, Crossopus, 70

1007

fimbriatus, Sorex, 70, 101
flava, Arvicola arvalis, 690
llavescens, ilus musculus, 870, 878
llavoscens, Nannugo pipistrellus, 205
flavescens, Talpa europrea, 4
tiavicans, Mustela putorius, 423
flavicollis, Apodemus, 828
flavicoUis, Apodemus flavicoUis, 829
flavicollis, Mus, 829, 831
flavus, Canis lupus, 313
flavus. Castor, 947
flavus, Lepus timidus, 502
flavus, ]\Ius musculus, 869
fluviatilis, Lutra, 356
fluviatilis, Sorex, 69
fodiens, Crossopus, 70, 73
fodieus, Cuuiculus, 490
fodiens, Neomys, 66
fodiens, Neomys fodiens, 69
fodieus, Sorex, 42, 69, 70
foetens, Yiverra, 423
foetidus, Putorius, 423
FoBtorius, 384, 415, 418, 428
foiua, Martes, 374
foina, Martes foiua, 375
foina, Mustela, 375
formicarius, Ursus, 286, 297
formicarius, Ursus arctos, 287
fretalis, Sorex araneus, 45
fridariensis, Apodemus, 825
fridariensis, Mus sylvaticus, 825
frisius, Talpa, 3
frugivorus, Musculus, 854
frumentarius, Cricetus, 602
fuliginosus, Arvicola musignani, 744
fuliginosus, Canis, 324
fuliginosus, Mus tectorum, 853
fuliginosus, Vulpes lagopus, 324
fulva, Arvicola arvalis, 690
fulva, Mustela Lutra, 415
fulvaster, Rattus domesticus, 853
fulvus, Arvicola, 632
fulvus, Canis lupus, 313
fulvus. Castor fiber, 947
fulvus, Lemmus, 683
fulvus, Mus cricetus, 602
furcifer, Tarandus, 981
furo-putorius, Mustela, 423
fusca, Arvicola agrestis, 759
fuscoater, Sciurus vulgaris, 910
fuscoatra, Sciurus vulgaris, 910
fuscus, Pitymys ibericus, 783
fuscus, Rattus domesticus, 853
fuscus, Ursus, 286
fuscus, Ursus arctos, 286

Gale, 384

gale, Mustela, 402

Gale my s, 20, 65

,, pyrenaicus, 21

„ „ pyrenaicus, 26

„ „ rufulus, 26

galinthias, Mustela, 414

galinthias, Putorius (Ictis) nivalis, 414

galinthias, Putorius nivalis, 414

gallsecius, Lepus granatensis, 517

galliardi, Arvicola arvalis, 683

gallia3. Castor, 947

gallica, Viverra Genetta, 451

gallicus, Castor fiber, 947

Galomys, 20

genei, Pipistrellus, 204

Genetta, 446

„ genetta, 447
,, ,, balearica, 452

,, „ genetta, 451

,, ,, rhodanica, 452

genetta, Genetta, 447, 451
genetta, Genetta genetta, 451
genetta, Viverra, 451
gerbei, Arvicola (Microtus), 773
gerbei, Pitymys, 773
gerbii, Arvicola (Microtus), 773
gerbii, Pitymys, 773
germauicus, Cervus elaphus, 965
germanicus, Mus cricetus, 602
germanicus, Rhinolophus ferrum-

equinum, 142
ghidinii, Vespertilio, 179
ghidinii, Vespertilio bechsteinii, 179
glareolus, Arvicola, 632, 641, 643
glareolus, Evotomys, 626, 645
glareolus, Evotomys glareolus, 632
glareolus, Hypudseus, 632
glareolus, Mus, 632
Glires, 481
Gliridai, 549
Glis, 572, 931

„ glis, 573

„ „ glis, 577

,, ,, italicus, 578

,, ,, melonii, 579

,, ,, pyrenaicus, 582

„ zemni, 1000
glis, Glis, 573, 577
glis, Glis glis, 577
glis, Myoxus, 577
glis, Sciurus, 577
gotthardi, Sciurus vulgaris, 910
grseca, Sciurus vulgaris, 910
grsecus, Canis, 315
grfecus, Spalax, 895
grsecus, Spalax grtecus, 1000
graicus, Capra ibex, 989
grampia, Felis, 464
grampia, Felis silvestris, 464
granarius, Sorex araneus, 52
granatensis, Lepus, 515, 516
granatensis, Lepus europreus, 516
granatensis, Lepus granatensis, 516
grandis, Ursus arctos, 286
gregarius, Mus, 668
grisea, Herpestes ichneumon, 441
grisea, Talpa europtea, 4

1008

griscogularis, Crossopus ciliatus, 70
griseus, Ursus arctos, 28G
groenlandicus, Canis, 324
Gulo, 433

„ gulo, 434
gulo, Gulo, 434 -
gulo, Mustela, 434
Guloninffi, 432
guttatus, Citellus, 829
guttatus, Mus citellus, 929
guttulatus, Spermophilus, 929
gymnesicus, Eliomys, 558
gymnesicus, Eliomys querciuus, 558

hallucalis, Evotomys glareolus, 643

hallucalis, Evotomys iiageri, 643

hamiltonii, Eliomys, 551

Hamster, 596

hamulicoruis, Rupicapra, 993

hartingi, ]\Iicrotus, 704

hayi, Mus sylvaticus, 810

hebridensis, Apodemus, 824

hebridensis, Mus, 824

hebridensis, Mus sylvaticus, 824

Heliomys, 596

hellenicus, Spalax moiiticola, 1000

helvetica, Rhinolophus euryale, 150

helveticus, Evotomys glareolus, 640

helveticus, Evotomys hercynicus, 640

helvolus, Mus musculus, 869

Hemiotomys, 659, 723

hercegovinensis, Spalax monticola,
1000

hercynicus, Evotomys hercynicus,
632, 637

hercynicus, Hypudteus, 632

hercynicus. Sores alpinus, 63

Herinaceus, 114

hermanni, Sorex, 42, 70

herminea, Mustela, 387

Herpestes, 440

hibericus, Putorius, 407

hibernica, Mustela, 398

hibernicus, Lepus, 531

hibernicus, Lepus timidus, 531

hibernicus, Mus, 858

hibernicus Mus (Epimys) norvegicus,
858

hibernicus, Putorius, 398

hibernicus, Putorius (Ictis), 398

hibernicus. Sores rusticus, 55

hippocrepis, Vespertilio, 142

hipposideros, Rhinolophus, 147, 150

hipposideros, Rhinolophus hippo-
sideros, 149

hipposideros, Vespertilio, 149

hipposiderus, Rhinolophus, 150

Hircus, 988

hirta, Arvicola, 673

hirtensis, Apodemus, 825

hirtensis, Mus, 825

hirtensis, Mus sylvaticus, 825

hirtus, Microtus agrcstis, 673
hispanica, Capra, 991
hispauica, Capra x^yrcnaica, 991
hispanica, Viverra genetta, 451
hispanicus, Cervus elaphus, 969
hispanicus, Erinaceus europaeus, 122
hispanicus, Lepus, 516
hispanicus, Mus spicilegus, 879
Histrix, 542
Homalurus, 29
homorodalmasiensis, Rhinolophus

unihastatus, 142
homorodensis, Rhinolophus ferrum-

equinum, 142
hortualis, Eliomys, 551
humeralis, Vespertilio, 169
hungaricus, Mesospalax, 1000
hungaricus, Spalax, 889, 894
hungaricus, Spalax hungaricus, 1000
hungaricus, Spalax typhlus, 894
huxleyi, Lepus, 491
huxleyi, Oryctolagus cuuiculus, 491
hyberna, Mustela erminea, 387
hybridus, Lepus, 508
hybridus, Lepus europseus, 508
hybridus, Mus decumanus, 858
Hydrogale, 65
Hydromustela, 415
hydrophilus, Sorex, 69
Hydrosores, 65
hydruntiua, Crocidura, 101
hypomelas, Vulpes, 331
Hypsugo, 202
Hypudseus, 623
Hystricidse, 542
Hystridse, 542
Hystris, 542

,, cristata, 543

iberica, Mustela nivalis, 407

iberica, Putorius nivalis, 407

ibericus, Arvicola, 782

ibericus, Arvicola (Microtus), 782

ibericus, Microtus, 782

ibericus, Pitymys, 780, 782

ibericus, Pitymys ibericus, 782

ibericus, Putorius nivalis, 407

Ibex, 988

ibex, Capra, 989

ibex. Ibex, 989

ichnusse, Crocidura, 998

ichnusse, Vulpes, 336

ichnusffi, Vulpes vulpes, 336

Ictis, 384

iculisma, Crocidura mimula, 98

ignotus, Crossopus ou Sorex, 43, 70

illyricus, Arvicola, 741

illyricus, Microtus musignaui, 741

litis, Mustela, 423

incertus, Arvicola, 690

incertus, Microtus, 690

incertus, Microtus arvalis, 691

1009

incortus, Pitymys, 691

incisivus, Vespertilio, 226

infoctus, Putorius, 423

infubcatus, Sciiu-us, 916

iufuscatus, Sciurus vulgaris, 916

iiiodorus, Sorex, 101

lusectivora, 1, 131

insul, INIicrotus agrestis, 669

insulauus, Rhinolopbus fcrrum-equi-

num, 1-il
insularis, Eptesicus serotinus, 226
insularis, Glis, 578
insularis, Lemmus, 668
intermedia, Arvicola, 672
intermedius, Dyromys uitedula, 569
intermedius, Mus, 803
intermedius, ]Mus (Epimys) rattus, 854
intermedius, IMus sylvatious, 803
intermedius, Mus rattus, 851
intermedins, ]\Iyoxus, 569
intermedius, Sorex, 62, 70
intcrstriatus, Mus, 539
interzonus, ^lus, 539
isabelliuus, Vespertilio, 226
islaudicus, Mus, 803
Isotus, 167

istericus, Evotomys glareolus, 637
ititricus, Spalax, 1000
italica, Arvicola amphibius, 740
italicus, Arvicola, 740
italicus, Erinaceus europajus, 123
italicus, Glis, 578
italicus, Glis glis, 578
italicus, Putorius nivalis, 405
italicus, Rbiuolophus ferrum-equi-

num, 142
italicus, Sciurus, 912
italicus, Sciurus vulgaris, 812, 913
iturissius, Lepus grauatensis, 518

jenaensis, Tbalassarctos, 293
jeudii, Heliomys, 602
jourensis, Capra icgagrus, 992
jubata, Alces, 978

kattlo, Felis, 471
kirschbaumii, Plecotus, 257
kisnyircsiensis, Khiuolopbus biba-

status, 150
kublii,>ripistrellus, 215
kuhlii, Vespertilio, 215, 231
kulilii, Vcsperugo, 216
kubli, Pipistrcllus, 216

labiosus, Sorex, 35
LagidiE, 484
Lagomys, 931
lagopus, Canis, 319
lagopus, Vulpes, 319
Lagos, 495

lanatus, Vespertilio, 184
lapponum, Tarandus, 980
lardarius, Vespertilio, 245

lasioptorus, Vespertilio, 244

Latax, 354

Lataxina, 354

latifrons, Arvicola agrestis, 671

latinorum, Martes martes, 373

latinorum, IMustela martes, 373

latipinnis, Vespertilio, 193

lebruni, Microtus (Chiouomys), 719

Icbrunii, Arvicola, 719

lebrunii, Microtus, 718

lebruuii, Microtus lebrunii, 719

Leggada, 863

leisleri, Nyctalus, 252

leisleri, Vespertilio, 252

leisleri, Vesperugo, 252

Lemmus, 614

lemmus, 615

,, crassideus, 621

lemmus, Lemmus, 615

lemmus, IMus, 615

lemmus, Myodes, 615

Icponticus, Microtus, 764

Lcporidie, 484

Leporina, 484

Lepus, 484. 495
,, crcticus, 512

europteus, 498
,, ,, corsicanus. 507

,, ,, europaeus, 502

hybridus, 508
,, meridiei, 506
,, ,, occidentalis, 504

,, ,, pyreuaicus, 506

,, ,, transsjivaiiicus, 509

,, granatensis, 515
,, ., galleecius, 517

., granatensis, 516

,, iturissius, 518

,, hibernicus, 531
,, mediterraueus, 513
,, parnassius, 519
,, timidus, 522
,, ., scoticus, 529

,, -, timidus, 526

,, ., varronis, 528

•leuotetiuops, Gorvus dama, 971

leucippe, Vespertilio, 219

leucocepbalus, ]\Ius sylvaticus, S03

Leucocyon, 318

Leucodon, 86

loucodon, Crocidura, 88

leucodon, Crocidura russula, 88

leucodon, Sorex, 88

leucodus, Crocidura, 88

leucogastcr, IMus, 854

Ijeuconoc, 166

Icucopictus, Arctomys citellus, 929

Leucorhyncbus, 65

leucotis, Sorex fodiens, 70

leucourus, Sciurus vulgaris, 907

leucurus, Arvicola, 722, 719

leucurus, Microtus lebrunii, 722
3 T

1010

leuciu'us, i\Iicrotus (Chionomys) le-

bi'uiii, 122
Icucurus, Sciurus, 907
Icucurus, Sciurus vulgaris, 907
leucurus, Sorex, 101
levcrnedii, Arvicola, 671
levernedii, IMicrotus agrestis, G71
levis, IMicrotus, 687
levis, Microtus arvalis, 687
lilfeus, Sciurus vulgaris, 913
lilfordi, Lcpus, 516
limbatus, Nannugo pipistrellus, 205
limnophilus, Vcspertilio, 189
lineatus, Canis vulpus, 330
lineatus, Sorcx, 69
liuneaua, Amphisorex, 70
Lipura, 931
liricaudatus, Sorex, 69
littoralis, Hypudteus paludosus, 738
longobarda, Sorex alpiuus, 62
longobardus, Sorex alpinus, 70
Lontra, 351
loriger, Sicista, 537
loriger, Sminthus, 537
lucanius, Sorex minutus, 60
lugubris, Vespertilio, 169
lupulinus, Felis, 471
Lupus, 301

lupus, Canis, 305, 313
lupus, Canis lupus, 313
luscus, Gulo, 131
lusitanica, Capra, 991
lusitanica, Capra pyrenaica, 991
lusitanica, Eliomys nitela, 560.
lusitanicus, Arvicola (Microtus), 776
lusitanicus, Eliomys, 560
lusitanicus, Eliomys quercinus, 560
lusitanicus, Microtus (Pitymys), 770
lusitanicus, I\lus spicilegus, 882
lusitanicus, Pitymys, 776
lutea, Canis vulpes, 331
lutea, Talpa europiea, 4
lutcscens, Lepus timidus, 531
Lutra, 854

„ lutra, 355
lutra, Lutra, 355
lutra, Mustcla, 355
Lutreola, 415
lutreola, Foetorius, 415
lutreola, IMustela, 415
lutreola, Putorius (Lutreola), 415
Lutrinae, 354
Lutris, 354
Lutrix, 354
Lutrogale, 355
Lutronectcs, 355
lycaon, Canis lupus, 313
Lyciscus, 304
Lyuceus, 470
Lyuchus, 471
lyncula, Fclis, 171
Lynx, 170

Lynx lynx. 471

,, pardellus, 475
lynx, Fclis, 171
lynx, Lynx, 471

JIachlis, 970

machlis, Alces, 978

inacropterus, Vesperugo pipistrellus,

204
Macrospalax, 1000
macrotrichus, Sorex, 35
Macrotus, 256
macrourus, Sorex, 70
maculata, Mustela erminea, 387
maculata, Talpa curopa^a, 4
maculatus, Lepus timidus, 502
maculatus, Mus agrarius, 836
maculatus. j\Ius musculus, S69
magnus, Vespertilio, 245
major, Canis lupus. 313
major, Crocidura, 101
major. Foetorius pusillus. 405
major I\Ius sylvaticus, 829
major, Mustela erminea, 389
major, Rhinolophus, 142
major, Ursus arctos, 286
major, Vespertilio, 245
major, Vespertilio ferrum-equinum,

142
majori, Vespertilio, 187
I\Iangusta, 440

manium, Putorius putorius, 423
maral, Cervus elaphus. 965
marginatus, Pipistrellus, 215
inarginatus, Vespertilio, 215
mariic, Microtus (Pitymys), 777
marite, Pitymys, 777
marianensis, INIeles meles, 352
marianensis, jMeles taxus, 352
marinus, Lutra vulgaris, 356
marinus, Ursus, 298
maritimus, Thalarctos, 298
maritimus, Ursus, 298
maritimus, Ursus (Thalassarctos), 298
Marmota, 931

,, bobak, 937

,, . marmota, 932
marmota, Marmota, 932
marmota, IMus, 932
marmotta, Arctomys, 932
JMarsipohcmus. 238
Martcs, 365, 440

bunites, 380

foiua, 874

foina, 375
mediterranea, 380

martes, 366
,, ,, latinorum, 373

,. ., martcs, 372

martes, Martes, 366, 372
martcs, JMartcs martcs, 372
martcs, Mustcla, 372

101

matschiei, Ovis, 9S7
maura, Cervus dama, 971
mauricus, Cervus, 970
maurus, Vesper ugo, 220
maxima, Vesperugo noctula, 244
maximus, Nyctalus, 244
maximus, Pterygistes, 244
Mediocricetus, G05
mediterranea, Felis, 408
mediterranea, ^fartes foina, BSD
mediterranea, ^Mustela, 380
mediterraneus, Cervus, 909
mediteiraueus, Lepus, 51:5
mediterraneus, ^[eles meles, ?.o2
mediterraneus, I'ipistrellus pipis-

trellus, 205
medius, Arvicola, 708
medius, Lepus, 502, 508
megalotos, Plecotus, 257
megapodius, Vespcrtilio, 187
mehelyi, Rhinolophus, 159
mebelyi, llbinolophus eur3-ale, 159
melanodon, Sorex, 35
luolanogaster, Canis, 381
melauogaster, Vulpes vulpes, 331
melanopterus, Vespertilio, 201
melas, Genetta, 451
melas, Genetta genetta, 451
Meles, 341

,, arcalus, 352

„ meles, 348

,, ,, mariaiiensis, 352
„ meles, 348
meles, ^leles meles, 348
meles, ileles, 343, 348
meles, Taxus, 348
meles, Ursus, 348
Melesium, 341
melina, Lynx vulgaris, 471
IMelinse, 341
melonii, Glis, 579
melonii, Glis glis, 579
meridlanus, IMicrotus arvalis, 080
meridiei, Lepus europtous, 500
meridionalis, Lepus, 506, 510
meridionalis, ]Mus, 844
meridionalis, JIus (Apodemus), 844
meridionalis, Putorius (Ictis) nivalis,

405
meridionalis, Putorius vulgaris, 405
meridionalis, Sciurus, 912
meridionalis, Sus, 960
meridionalis, Sus scrofa, 9G0
meridionalis, Vulpes vulgaris, 331
ilesocricetus, G05

,, newtoni, GOG

Mesospalax, 1000
messorius, Mus, 844
Meteorus, 225, 238
IMicrochiroptera, 134
Micromys, 791, 840

,, minutus, 841

Micromys minutus pratensis, 840
„ „ soricinus, 841

INIicrospalax, 888
ilicrotinse, 610
Microtus, 658, 659, 712, 752
,, agrestis, 662

„ ,, agr-estis, 668

bailloni, 672
,, ,, exsol, 669

,, ,, hirtus, 673

,, ,, levernedii, 071

„ „ negleetus, 675

,, „ rozianus, OSU

,, angularii^, 70f;
,, arvalis, 081
,, ,, arvalis, 083

,, ,, duplicatus, G8G

,. levis, 087
,, ,, meridianus, 080

asturianus, 093
cabrer;e, 701
dentatus, 703
bartingi, 704
iucertus, G9U
lebrunii, 718

,, lebrunii, 719
leucurus, 722
nivalis, 713

aquitanius, 717
,, nivalis, 716
oreadensis, 694
,, ratticeps, 708

,, sandayensis, 690

,, ,, sandayensis,

697
,, ,, westrse, 698

,, sarnius, 700
,, ulpius, 723

microurus, Leucodon, 88
^licrurus, 752
midas, Dysopes, 277
milleri, Neomys, 78
mimula, Crocidura, 94, 95
raimula, Crocidura mimula, 95
miniatus, ^Eus, 844
minima, Vesperugo noctula, 245
minimus, Mus, 844
minimus, Mus minutus, 844
minimus, Mus (Apodemus) minutus,

844
minimus, Rbinolopbus, 151
minimus, Pi.binolophus bipposiderus,

151
minimus, Sorex, 55
Miniopteri, 268
MiniopteriniP, 268
Miniopteris, 268
Miniopterus, 268

„ scbreibcrsii^ 269

Minneopterus, 268
minor, Arvicola ampbibius, 740
minor, Canis lupus, 313

1012

minor, Cervns elaphus, 909

minor, Crocidura aranoa, 95, 101

minor, Lutra, 415

minor, Mus rutilus, 632

minor, IMustela, 402

minor, Neomys fodiens, 70

minor, Rhinolophns, 149

minor, Spalax, 745

minor, Ursus arctos, 28G

minor, Vespertilio lerrum-enuiuum.

149
minous, Acomys, 883
miuous, Acomys dimidiatus, 883
minuta, Croridura, 95
minuta, Phyllorbina, 149
minutellus, Vespertilio, 184
minutissimus, Vespertilio, 204
mimitus, INIieromys, 841
minutus, Mus, 844
minutus, Putorius, 402
minutus, Putorius (Ictis) vulgaris, 403
minutus, Rhinoloplius liipposiderus,

154
minutus, Sorex, 53, 55
minutus, Sorex minutus, 55
minutus, Vespertilio, 154
Minyopterus, 208
mollis, Sorex vulgaris, 43
mollissimus, JIusculus, 871
^Tolossi, 270
]\rolossida3, 276

montanus, Cervus, vulgaris, 900
montanus, Plecotus auritus, 257
]Tiontieola, Arvicola, 749
monticola, Arvicola scherman, 749
monticola, jMosospalax, 1000
monticola, Putorius nivalis, 403
monticola, Spalax, 894
monticola, Spalax monticola, 1000
Mops, 270

moreotica, Canis aureas, 315
moschata, Crocidura, 101
multiplex, Arvicola, 704
multiplex, Pitjmiys, 704
]\Iungo, 440
Mungos, 440

,, widdringtonii, 441
muralis, ^lus, 874
muralis, I\Ius musculus, 874
Muridie, 591
INIurina, 530
IMurinse, 791

murinus, Vespertilio, 192, 238
Mus, 791, 840, 848, 863

,, ffproensis, 875

,, muralis, 874

,, musculus, 805

,, ,. azoricus, 871

,, „ musculus, 809

,, spicilegus, 877

,, ,, hispanicus, 879

,, .. lusitanicus, 882

j\IuR spicilegus spicilegus, 878
musaraneus, Sorex, 101
^Muscardinidse, 549
Muscardinus, 583

,, avellanarius, 583

muscardinus, Myoxus, 583
Muscardinus pulcher, 590
Musculus, 803
musculus, ]\[us, 865, 869
musculus, Mus musculus, 869
musculus, Sorex, 70
musiguani, Arvicola, 744
musignanoi, Arvicola, 744
Musimou, 986
musimon, .T^goceros, 987
musimon, Ovis, 987
musiniani, ^licrotus, 735
Musmon, 980
musmon, Ovis, 987
Mustela, 365, 381, 384
,, africana, 412
,, erminea, 385

pestiva, 389
,, ., erminea, 387

,, ., ricinre, 397

stabilis, 390
,, galinthias, 414
bibernica, 398
,, lutreola, 415
,, nivalis, 401
,, ., boccamela, 405

„ ,, iberica, 407

,, nivalis, 402
putorius, 418

aureolus, 425
putorius, 423
Mustelidfp, 340
Mustelina, 384
Musteliua-, 364
:\rygale, 21
Mygalina, 20
IMvnomes, 659
Mvodes, 014, 023
Myogale, 21
ilyogalea, 21
]\[yopus. Oil

,, schisticolor. Oil
^lyosictis, 65
myosotis, Myotis, 193
mj'osotis, Vespertilio, 192
:\Iyospalax, 887
:\[yotis, 16()

,, bechsteinii, 179
,, eapaccinii, 187
,, dasycneme, 189
,, daubeutonii, 184
,, emarginatus, 177

myotis, 192
,, mystaeinus, 109
,, nattereri, 174
oxygnatbus, 199
myotis, Myotis, 192, 193

1013

myotis, YeRportilio, 102
Myoxidse, 549
Myoxiua, 549
Mj'oxus, 56G, 572
Myrmarctos, 285
myrmephagus, Ursus, 28G
mystacinus, Mus, 794
mystacinus, INIyotis, 169
mystacinus, Vespertilio, 169, 189

nageri, Evotorays, 641

nageri, Evotomys giareolus, 641

nageri, Hypudseus, 641

naias, Neomys fodiens, 70

Nannomys, 863

Nanuospalax, 888

Nannugo, 203

nanus, Neomys fodiens, 71

natans, Sorex, 70

nathusii, Pipistrellus, 213

nathusii, Vespertilio, 213

nathusii, Vesperugo, 213

nattereri, jMyotis, 174

uattereri, Vespertilio, 174

nebrodonsis, Arvicola, 770

nebrodensis, Pitymys, 770

nebrodensis, Pitymys savii, 770

neglectuR, Arvicola, 675

neglectus, Microtus agrestis, 672, 673,

675
neglectus, Vespertilio, 177
nehriugi, Cricetus, 605
nelaringi, Cricetus cricetus, 605
nemoralis, IMus, 854
Neocyon, 304
Neogale, 384
Neomys, 65

,, auomaluR, 81

„ fodiens, 66, 70

,, ,, bicolor, 7:?

,, ,, fodiens, 69

,, milleri, 78
nesiotes, Erinaceus, 129
nesiotes, Erinaceus europa^us, 129
nowtoni, Cricetus, 606
newtoni, Cricetus (Mesocricetus), 606
newtoni, ]\Iesocriceti;s, 606
niger, Arvicola terrestris, 746
niger, Canis lupus, 313
niger, Capreolus vulgaris, 974
niger, Castor, 947
niger, Cricetus cricetus, 602
niger, Cricetus vulgaris, 602
niger, Dama platyceros, 971
niger, Eepus timidus, 502
niger, ilus musculus, 869
niger, ^Iiis sylvaticus, 803
niger, Sciurus vulgaris, 905
niger, Ursus arctos, 285
nigra, Arctomys marmota. 932
nigra, Arvicola agrestis, 671
nigra, Canis vulpes, 33^

nigra, Sorex vulgaris, 48

nigra, Talpa ouropa^a, 4

nigra, Vespertilio pipistrellus, 204

nigrans, Vespertilio, 220

nigrescens, Sciurus vulgaris, 910

nigricans, Arvicola amphibius, 730

nigricans, Brachyotus mystacinus,

169
nigricans. Hamster, 602
nigricans, Lepus timidus, 502
nigricans, ]\Ius arvalis, 668
nigricans, Nannugo pipistrellus, 205
nigricans, Pipistrellus, 204
nigricans, Sorex fodiens, 70
nigricans, Vespertilio mystacinus, 169
nigripes, Sorex, 70
nigro-argenteus, Canis, 330
nigro-fuscus, Vespertilio mystacinus,

169
nigrocaudatus, Canis vulpus, 330
nilssoni, Eptesicus, 234
nilssoni, Vesperugo, 234
nilssonii, Eptesicus, 234
nilssonii, Vespertilio, 234
nilssonii, Vesperugo, 231
uitedula, Dyromys, 567, 568
nitodula, Dyromys nitedula, 568
nitedula, Mus, 551, 568
nitedulus, Eliomys, 568
nitela, Myoxus, 551
nivalis, Arvicola, 716, 719, 722
nivalis, ^Microtus, 713, 716
nivalis, ]\Iicrotus (Cbionorays), 716
nivalis, Microtus nivalis, 716
nivalis, Mustela, 401, 402
nivalis, ]Mustela nivalis, 402
nivalis, Putorius, 403
nivalis, Putorius (Ictis), 403
niveus, Mus musculus, 869
nivicola, Arvicola, 716
Noctula, 225
noctula, Nyctalus, 245
noctula, Ptcrygistes, 245
noctula, Vespertilio, 245
noctula, Vesperugo, 245
Noctulinia, 242
nordmauui, Sniinthus, 537
normalis, Ursus arctos, 286
norvegicus, Epimys, 858
norvegicus, Evotomys, 638
norvegicus, Evotomys giareolus, 638
norvegicus, Mus, 858
norvegicus, ]Mus (Epimys), 858
norvegicus, Ursus, 286
norwegicus, Epimys, 858
norwegicus, T^emmus, 615
nuda, Sorex vulgaris, 42
nudipes, Lutra, 356
numantius, Sciurus vulgaris, 914
Nutria, 355
Nyctalus, 242

azoreum, 254

1014

Nyctalus leisleri, 252

,, maximus, 244

,, uoctula, 245
Nyctinoma, 276
Nyctinomes, 276
Nyctinomia, 276
Nyctinomops, 276
Nyctiuomus, 276

,, teuiotis, 277

Nyctipteuus, 225
Nystactes, 166

obscurus, RhinolophuR ferrum-equi-

num, 143
occidentalis, Ennaceus europreus, 120
occideutalis, Lepus europppns, 504
occidentalis, Ovis musimon, 987
occidentalis, Talpa, 15
occideutalis, Talpa cfeca, 15
Ochetomys, 723

ochromixtus, Pipistrellus savii, 220
ochromixtus, Vespertilio, 220
Odmrclurus, 446
CEdocepbalus, 542
okenii, Vespertilio, 226
Ommatostergus, 888
orcadensis, Microtus, 694
orientalis, Lupus^ 313
ornata, Rupicapra, 994
orsinii, Vespertilio, 269
Oryctolagus, 484

cuniculus, 485

cuni cuius, 490
,, ,, huxleyi, 491

oryzivorus, Mus, 844
Otisorex, 29
otus, Vespertilio, 256
Ovis, 986

,, musimon, 987
oxygnathus, ]\Iyot4s, 199
oxygnathus, Myotis myotis, 199
oxygnathus, Vespertilio, 199
Oxygons, 304
Oxyrhin, 29

pachygnathus, Vespertilio, 252
Pachyomus, 225
Pachyura, 81

,, etrusca, 83
pachyurus, Sorex, 83
pallidus, Eliomys, 559
pallidus, Rhinolophus hipposideros,

151
Xjallidus, Sorex arancus, 42
Palmatus, 970
palmatus, Alces, 978
Paludicola, 659, 712, 723
paludosus, Mus, 738
palustris, Vespertilio, 245
Panugo, 242
Paralces, 976
pardella. Lynx, 475

pardella. Lynx (Eucorvaria), 475

pardellus. Lynx, 475

Pardina, 470

pardina, Felis, 475

parnassius, Lepus, 519

parnassius, Lepus europffius, 519

parva, Rupicapra, 995

parva, Rupicapra rupicapra, 995

parvulus, Drymomys, 869

parvulus, Mus, 844

parvus, ]\Ius sylvaticus, 803

pascuus, Pitymys ibericus, 783

Paurodus, 86

pecchioli, Mus, 810

pelandonius, Pitymys, 778

pellucens, Vespertilio, 187

pondulinus, Mus, 844

peninsulse, Genetta, 451

peninsula;, Genetta genetta, 451

pennanti, Ampliisorex, 73

peregusna, Mustela, 429

peregusna, Vormela, 429

pcrsonatus, Sorex, 35

pcrspicillatus, Vespertilio, 142

portinax, Arvicola, 740

Petauristidfe, 940

petropbilus, Hypudteus, 716

pbasma, Rhinolophus, 151

Phyllostoraata, 130

Phyllotis, 137

picta, Capra aegagrus, 992

picteti, Mus, 854

pictus, j5^igoceros, 992

Pinalia, 65

Pinemys, 752

pipistrelle, Vespertilio, 204

Pipistrellus, 202

kuhlii, 215
nathusii, 213
pipistrellus, 204
savii, 219

pipistrellus, Nannugo, 204

pipistrellus, Pipistrellus, 204

pipistrellus, Vespertilio, 204
pipistrellus, Vesperugo, 204
piscatoria, Mustela Lutra, 355
Pitymys, 752

,, atticus, 787
dacius, 700
depressus, 779
druentius, 762'
,, duodecimeostatus, 784
fatioi, 763
gerbii, 773
„ ibericus, 780
,, ., centralis, 782

,, ,, ibericus, 782

,, ,, pascuus, 783

,, ,, regulus, 784

,, lusitanicus, 776
,, marine, 777
,, multiplex, 764

INDEX

1015

Pitymys ncbrodcusis, 770
„ pelandonius, 778
,, planiccps, 172
„ proviucialis, 785
,, pyreiiaicus, 770
., ,, brunucus, 772

,, ., pyrenaicus, 771

,, savii, 768

subterraueus, 755

capuciiius, 700
subterraueus,
758
,, thomasi, 786
planiccps, Pitymys, 772
Platyceros, 970
platyccros, Ccrvus, 970
platyrhynchus, Cervus (Taranclus),

985
platyrhynchus, Rangifer, 985
Plecotus, 256

„ auritus, 256
Plerodus, 81

plumbeus, Cervus capreolus, 974
polaris, Ursus, 298
poliogaster, Crocidura, 101
polonicus, Spalax, 1000
polonicus, Ursus arctos, 286
poschiavinus, Mus, 869
j)0schiavinus, Mus musculus, 870
pratensis, Arvicola, 632
pratensis, I\Iicromys minutus, 810
pratensis, Mus, 846
pratensis, ]Mus minutus, 846
pratensis, ]Mus (Apodemus), minutus,

846
Praticola, 659, 712, 728
princeps, jNIus sylvaticus, 829
principalis, Arvicola arvalis, 083
Procerus, 979
Procervus, 979
proprius, Castor, 948
proterus, Vespertilio, 245
proviucialis, Pitymys, 785
Psammomys, 752
Pscudoconomys, 863
psilurus, Sorex, 70
Pternopterus, 167
Pteromidge, 940
Pteromyinse, 940
Pteromvs, 941
Pterygistes, 542
pulcher, Muscardiuus, 590
pulcbra, Crocidura russula, 103
pullatus, Vcsperugo kuhlii, 216
pumilio, Sorex, 55
pumik;s, ]Mus, 844
pumilus. Sores, 55
pusillus, Fcetorius, 403
pusillus, Vespertilio, 204
Putorius, 418
putorius, Fcetorius, 123
putorius, Mustcla, 418, 123

putorius, ^lustela putorius, 423
putorius, Putorius, 423
pygmteus, Sorex, 55
pygma3Us, Vespertilio, 204
pyrenseus, Ursus, 286
pyrenaica, Capra, 889, 990, 991
pyrenaica, Capra pyrenaica, 990
pyrenaica, jMygale, 26
pyrenaica, IMyogale, 26
pyrehaica, Rupicapra, 995
pyrenaica, Rupicapra rupicapra, 995
pyrenaicus, Arvicola, 771
pyrenaicus, Galemys, 21
pyrenaicus, Galemys pyrenaicus, 26
pyreuaicus, Glis glis, 582
pyrenaicus, Lepus europaeus, 506
pyreuaicus, Pitymys, 770, 771
pyreuaicus, Pitymys pyreuaicus, 771
pyrenaicus. Sores araneus, 41
pyrenaicus, Ursus, 286
pyreuaicus, Ursus arctos, 287

quadricaudatus, Sorex, 42
I quercinus, Eliomys, 551, 558
quercinus, Mus, 551
quercinus, Myoxus, 551

Rangifer, 979

,, fennicus, 981
,, platyrhynchus, 9S5
., tarandus, 980

rangifer, Cervus tarandus, 980
ratticeps, Arvicola, 708
ratticeps, Microtus, 70S
Rattus, 848

rattus, Epimys, 849, 853
rattus, Epimys rattus, 853
rattus, Mus, 853
rattus, ]\Ius (Epimys), 853
regularis, Pitymys ibericus, 784
regulus, Pitymys ibericus, 784
remifer, Sorex, 69
rhcnanus, Cervus, 9()6
Rhiuocrepis, 137
Rliinolophidic, 136
Rhiuolophina, 136
Rhinolophus, 137

„ blasii, 162

,, euryale, 155

,, ferrum-equiuum, 139

fer-
rum-equiuum, 142
I. ,' insu-

lanus, 147
,, liipposidoros, I 17

hipposi-

dcros, 149

>, !■ minimus,

151

„ minutus,

154

1016

Khinoloplius mohelyi, 15U

rliinclophus, Sorex, 35

rhodanica, Gcnetta, 452

rhodanica, Genetta gonctta, 452

ricinffi, IMustcla crmiuca, 397

ricinac, Putorius erminea, 3U7

riciiue, Putorius crmiueus, 397

riparius, Arvicola, 6'di

rivalis, Sorcx, 70

robubtus, Dyromys, 572

Kodeutia, 481

roensis, Lutra, 35G

roiuana, Talpa, 18

Romicia, 202

rossicus, Ursus arctos, 286

roumaaicus, Erinacous, 127

roumauicus, Erinaceus eui'opteus,

127 _ . ^

rozianus, Arvicola, G80
roisianus, Microtus agrestis, 680
rubcns, Mus, 836

rubidus, Evotomys hercynicus, 632
rubidus, Lemmus, 632
rufa, Arvicola agrestis, 671
rufa, Crocidura, 101
rufa, Talpa europtea, 4
rufescens, Arvicola, 632
rufescens, Vespertilio, 177, 245
rufescens, Vespertilio pipistrellus, 204
rufesceute-fuscus, Hypudaeus, 758
rufocanus, Evotomys, 648
rufocauus, Evotomys (Craseomys), 648
rufocanus, Hypudaeus, 648
rufofuscus, Brachyotus mystacinus,

169
rufofuscus, Hypudasus, 758
rufula, ^Myogale pyrenaica, 26
rufula, Myogalea, 26
rufulus, Galemys pyrenaicus, 26
rufus, Lopus timidus, 502
rufus, Sciurus, 914
rufus, ticiurus vulgaris, 905, 909, 910
rufus, Ursus arctos, 286
Kupicapra, 992

,, oruata, 994

,, parva, 995

,, pyrenaica, 995

,, rupicapra, 993

rupicapra, Autilope, 995
rupicapra, CapcUa, 993
rupicapra, Capra, 993
rupicapra, Kupicapra, 993, 995
russicus, Pteromys, 941
russicus, Sciuroptcrus, 941
russula, Crocidura, 99, 101
russula, Crocidura russula, 101
russulus, Sorex, 101
russus, Sciurus vulgaris, 909
rusticus, Sorex, 55
rutilans, Sciurus vulgaris, 910
rutilus, Evotomys, 646
rutilus, Mus, 646

Sacalius, 304

sandayensis, IMicrotus, 696, 697

sandayensis, i\Iicrotus orcadeusis, 697

sandayensis , Slicrotus sandayensis, 097

santonus, Sorex, 40

santouus, Sorex araneus, 40

sapidus, Arvicola, 732, 733

sapidus, Arvicola sapidus, 733

sarda, Felis, 408

sarda, Eelis libyca, 468

sarda, Felis ocreata, 468

sardous, Sus scrofa, 960

sardus, Eliomys, 560

sarmatica, Slustela, 429

sarmaticus, Foetorius, 429

sarmaticus, Putorius, 429

sarnius, Microtus, 700

savii, Arvicola, 768

savii, Dysopes, 277

savii, Pipistrellus, 219

savii, Pitymys, 768

savii, Vespertilio, 219

scalops, Talpa, 4

scandinavicus, Ursus arctos, 286

scherman, Arvicola, 744, 745

scherman, Arvicola scherman, 745

scherman, I\Ius, 745

schermaus, Mus, 745

schiuzii, Vespertilio, 169

schisticolor, Lemmus, 611

schisticolor, INIyodes, 611

schisticolor, Myopus, 611

schrankii, Vespertilio. 169, 177

schreiberii, jMiniopterus, 269

schreibersii, JMiniopterus, 269

schreibersii, Vespertilio, 269

Sciuridffi, 897

Sciuriua, 940

SciuriuiE, 940

Sciuropterus, 941

russicus, 941

Sciurus, 898

,, vulgaris, 898

,, alpinus, 914
'., „ keticus, 923

,, ,, fuscoater, 910

,, ., infuscatus, 916

,, ,, italicus, 912

,, ,, leucourus, 907

,, ,, Ulceus, 913

,, ,, numautius, 914

,, ,, russus, 909

,, ,, segurtc, 917

,, ,, varius, 906

,, ,, vulgaris, 905

scoticus, Cervus elaphus, 968

scoticus, Lepus medius, 529

scoticus, Lepus timidus, 529

Scrofa, 956

scrofa, Sus, 957

segurte, Sciurus vulgaris, 917

selysii, Arvicola, 768

1017

selysii, Arvicola (Microtus), 762
Selysius, 166
Semicricetus, 605
septentrionalis, Lcpus, 526
serbicus, Spalax monticola, 1000
serotiua, Nocfcula, 245
serotine, Vespertilio, 226
serotinus, Eptesicus, 226
serotinus, Vesx^ertilio, 226
serotinus, Vespcrugo, 226
setosus, Sus, 957
sibiricus, Pteromys, 941
Sicista, 536

,, loriger, 537
,, trizona, 539
Sicistinpe, 536
sicula, Crocidura, 108
siculse, Myoxus, 854
siculus, Eptesicus, 239
siculus, Putorius (Ictis) nivalis, 405
siculus, Putorius nivalis, 405
siculus, Vesperus, 238
signatus, Canis lupus, 314
silacca, Vulpes vulpos, 333
silaceus, Vulpos vulpos, 333
silvestris, Folis, 457, 402
silvestris, Felis (Gatus), 462
silvestris, Felis silvestris, 462
simplex, Arvicola arvalis, 683
Simplicidcutata, 535
skomerensis, Evotomys, 044
Sminthi, 530
Sminthinse, 536
Sminthus, 536
sodalis, Eptesicus, 231
sodalis, Vespertilio, 231
solitarius. Castor fiber, 947
Sorex, 29

„ alpinus, 60
,, ,, alpinus, 62

,, ,, hercynicus, 63

,, araneus, 31
,, ,, araneus, 35

,, ,, bergensis, 41

„ ,, castaneus, 37

„ ,, euronotus, 41

„ ,, fretalis, 45

,, ,, granarius, 52

„ ,, pyrenaicus, 44

,, ,, santonus, 40

,, ,, tetragonurus, 42

,, minutus, 53
,, ,, lucanius, 60

,, ,, minutus, 55

SoricidtB, 28

soricinus, Micromys minutus, 844
soriciuus, Mus, 844
sowerbyi, Crossopus, 73
Spalacidfu, 887
Spalax, 887, 1000
„ dolbrogese, 889
„ grsecus, 895

Spalax hungaricus, 894

speciosus, Muscardinus avellanarius,

590
speciosus, Myoxus, 590
spelrea, Myotis murina, 193
Spermophilus, 924
spetsbergensis, Cervus tarandus, 985
spicilegus, Mus, 877, 878
spicilegus, Mus spicilegus, 878
spitzbergeneusis, Alopox, 324
spitzbergenensis, Canis lagopus, 324
spitzbergensis, Rangifer, 985
spitzbergensis, Rangifer arcticus, 985
spitzbergensis, Thalassarctos, 298
spitzbergensis, Vulpes lagopus, 324
stabilis, Mustela crminca, 390
stabilis, Putorius ermineus, 390
stabilis, Putorius (Ictis) ermineus, 390
stagnatilis, Sorcx, 70
staufferi, Vespertilio, 184
stenorostris, Ursus arctos, 287
stenotus, Vespertilio, 204

stimmiugi, Arvicola (Microtus) rat-
ticeps, 708

stimmingi, Microtus ratticeps, 708

striatus, Mus musculus, 809

suaveolens, Crocidura, 83

subcEeruleus, Mus, 853

submurinus, Vespertilio, 193

subpalmatus, Putorius (Ictis) nivalis,
412

subterraneus, Arvicola, 758

subterraneus, Pitymys, 755, 759

subterraneus, Pitymys subterraneus,
758

subtilis, Sicista, 539

suecicus, Evotomys glareolus, 636

suecicus, Evotomys hercynicus, 686

Suidaj, 956

suillus, Erinaceus, 120

surmolottus, Mus, 858

Sus, 956
„ attila, 960
„ meridionalis, 960
,, scrofa, 957

suslica, Citellus, 929

suslica, Mus, 929

sylvatica, Gapra rupicapra, 993

sylvatica, Martes, 372

sylvaticus, Apodemus, 797, 803

sylvaticus, Apodemus sylvaticus, 803

sylvaticus, Lepus borealis, 526

sylvaticus, Mus, 803, 829

sylvestris, Felis, 462

sylvestris, JIus, 854

sylvestris, Mustela, 372

Sylvicola, 659

Synotus, 263

syrmiensis, Spalax monticola, 1000

tseniotis, Nyctiuomus, 277
Talpa, 8

3 u

1018

Talpa caeca, 15

„ europnea, 3

,, occidentalis, 15

,, romana, 18
Talpidis, 2
Talpinse, 2
Talpoides, 887
Tarandus, 979
taraudus, Cervus, 980
tarandus, Rangifer, 980
tartessia, Felis, 4G5
tartessia, Felis silvestris, 465
Taxus, 341
taxus, Meles, 348
tectorum, ]\Ius, 854
teuebricus, Arvieola, 735
tenebricus, Arvieola sapidus, 735
teniotis, Cephalotes, 277
teniotis, Nyctinomus, 277
terrestris, Arvieola, 738, 745, 746,

749
terrestris, Mus, 738, 745
Terrieola, 752
tetragonurus, Sorex, 42
tetragouurus, Sorex araneus, 42, 43
Tetramerodou, 659
Thalaretos, 297

,, maritimus, 298

Thalassarctos, 297
Thalassiarcbus, 297
tbomasi, Mierotus (Pitymys), 78G
tbomasi, Pitymys, 786
tboracinus, Sorex, 101
thotfci, Capreolus capreolus, 975
Tbous, 304

tigrina, Arctomys marmota, 932
timidus, Lepus, 502, 508, 513, 522,

527
timidus, Lepus timidus, 526
toscanus, Euryalus, 155
toscanus, Rbinolopbus euryale. 155
Tragus, 988
tragus, Rupieapra, 993
Tralatitus, 167

traussylvanicus, Capreolus, 975
transsylvanicus, Capreolus capreolus,

975
transsylvanicus, Lepus, 509
transsylvanicus, Lepus europseus, 509
transsylvanicus, Spalax bungaricus,

1000
transsylvanus, Eptesicus serotinus,

226
transsylvanus, Vesperus serotinus, 226
transsylvaticus, Lepus europseus, 509
transylvanus, Vesperus serotinus, 226
Trilatitus, 166
tripartitus, Mus, 639
tristriatus, Mus, 539
triticeus, Mus, 844 i
trizona, Sicista, 539
trizonus, Mus, 539

troglopbilus, Rbinolopbus bipposi-

deros, 150
turcicus, Spalax monticola, 1000
typicus, Lepus mcditerraneus, 513
typicus, Lepus timidus, 529
typicus, ileles meles, 348
typicus, Mus sylvaticus, 829
typicus, Putorius nivalis, 403
typicus, Rbinolopbus ferrum-cqui-

num, 143
typicus, Rbinolopbus bipposiderus,

150
typicus, Sciurus vulgaris, 905
typus, Isotus nattereri, 174
typus, Myotis murinus, 193
tyi)us, Pipistrellus, 204
typus, Plecotus auritus, 257
typus, Rbinolopbus bipposideros, 150

ulpius, Mierotus, 723

ulpius, Mierotus (Cbionomys), 723

ungula, Vespertilio, 142

Ungulata, 955

unieolor, Sorex, 101

unicolor, Vesperugo natbusii, 213

unifer, Rbinolopbus, 142

unibastatus, Rbinolopbus, 142

Ursaretos, 285

Ursidffi, 284

ursinii, Vespertilio, 269

Ursula, Vespertilio, 216

Ursus, 285

,, arctos, 285
ursus, Ursus, 285

vagans, Erinaceus algirus, 133
vagus, Smintbus, 539
variabilis, Arvieola arvalis, 683
variabilis, Lepus, 526, 528, 529, 531
variegata, Talpa europsea, 4
variegatus, Canis vulpus, 330
variegatus, Castor fiber, 947
varius, Capreolus vulgaris, 974
varius. Castor, 947
varius, Cervus elapbus, 965
varius, Cricetus vulgaris, 602
varius, Dama platyceros, 971
varius, Mus museulus, 869
varius, Mus sylvaticus, 803
varius, Rattus domesticus, 853
varius, Sciurus vulgaris, 906
varronis, Lepus, 528
varronis, Lepus medius, 528
varronis, Lepus timidus, 528
vasconiaj, Evotomys, 639
vascouiai, Evotomys glareolus, 639
velatus, Plecotus, 259
vermicula, Lepus, 490
vernicularis, Lepus, 490
Vesperugo, 202, 224, 238, 242
Vesperus, 224. 238
Vespertilio, 166, 225, 238

1019

Vespertilio murinus, 238
Vespertiliones, 165
Vespertiliouidae, 165
Vespertilioninse, 165
victoria?, Capra pyrenaica, 991
virgata, Pelis, 472
virgulosus, Mus, 539
Vison, 415

vison, Mustela putorius, 423
vispistrellus, Vespertilio, 215
vistulinus, Castor, 948
Viverridas, 440
volans, Pteromys, 941
volans, Sciurus, 941
Vormela, 428

„ peregusna, 429
vulgaris, Arvicola, 683
vulgaris, Capreolus, 974
vulgaris, Cervus, 965
vulgaris, Cricetus, 602
vulgaris, Dama, 971
vulgaris, Foetorius, 403
vulgaris, Genetta, 451
vulgaris, Genetta genetta, 452
vulgaris, Glis, 577
vulgaris, Gulo, 434
vulgaris, Lutra, 355
vulgaris, Lynx, 471
vulgaris, Martes, 372
vulgaris, Mustela, 402
vulgaris, Plecotus, 256
vulgaris, Pteromys, 941
vulgaris, Putorius, 423
viilgaris, Putorius nivalis, 403
vulgaris, Putorius (Ictis) nivalis, 403

vulgaris, Sciurus, 898, 905, 907, 909,

910, 912, 914
vulgaris, Sciurus vulgaris, 905
vulgaris, Sorex, 35, 42
vulgaris, Talpa, 4
vulgaris, Taxus, 348
vulgaris, Vulpes, 330
Vulpes, 325

,, ichnusse, 330

,, vulpes, 326

,, ,, crucigera, 331

,, ,, silacea, 333

,, ,, vulpes, 330

vulpes, Canis, 330, 331
vulpes, Vulpes, 326, 330, 331
vulpes, Vulpes vulpes, 330
Vulpicanis, 304
vulpinus, Felis, 471
vulpus, Canis, 330

westrte, Microtus sandayensis, 698
widdringtoni, Herpestes ichneumon,

441
vyiddringtonii, Herpestes, 441
widdringtonii, Mungos, 441
wiedii, Vespertilio, 226
wingei, Dyromys nitedula, 570
wingei, Myoxus, 570
wintoni, Apodemus flavicoUis, 831
wintoni, Mus sylvaticus, 831

Zapodidae, 535
zemni, Glis, 1000
Zibellina, 365

LONDON :

PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,

DUKE STREET, STAMFORD STREET. S.E. , AND GREAT WINDMILL STREET, W.

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